Life histories in Fiji as reconstructed from first millennium CE Sigatoka Sand Dune burials using isotopes.

This paper aims to re-examine the dietary practices of individuals buried at Sigatoka Sand Dunes site (Fiji) in Burial Ground 1 excavated by Simon Best in 1987 and 1988 using two approaches and a reassessment of their archaeological, bioarchaeological and chronological frame. First, stable carbon and nitrogen isotope analysis was applied to document dietary changes between childhood and adulthood using an intra-individual approach on paired bone-tooth. Second, the potential adaptation of the individuals to their environment was evaluated through regional and temporal comparisons using inter-individual bone analysis. Ten AMS radiocarbon dates were measured directly on human bone collagen samples, placing the series in a range of approximately 600 years covering the middle of the first millennium CE (1,888 to 1,272 cal BP). δ13C and δ15N ratios were measured on bone and tooth collagen samples from 38 adult individuals. The results show that δ15N values from tooth are higher than those s from bone while bone and tooth δ13C values are similar, except for females. Fifteen individuals were included in an intra-individual analysis based on paired bone and tooth samples, which revealed six dietary patterns distinguished by a differential dietary intake of marine resources and resources at different trophic levels. These highlight sex-specific differences not related to mortuary practices but to daily life activities, supporting the hypothesis of a sexual division of labour. Compared to other Southwest Pacific series, Sigatoka diets show a specific trend towards marine food consumption that supports the hypothesis of a relative food self-sufficiency requiring no interactions with other groups.

"Feeling at home in Vanuatu": Integration of newcomers from the East during the last millennium.

Several localities across the Vanuatu archipelago (Melanesia), so-called 'Polynesian Outliers', are inhabited by communities that display Polynesian linguistic and cultural features although being located outside the Polynesian Triangle. Several introductions of Polynesian genetic components to Central and Southern Vanuatu during the last millenium have resulted in the cultural distinctiveness observed among the Polynesian Outliers in Vanuatu. However, social, political or economic process surrounding the exchange of genes between Polynesian and local individuals remain unidentified. Recent bioanthropological studies suggest the existence of female mobilities from neighboring regions to Vanuatu but also to the Polynesian Outliers of Taumako (Solomon Islands) within patrilocal societies. We aim to examine the hypothesis that Polynesian biological affinities observed in ancient individuals from Vanuatu are gendered or sex-specific, and that some of the Polynesian migrations during the last millennium may have involved practices of exogamy. By reconstructing phenotypes and biological identities from 13 archaeologically-recovered human skulls (400-300 years ago) from "Polynesian-related" regions of Vanuatu, we provide new insights to better contextualize the settlement patterns of Polynesian individuals. Eastern-Pacific associated phenotype are observable in 4 women from the Eretok burial complex (Efate region) and the Polynesian Outlier of Futuna, who were buried in close proximity to individuals with Western-Pacific associated phenotype. We suggest that close integration of individuals from the East into the local Vanuatu society, as well as the practice of exogamy, might have been key processes contributing to the preservation of Polynesian cultural features in Vanuatu over the past millennium. Our finding are cross-referenced with oral records from these two areas, as well as the known genetic makeup of the Vanuatu Polynesian Outliers.

The genomic landscape of contemporary western Remote Oceanians.

The Vanuatu archipelago served as a gateway to Remote Oceania during one of the most extensive human migrations to uninhabited lands ∼3,000 years ago. Ancient DNA studies suggest an initial settlement by East Asian-related peoples that was quickly followed by the arrival of Papuan-related populations, leading to a major population turnover. Yet there is uncertainty over the population processes and the sociocultural factors that have shaped the genomic diversity of ni-Vanuatu, who present nowadays among the world's highest linguistic and cultural diversity. Here, we report new genome-wide data for 1,433 contemporary ni-Vanuatu from 29 different islands, including 287 couples. We find that ni-Vanuatu derive their East Asian- and Papuan-related ancestry from the same source populations and descend from relatively synchronous, sex-biased admixture events that occurred ∼1,700-2,300 years ago, indicating a peopling history common to the whole archipelago. However, East Asian-related ancestry proportions differ markedly across islands, suggesting that the Papuan-related population turnover was geographically uneven. Furthermore, we detect Polynesian ancestry arriving ∼600-1,000 years ago to Central and South Vanuatu in both Polynesian-speaking and non-Polynesian-speaking populations. Last, we provide evidence for a tendency of spouses to carry similar genetic ancestry, when accounting for relatedness avoidance. The signal is not driven by strong genetic effects of specific loci or trait-associated variants, suggesting that it results instead from social assortative mating. Altogether, our findings provide an insight into both the genetic history of ni-Vanuatu populations and how sociocultural processes have shaped the diversity of their genomes.

Genomic insights into population history and biological adaptation in Oceania.

The Pacific region is of major importance for addressing questions regarding human dispersals, interactions with archaic hominins and natural selection processes1. However, the demographic and adaptive history of Oceanian populations remains largely uncharacterized. Here we report high-coverage genomes of 317 individuals from 20 populations from the Pacific region. We find that the ancestors of Papuan-related ('Near Oceanian') groups underwent a strong bottleneck before the settlement of the region, and separated around 20,000-40,000 years ago. We infer that the East Asian ancestors of Pacific populations may have diverged from Taiwanese Indigenous peoples before the Neolithic expansion, which is thought to have started from Taiwan around 5,000 years ago2-4. Additionally, this dispersal was not followed by an immediate, single admixture event with Near Oceanian populations, but involved recurrent episodes of genetic interactions. Our analyses reveal marked differences in the proportion and nature of Denisovan heritage among Pacific groups, suggesting that independent interbreeding with highly structured archaic populations occurred. Furthermore, whereas introgression of Neanderthal genetic information facilitated the adaptation of modern humans related to multiple phenotypes (for example, metabolism, pigmentation and neuronal development), Denisovan introgression was primarily beneficial for immune-related functions. Finally, we report evidence of selective sweeps and polygenic adaptation associated with pathogen exposure and lipid metabolism in the Pacific region, increasing our understanding of the mechanisms of biological adaptation to island environments.

Bone loss markers in the earliest Pacific Islanders.

Kingdom of Tonga in Polynesia is one of the most obese nations where metabolic conditions, sedentary lifestyles, and poor quality diet are widespread. These factors can lead to poor musculoskeletal health. However, whether metabolic abnormalities such as osteoporosis occurred in archaeological populations of Tonga is unknown. We employed a microscopic investigation of femur samples to establish whether bone loss afflicted humans in this Pacific region approximately 3000 years ago. Histology, laser confocal microscopy, and synchrotron Fourier-transform infrared microspectroscopy were used to measure bone vascular canal densities, bone porosity, and carbonate and phosphate content of bone composition in eight samples extracted from adult Talasiu males and females dated to 2650 BP. Compared to males, samples from females had fewer vascular canals, lower carbonate and phosphate content, and higher bone porosity. Although both sexes showed evidence of trabecularised cortical bone, it was more widespread in females (35.5%) than males (15.8%). Our data suggest experiences of advanced bone resorption, possibly as a result of osteoporosis. This provides first evidence for microscopic bone loss in a sample of archaeological humans from a Pacific population widely afflicted by metabolic conditions today.

Three Phases of Ancient Migration Shaped the Ancestry of Human Populations in Vanuatu.

The archipelago of Vanuatu has been at the crossroads of human population movements in the Pacific for the past three millennia. To help address several open questions regarding the history of these movements, we generated genome-wide data for 11 ancient individuals from the island of Efate dating from its earliest settlement to the recent past, including five associated with the Chief Roi Mata's Domain World Heritage Area, and analyzed them in conjunction with 34 published ancient individuals from Vanuatu and elsewhere in Oceania, as well as present-day populations. Our results outline three distinct periods of population transformations. First, the four earliest individuals, from the Lapita-period site of Teouma, are concordant with eight previously described Lapita-associated individuals from Vanuatu and Tonga in having almost all of their ancestry from a "First Remote Oceanian" source related to East and Southeast Asians. Second, both the Papuan ancestry predominating in Vanuatu for the past 2,500 years and the smaller component of Papuan ancestry found in Polynesians can be modeled as deriving from a single source most likely originating in New Britain, suggesting that the movement of people carrying this ancestry to Remote Oceania closely followed that of the First Remote Oceanians in time and space. Third, the Chief Roi Mata's Domain individuals descend from a mixture of Vanuatu- and Polynesian-derived ancestry and are related to Polynesian-influenced communities today in central, but not southern, Vanuatu, demonstrating Polynesian genetic input in multiple groups with independent histories.

Language continuity despite population replacement in Remote Oceania.

Recent genomic analyses show that the earliest peoples reaching Remote Oceania-associated with Austronesian-speaking Lapita culture-were almost completely East Asian, without detectable Papuan ancestry. However, Papuan-related genetic ancestry is found across present-day Pacific populations, indicating that peoples from Near Oceania have played a significant, but largely unknown, ancestral role. Here, new genome-wide data from 19 ancient South Pacific individuals provide direct evidence of a so-far undescribed Papuan expansion into Remote Oceania starting ~2,500 yr BP, far earlier than previously estimated and supporting a model from historical linguistics. New genome-wide data from 27 contemporary ni-Vanuatu demonstrate a subsequent and almost complete replacement of Lapita-Austronesian by Near Oceanian ancestry. Despite this massive demographic change, incoming Papuan languages did not replace Austronesian languages. Population replacement with language continuity is extremely rare-if not unprecedented-in human history. Our analyses show that rather than one large-scale event, the process was incremental and complex, with repeated migrations and sex-biased admixture with peoples from the Bismarck Archipelago.

Population Turnover in Remote Oceania Shortly after Initial Settlement.

Ancient DNA from Vanuatu and Tonga dating to about 2,900-2,600 years ago (before present, BP) has revealed that the "First Remote Oceanians" associated with the Lapita archaeological culture were directly descended from the population that, beginning around 5000 BP, spread Austronesian languages from Taiwan to the Philippines, western Melanesia, and eventually Remote Oceania. Thus, ancestors of the First Remote Oceanians must have passed by the Papuan-ancestry populations they encountered in New Guinea, the Bismarck Archipelago, and the Solomon Islands with minimal admixture [1]. However, all present-day populations in Near and Remote Oceania harbor >25% Papuan ancestry, implying that additional eastward migration must have occurred. We generated genome-wide data for 14 ancient individuals from Efate and Epi Islands in Vanuatu from 2900-150 BP, as well as 185 present-day individuals from 18 islands. We find that people of almost entirely Papuan ancestry arrived in Vanuatu by around 2300 BP, most likely reflecting migrations a few hundred years earlier at the end of the Lapita period, when there is also evidence of changes in skeletal morphology and cessation of long-distance trade between Near and Remote Oceania [2, 3]. Papuan ancestry was subsequently diluted through admixture but remains at least 80%-90% in most islands. Through a fine-grained analysis of ancestry profiles, we show that the Papuan ancestry in Vanuatu derives from the Bismarck Archipelago rather than the geographically closer Solomon Islands. However, the Papuan ancestry in Polynesia-the most remote Pacific islands-derives from different sources, documenting a third stream of migration from Near to Remote Oceania.

Genomic insights into the peopling of the Southwest Pacific.

The appearance of people associated with the Lapita culture in the South Pacific around 3,000 years ago marked the beginning of the last major human dispersal to unpopulated lands. However, the relationship of these pioneers to the long-established Papuan people of the New Guinea region is unclear. Here we present genome-wide ancient DNA data from three individuals from Vanuatu (about 3,100-2,700 years before present) and one from Tonga (about 2,700-2,300 years before present), and analyse them with data from 778 present-day East Asians and Oceanians. Today, indigenous people of the South Pacific harbour a mixture of ancestry from Papuans and a population of East Asian origin that no longer exists in unmixed form, but is a match to the ancient individuals. Most analyses have interpreted the minimum of twenty-five per cent Papuan ancestry in the region today as evidence that the first humans to reach Remote Oceania, including Polynesia, were derived from population mixtures near New Guinea, before their further expansion into Remote Oceania. However, our finding that the ancient individuals had little to no Papuan ancestry implies that later human population movements spread Papuan ancestry through the South Pacific after the first peopling of the islands.

The genetic history of Ice Age Europe.

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.

Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe.

How modern humans dispersed into Eurasia and Australasia, including the number of separate expansions and their timings, is highly debated [1, 2]. Two categories of models are proposed for the dispersal of non-Africans: (1) single dispersal, i.e., a single major diffusion of modern humans across Eurasia and Australasia [3-5]; and (2) multiple dispersal, i.e., additional earlier population expansions that may have contributed to the genetic diversity of some present-day humans outside of Africa [6-9]. Many variants of these models focus largely on Asia and Australasia, neglecting human dispersal into Europe, thus explaining only a subset of the entire colonization process outside of Africa [3-5, 8, 9]. The genetic diversity of the first modern humans who spread into Europe during the Late Pleistocene and the impact of subsequent climatic events on their demography are largely unknown. Here we analyze 55 complete human mitochondrial genomes (mtDNAs) of hunter-gatherers spanning ∼35,000 years of European prehistory. We unexpectedly find mtDNA lineage M in individuals prior to the Last Glacial Maximum (LGM). This lineage is absent in contemporary Europeans, although it is found at high frequency in modern Asians, Australasians, and Native Americans. Dating the most recent common ancestor of each of the modern non-African mtDNA clades reveals their single, late, and rapid dispersal less than 55,000 years ago. Demographic modeling not only indicates an LGM genetic bottleneck, but also provides surprising evidence of a major population turnover in Europe around 14,500 years ago during the Late Glacial, a period of climatic instability at the end of the Pleistocene.

Early Lapita skeletons from Vanuatu show Polynesian craniofacial shape: Implications for Remote Oceanic settlement and Lapita origins.

With a cultural and linguistic origin in Island Southeast Asia the Lapita expansion is thought to have led ultimately to the Polynesian settlement of the east Polynesian region after a time of mixing/integration in north Melanesia and a nearly 2,000-y pause in West Polynesia. One of the major achievements of recent Lapita research in Vanuatu has been the discovery of the oldest cemetery found so far in the Pacific at Teouma on the south coast of Efate Island, opening up new prospects for the biological definition of the early settlers of the archipelago and of Remote Oceania in general. Using craniometric evidence from the skeletons in conjunction with archaeological data, we discuss here four debated issues: the Lapita-Asian connection, the degree of admixture, the Lapita-Polynesian connection, and the question of secondary population movement into Remote Oceania.

Diet and human mobility from the lapita to the early historic period on Uripiv island, Northeast Malakula, Vanuatu.

Vanuatu was first settled ca. 3000 years ago by populations associated with the Lapita culture. Models of diet, subsistence practices, and human interaction for the Lapita and subsequent occupation periods have been developed mainly using the available archaeological and paleoenvironmental data. We test these models using stable (carbon, nitrogen, and sulfur) and radiogenic (strontium) isotopes to assess the diet and childhood residency of past communities that lived on the small (<1 km2) island of Uripiv, located off the northeast coast of Malakula, Vanuatu. The burials are from the initial Lapita occupation of the island (ca. 2800-2600 BP), the subsequent later Lapita (LL, ca. 2600-2500 BP) and post-Lapita (PL, ca. 2500-2000 BP) occupations, in addition to a late prehistoric/historic (LPH, ca. 300-150 BP) occupation period. The human stable isotope results indicate a progressively more terrestrial diet over time, which supports the archaeological model of an intensification of horticultural and arboricultural systems as local resources were depleted, populations grew, and cultural situations changed. Pig diets were similar and included marine foods during the Lapita and PL periods but were highly terrestrial during the LPH period. This dietary pattern indicates that there was little variation in animal husbandry methods during the first 800 years of prehistory; however, there was a subsequent change as animal diets became more controlled in the LPH period. After comparison with the local bioavailable 87Sr/86Sr baseline, all of the Lapita and LPH individuals appeared to be 'local', but three of the PL individuals were identified as "non-local." We suggest that these "non-locals" moved to the island after infancy or childhood from one of the larger islands, supporting the model of a high level of regional interaction during the post-Lapita period.

Lapita diet in remote oceania: new stable isotope evidence from the 3000-year-old Teouma site, Efate Island, Vanuatu.

Remote Oceania was colonized ca. 3000 BP by populations associated with the Lapita Cultural Complex, marking a major event in the prehistoric settlement of the Pacific Islands. Although over 250 Lapita sites have been found throughout the Western Pacific, human remains associated with Lapita period sites are rare. The site of Teouma, on Efate Island, Vanuatu has yielded the largest burial assemblage (n=68 inhumations) of Lapita period humans ever discovered, providing a unique opportunity for assessing human adaptation to the environment in a colonizing population. Stable isotope ratios (δ13C, δ15N, δ34S) of human bone collagen from forty-nine Teouma adults were analyzed against a comprehensive dietary baseline to assess the paleodiet of some of Vanuatu's earliest inhabitants. The isotopic dietary baseline included both modern plants and animals (n=98) and prehistoric fauna from the site (n=71). The human stable isotope data showed that dietary protein at Teouma included a mixture of reef fish and inshore organisms and a variety of higher trophic marine (e.g. marine turtle) and terrestrial animals (e.g. domestic animals and fruit bats). The domestic pigs and chickens at Teouma primarily ate food from a C3 terrestrial environment but their δ15N values indicated that they were eating foods from higher trophic levels than those of plants, such as insects or human fecal matter, suggesting that animal husbandry at the site may have included free range methods. The dietary interpretations for the humans suggest that broad-spectrum foraging and the consumption of domestic animals were the most important methods for procuring dietary protein at the site. Males displayed significantly higher δ15N values compared with females, possibly suggesting dietary differences associated with labor specialization or socio-cultural practices relating to food distribution.