Institute of Archaeology

The Bronze Age of Eurasia (around 3000–1000 BC) was a period of major cultural changes. However, there is debate about whether these changes resulted from the circulation of ideas or from human migrations, potentially also facilitating the spread of languages and certain phenotypic traits. We investigated this by using new, improved methods to sequence low-coverage genomes from 101 ancient humans from across Eurasia. We show that the Bronze Age was a highly dynamic period involving large-scale population migrations and replacements, responsible for shaping major parts of present-day demographic structure in both Europe and Asia. Our findings are consistent with the hypothesized spread of Indo-European languages during the Early Bronze Age. We also demonstrate that light skin pigmentation in Europeans was already present at high frequency in the Bronze Age, but not lactose tolerance, indicating a more recent onset of positive selection on lactose tolerance than previously thought. An analysis of 101 ancient human genomes from the Bronze Age (3000–1000 bc) reveals large-scale population migrations in Eurasia consistent with the spread of Indo-European languages; individuals frequently had light skin pigmentation but were not lactose tolerant. Was the Bronze Age of a period of major cultural changes because of circulation of ideas or because of large-scale migrations? The authors sequence and analyse low-coverage genomes from 101 ancient humans from across Eurasia to reveal large-scale population migrations and replacements during this time. Analyses indicate that light skin pigmentation was already frequent among Europeans in the Bronze Age but not lactose tolerance, indicating a more recent onset of positive selection on the latter trait than previously believed. The reported findings are also consistent with the spread of Indo-European languages during the Early Bronze Age reported on page 207 of this issue.

Extended provisioning of offspring and long postmenopausal life spans are characteristic of all modern humans but no other primates. These traits may have evolved in tandem. Analysis of relationships between women's time allocation and children's nutritional welfare among the Hadza of northern Tanzania yields results consistent with this proposition. Implications for current thought about the evolution of hominid food sharing, life history, and social organization are discussed.

Abstract Domestication of horses fundamentally transformed long-range mobility and warfare 1 . However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling 2–4 at Botai, Central Asia around 3500 bc 3 . Other longstanding candidate regions for horse domestication, such as Iberia 5 and Anatolia 6 , have also recently been challenged. Thus, the genetic, geographic and temporal origins of modern domestic horses have remained unknown. Here we pinpoint the Western Eurasian steppes, especially the lower Volga-Don region, as the homeland of modern domestic horses. Furthermore, we map the population changes accompanying domestication from 273 ancient horse genomes. This reveals that modern domestic horses ultimately replaced almost all other local populations as they expanded rapidly across Eurasia from about 2000 bc , synchronously with equestrian material culture, including Sintashta spoke-wheeled chariots. We find that equestrianism involved strong selection for critical locomotor and behavioural adaptations at the GSDMC and ZFPM1 genes. Our results reject the commonly held association 7 between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 bc 8,9 driving the spread of Indo-European languages 10 . This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early second millennium bc Sintashta culture 11,12 .

BACKGROUND: Canagliflozin reduces the risk of kidney failure in patients with type 2 diabetes mellitus and chronic kidney disease, but effects on specific cardiovascular outcomes are uncertain, as are effects in people without previous cardiovascular disease (primary prevention). METHODS: In CREDENCE (Canagliflozin and Renal Events in Diabetes With Established Nephropathy Clinical Evaluation), 4401 participants with type 2 diabetes mellitus and chronic kidney disease were randomly assigned to canagliflozin or placebo on a background of optimized standard of care. RESULTS: Primary prevention participants (n=2181, 49.6%) were younger (61 versus 65 years), were more often female (37% versus 31%), and had shorter duration of diabetes mellitus (15 years versus 16 years) compared with secondary prevention participants (n=2220, 50.4%). Canagliflozin reduced the risk of major cardiovascular events overall (hazard ratio [HR], 0.80 [95% CI, 0.67-0.95]; P=0.01), with consistent reductions in both the primary (HR, 0.68 [95% CI, 0.49-0.94]) and secondary (HR, 0.85 [95% CI, 0.69-1.06]) prevention groups (P for interaction=0.25). Effects were also similar for the components of the composite including cardiovascular death (HR, 0.78 [95% CI, 0.61-1.00]), nonfatal myocardial infarction (HR, 0.81 [95% CI, 0.59-1.10]), and nonfatal stroke (HR, 0.80 [95% CI, 0.56-1.15]). The risk of the primary composite renal outcome and the composite of cardiovascular death or hospitalization for heart failure were also consistently reduced in both the primary and secondary prevention groups (P for interaction >0.5 for each outcome). CONCLUSIONS: Canagliflozin significantly reduced major cardiovascular events and kidney failure in patients with type 2 diabetes mellitus and chronic kidney disease, including in participants who did not have previous cardiovascular disease. CLINICAL TRIAL REGISTRATION: URL: https://www.clinicaltrials.gov. Unique identifier: NCT02065791.

Zooarcheological evidence suggests that pigs were domesticated in Southwest Asia ~8,500 BC. They then spread across the Middle and Near East and westward into Europe alongside early agriculturalists. European pigs were either domesticated independently or more likely appeared so as a result of admixture between introduced pigs and European wild boar. As a result, European wild boar mtDNA lineages replaced Near Eastern/Anatolian mtDNA signatures in Europe and subsequently replaced indigenous domestic pig lineages in Anatolia. The specific details of these processes, however, remain unknown. To address questions related to early pig domestication, dispersal, and turnover in the Near East, we analyzed ancient mitochondrial DNA and dental geometric morphometric variation in 393 ancient pig specimens representing 48 archeological sites (from the Pre-Pottery Neolithic to the Medieval period) from Armenia, Cyprus, Georgia, Iran, Syria, and Turkey. Our results reveal the first genetic signatures of early domestic pigs in the Near Eastern Neolithic core zone. We also demonstrate that these early pigs differed genetically from those in western Anatolia that were introduced to Europe during the Neolithic expansion. In addition, we present a significantly more refined chronology for the introduction of European domestic pigs into Asia Minor that took place during the Bronze Age, at least 900 years earlier than previously detected. By the 5th century AD, European signatures completely replaced the endemic lineages possibly coinciding with the widespread demographic and societal changes that occurred during the Anatolian Bronze and Iron Ages.

Abstract The origins of agriculture involved pathways of domestication in which human behaviours and plant genetic adaptations were entangled. These changes resulted in consequences that were unintended at the start of the process. This paper highlights some of the key innovations in human behaviours, such as soil preparation, harvesting and threshing, and how these were coupled with genetic ‘innovations’ within plant populations. We identify a number of ‘traps’ for early cultivators, including the needs for extra labour expenditure on crop-processing and soil fertility maintenance, but also linked gains in terms of potential crop yields. Compilations of quantitative data across a few different crops for the traits of non-shattering and seed size are discussed in terms of the apparently slow process of domestication, and parallels and differences between different regional pathways are identified. We highlight the need to bridge the gap between a Neolithic archaeobotanical focus on domestication and a focus of later periods on crop-processing activities and labour organization. In addition, archaeobotanical data provide a basis for rethinking previous assumptions about how plant genetic data should be related to the origins of agriculture and we contrast two alternative hypotheses: gradual evolution with low selection pressure versus metastable equilibrium that prolonged the persistence of ‘semi-domesticated’ populations. Our revised understanding of the innovations involved in plant domestication highlight the need for new approaches to collecting, modelling and integrating genetic data and archaeobotanical evidence.

Abstract There have been two trends in paleocological research. The first concentrated on the identification of specimens to species and genus level. This methodology enabled precise paleocological reconstruction based on modern analogues. On the down side, this type of analysis left out many animal bone fragments that were not identifiable. The second trend incorporated higher-level taxonomy. Since many tribes and families are similar in their habitat requirements, the analysis of the unidenti-fied fragments has the potential to alter previous conclusions (based on specimens identified to species alone), providing a more comprehensive picture of the environment. This paper addresses the question of whether identification to higher-level taxonomy may change previous paleoecologi-cal inferences from prior analyses of Stratum I26 from the site of ‘Ubeidiya. Two databases are compared. The first is the published database (Tchernov 1986a and b), which includes only specimens identified to species and genus; the second in-cludes all fragments identified at higher taxonomic levels. Different paleoecological models are applied to both databases. Results indicate that NISP increased from ca. 60 to 2300 with inclusion of the fragments. Species richness, based on mor-photyping, increased ~3 fold. The paleoecological reconstruction shifted from a habitat dominated by woodland and wood-land-bushland to one which included grassland environments. These results suggest that using higher-level taxonomy is of great importance in paleoecological reconstruction (as op-posed to Bar Oz and Dayan, 2002). The discrepancy arises from the fact that a low diversity site (such as those from Le-vanite Epipalaeolithic) may be heavily dominated by a single species (e.g. gazelle) while the high richness and diversity in ‘Ubeidiya make palaeolenvironmental reconstructions much more susceptible to a change in analytical methodology. One cannot

Farming was established in Central Europe by the Linearbandkeramik culture (LBK), a well-investigated archaeological horizon, which emerged in the Carpathian Basin, in today's Hungary. However, the genetic background of the LBK genesis is yet unclear. Here we present 9 Y chromosomal and 84 mitochondrial DNA profiles from Mesolithic, Neolithic Starčevo and LBK sites (seventh/sixth millennia BC) from the Carpathian Basin and southeastern Europe. We detect genetic continuity of both maternal and paternal elements during the initial spread of agriculture, and confirm the substantial genetic impact of early southeastern European and Carpathian Basin farming cultures on Central European populations of the sixth-fourth millennia BC. Comprehensive Y chromosomal and mitochondrial DNA population genetic analyses demonstrate a clear affinity of the early farmers to the modern Near East and Caucasus, tracing the expansion from that region through southeastern Europe and the Carpathian Basin into Central Europe. However, our results also reveal contrasting patterns for male and female genetic diversity in the European Neolithic, suggesting a system of patrilineal descent and patrilocal residential rules among the early farmers.

An underappreciated feature of complex societies is the production and consumption of alcoholic beverages, in particular wines and beers. A variety of data are reviewed which suggest significant expansion of alcoholic beverage production and consumption in many areas of Western Asia during the 4th and 3d millennia B.C. Production of beverages formed part of the processes by which emerging elites expanded control over craft production, established symbols, created manipulable surpluses, and renegotiated gender roles. Consumption of beverages was an important element of nutrition, ritual, and political economy in the early societies of Mesopotamia, Egypt, Syria, and the Levant. Review of evidence from the Aegean indicates similar processes at work but with emphasis on competitive feasting and hospitality. These different uses of alcoholic beverages represent significant regularities in the emergence of social complexity and the rise of the state.

In the second article of this series, brief mention was made of a cave at Kastritsa, near Lake Ioannina, whose talus had yielded an industry reminiscent of the last phase at the shelter of Asprochaliko. The evidence for an increase in the size of Lake Ioannina during the Last Glaciation was also outlined. The purpose of this paper is to consider the concomitant environmental changes in relation to the evidence for human occupation. Lake Ioannina lies at 469 metres above sea-level. To the north-east stands the Mitsikeli ridge, with a maximum elevation of 1,810 metres and separated from the Pindus range by the gorge of the upper Arakhthos (fig. 1). To the south-west lies the Tomarokhoria plateau, which in places rises to over 1,900 metres. The basin, a typical polje , is cradled by limestones which range in age from the Upper Triassic to the Upper Eocene. The Flysch of the Pindus borders it on the south-east. The lake lies along the axis of the Perama syncline, which runs north-west-south-east parallel to the Mitsikeli anticline and the Stavraki anticline; faulting is common. The basin would seem to owe its existence primarily to structural factors, although solution doubtless contributed to its present form.

BACKGROUND: Cattle domestication started in the 9(th) millennium BC in Southwest Asia. Domesticated cattle were then introduced into Europe during the Neolithic transition. However, the scarcity of palaeogenetic data from the first European domesticated cattle still inhibits the accurate reconstruction of their early demography. In this study, mitochondrial DNA from 193 ancient and 597 modern domesticated cattle (Bos taurus) from sites across Europe, Western Anatolia and Iran were analysed to provide insight into the Neolithic dispersal process and the role of the local European aurochs population during cattle domestication. RESULTS: Using descriptive summary statistics and serial coalescent simulations paired with approximate Bayesian computation we find: (i) decreasing genetic diversity in a southeast to northwest direction, (ii) strong correlation of genetic and geographical distances, iii) an estimated effective size of the Near Eastern female founder population of 81, iv) that the expansion of cattle from the Near East and Anatolia into Europe does not appear to constitute a significant bottleneck, and that v) there is evidence for gene-flow between the Near Eastern/Anatolian and European cattle populations in the early phases of the European Neolithic, but that it is restricted after 5,000 BCE. CONCLUSIONS: The most plausible scenario to explain these results is a single and regionally restricted domestication process of cattle in the Near East with subsequent migration into Europe during the Neolithic transition without significant maternal interbreeding with the endogenous wild stock. Evidence for gene-flow between cattle populations from Southwestern Asia and Europe during the earlier phases of the European Neolithic points towards intercontinental trade connections between Neolithic farmers.

My research has shown that the type of regional archaeological data analysis required by landscape archaeological approaches is an area where both theory and method are still in their infancy. High-level theories about the occurrence, scope, and effects of processes such as centralization, urbanization, and Hellenization/Romanization cannot yet be supported by middle range theory, which itself cannot be developed until the basic business of generating information of sufficient quality about the archaeological record has been tackled. Currently, archaeological data can be made to fit almost any interpretation generated, ultimately, on the basis of the ancient written sources. If we are to escape from this selfreinforcing cycle, research should perhaps no longer be focused on the classical themes generated by culture-historical approaches, but should seek its own proper field of operation. In the area of methods and methodology, I have demonstrated the pervasive influence of systematic research and visibility biases on the patterns that are present in the archaeological data generated over the past 50 years or so. There are mechanisms at work, both in the traditional archaeological interpretation of limited numbers of excavated sites and historical sources, and in the landscape archaeological approach, that cause the systematic undervaluation of unobtrusive remains. The significance of systematic biases in both the coarse site-based data sets resulting from desktop and ‘topographic’ studies and the more detailed site-based or ‘continuous’ data resulting from intensive field surveys has become much clearer as a result of the studies reported here. This should have practical consequences for the ways in which we study the existing archaeological record, plan future landscape archaeological research, and conduct field surveys. Site databases, the traditional starting point for regional archaeological studies, can no longer be taken at face value; rather, they require careful source criticism before being used to support specific arguments and hypotheses about settlement and land use dynamics. My studies have also shown that future data collection, whether through field survey, excavation or other methods, has to take place in a much more methodical manner if we are to produce data that are sufficiently standardized to be successfully exchanged, compared, and interpreted by others – guidelines for which should become embodied in an international standard defining ‘best practice in landscape archaeology’.

Through integrated analysis of archaeobotanical and artefactual distributions across a settlement, the authors discover ‘neighbourhoods’ using different cultivation areas in the surrounding landscape. Differences between groups also emerge over the life of the settlement in the use of special plants, such as opium poppy and feathergrass. Spatial configurations of cultivation and plant use map out the shifting social geographies of a Neolithic community.

Diagnosis of the congenital form of syphilis is an important part of the palaeopathology of this disease. In theory, there are clear clinical signs to be found in the long bones and teeth, but it has rarely been possible to recognise the latter with a confidence in archaeological material, partly because the original descriptions of the dental deformities are sometimes contradictory and partly because it is nowadays difficult to find reference specimens in museums. This article describes two such specimens which have recently been rediscovered, and discusses the form of the dental defects which they show (Hutchinson's incisors, Moon's molars, and mulberry molars) in relation to the developmental sequence of the teeth.

Around 8200 calBP, large parts of the now submerged North Sea continental shelf (‘Doggerland’) were catastrophically flooded by the Storegga Slide tsunami, one of the largest tsunamis known for the Holocene, which was generated on the Norwegian coastal margin by a submarine landslide. In the present paper, we derive a precise calendric date for the Storegga Slide tsunami, use this date for reconstruction of contemporary coastlines in the North Sea in relation to rapidly rising sea-levels, and discuss the potential effects of the tsunami on the contemporaneous Mesolithic population. One main result of this study is an unexpectedly high tsunami impact assigned to the western regions of Jutland.

Abstract Farming was first introduced to southeastern Europe in the mid-7 th millennium BCE – brought by migrants from Anatolia who settled in the region before spreading throughout Europe. To clarify the dynamics of the interaction between the first farmers and indigenous hunter-gatherers where they first met, we analyze genome-wide ancient DNA data from 223 individuals who lived in southeastern Europe and surrounding regions between 12,000 and 500 BCE. We document previously uncharacterized genetic structure, showing a West-East cline of ancestry in hunter-gatherers, and show that some Aegean farmers had ancestry from a different lineage than the northwestern Anatolian lineage that formed the overwhelming ancestry of other European farmers. We show that the first farmers of northern and western Europe passed through southeastern Europe with limited admixture with local hunter-gatherers, but that some groups mixed extensively, with relatively sex-balanced admixture compared to the male-biased hunter-gatherer admixture that prevailed later in the North and West. Southeastern Europe continued to be a nexus between East and West after farming arrived, with intermittent genetic contact from the Steppe up to 2,000 years before the migration that replaced much of northern Europe’s population.

Although sled dogs are one of the most specialized groups of dogs, their origin and evolution has received much less attention than many other dog groups. We applied a genomic approach to investigate their spatiotemporal emergence by sequencing the genomes of 10 modern Greenland sled dogs, an ~9500-year-old Siberian dog associated with archaeological evidence for sled technology, and an ~33,000-year-old Siberian wolf. We found noteworthy genetic similarity between the ancient dog and modern sled dogs. We detected gene flow from Pleistocene Siberian wolves, but not modern American wolves, to present-day sled dogs. The results indicate that the major ancestry of modern sled dogs traces back to Siberia, where sled dog-specific haplotypes of genes that potentially relate to Arctic adaptation were established by 9500 years ago.

Abstract Horses revolutionized human history with fast mobility 1 . However, the timeline between their domestication and their widespread integration as a means of transport remains contentious 2–4 . Here we assemble a collection of 475 ancient horse genomes to assess the period when these animals were first reshaped by human agency in Eurasia. We find that reproductive control of the modern domestic lineage emerged around 2200 bce , through close-kin mating and shortened generation times. Reproductive control emerged following a severe domestication bottleneck starting no earlier than approximately 2700 bce , and coincided with a sudden expansion across Eurasia that ultimately resulted in the replacement of nearly every local horse lineage. This expansion marked the rise of widespread horse-based mobility in human history, which refutes the commonly held narrative of large horse herds accompanying the massive migration of steppe peoples across Europe around 3000 bce and earlier 3,5 . Finally, we detect significantly shortened generation times at Botai around 3500 bce , a settlement from central Asia associated with corrals and a subsistence economy centred on horses 6,7 . This supports local horse husbandry before the rise of modern domestic bloodlines.

Perhaps nowhere in European prehistory does the idea of clearly-defined cultural boundaries remain more current than in the initial Neolithic, where the southeast–northwest trend of the spread of farming crosses what is perceived as a sharp divide between the Balkans and central Europe. This corresponds to a distinction between the Vinča culture package, named for a classic site in Serbia, with its characteristic pottery assemblage and absence of longhouses, and the Linearbandkeramik (LBK), with equally diagnostic but different pottery, and its apparently culturally-diagnostic longhouses, extending in a more northerly belt through central Europe westward to the Dutch coast. In this paper we question the concept of such a clear division through a presentation of new data from the site of Szederkény-Kukorica-dűlő. A large settlement in southeast Transdanubia, Hungary, excavated in advance of road construction, Szederkény is notable for its combination of pottery styles, variously including Vinča A, Ražište and LBK, and longhouses of a kind otherwise familiar from the LBK world. Formal modelling of its date establishes that the site probably began in the later 54th century cal BC, lasting until the first decades of the 52nd century cal BC. Occupation, featuring longhouses, pits and graves, probably began at the same time in the eastern and western parts of the settlement, starting a decade or two later in the central part; the western part was probably the last to be abandoned. Vinča pottery is predominantly associated with the eastern and central parts of the site, and Ražište pottery with the west. Formal modelling of the early history of longhouses in the LBK world suggests their emergence in the Formative LBK of Transdanubia c. 5500 cal BC followed by rapid dispersal in the middle of the 54th century cal BC, associated with the ‘earliest’ (älteste) LBK. The adoption of longhouses at Szederkény thus appears to come a few generations after the start of this ‘diaspora’. Rather than explaining the mixture of things, practices and perhaps people at Szederkény with reference to problematic notions such as hybridity, we propose instead a more fluid and varied vocabulary, encompassing combination and amalgamation, relationships and performance in the flow of social life, and networks; this makes greater allowance for diversity and interleaving in a context of rapid change.

Abstract The harbor town of Berenike functioned in the long-distance trade between the Mediterranean and the Indian Ocean Basin from the third century BC to the early sixth century AD. This contribution aims to view the Berenike material within its wider historical context through a preliminary analysis of the combined archaeological and textual evidence. By comparing the results of the work of different specialists, the potential of a multi-disciplinary approach becomes apparent. A number of caveats are given, which illustrate the interpretative problems arising from comparing the results of different disciplines. Lastly, this attempt at integration shows that the discrepancies between the different sources offer important insights. Le port maritime de Berenike fonctionnait dans le commerce de longue distance entre la Méditerranée et le bassin de la Mer des Indes depuis le troisième siècle avant JC jusqu'au début du sixième siècle après JC. Cette contribution examine le matériel de Berenike dans un contexte plus large par une analyse préliminaire croisant des données archéologiques et textuelles. En comparant les résultats proposés par plusieurs spécialistes, le potentiel du travail multidisciplinaire devient évident. Un nombre de caveats est présenté pour illustrer des problèmes d'interprétation, provenant des comparaisons de résultats de plusieurs disciplines. Enfin, cet essai d'intégration montre que les divergences entre plusieurs sources offre des aperçus importants.