"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.

Ancient DNA reveals five streams of migration into Micronesia and matrilocality in early Pacific seafarers.

Micronesia began to be peopled earlier than other parts of Remote Oceania, but the origins of its inhabitants remain unclear. We generated genome-wide data from 164 ancient and 112 modern individuals. Analysis reveals five migratory streams into Micronesia. Three are East Asian related, one is Polynesian, and a fifth is a Papuan source related to mainland New Guineans that is different from the New Britain-related Papuan source for southwest Pacific populations but is similarly derived from male migrants ~2500 to 2000 years ago. People of the Mariana Archipelago may derive all of their precolonial ancestry from East Asian sources, making them the only Remote Oceanians without Papuan ancestry. Female-inherited mitochondrial DNA was highly differentiated across early Remote Oceanian communities but homogeneous within, implying matrilocal practices whereby women almost never raised their children in communities different from the ones in which they grew up.

Frontier Lapita interaction with resident Papuan populations set the stage for initial peopling of the Pacific.

The initial peopling of the remote Pacific islands was one of the greatest migrations in human history, beginning three millennia ago by Lapita cultural groups. The spread of Lapita out of an ancestral Asian homeland is a dominant narrative in the origins of Pacific peoples, and although Island New Guinea has long been recognized as a springboard for the peopling of Oceania, the role of Indigenous populations in this remarkable phase of exploration remains largely untested. Here, we report the earliest evidence for Lapita-introduced animals, turtle bone technology and repeated obsidian import in southern New Guinea 3,480-3,060 years ago, synchronous with the establishment of the earliest known Lapita settlements 700 km away. Our findings precede sustained Lapita migrations and pottery introductions by several centuries, occur alongside Indigenous technologies and suggest continued multicultural influences on population diversity despite language replacement. Our work shows that initial Lapita expansion throughout Island New Guinea was more expansive than previously considered, with Indigenous contact influencing migration pathways and island-hopping strategies that culminated in rapid and purposeful Pacific-wide settlement. Later Lapita dispersals through New Guinea were facilitated by earlier contact with Indigenous populations and profoundly influenced the region as a global centre of cultural and linguistic diversity.

Fossils, fish and tropical forests: prehistoric human adaptations on the island frontiers of Oceania.

Oceania is a key region for studying human dispersals, adaptations and interactions with other hominin populations. Although archaeological evidence now reveals occupation of the region by approximately 65-45 000 years ago, its human fossil record, which has the best potential to provide direct insights into ecological adaptations and population relationships, has remained much more elusive. Here, we apply radiocarbon dating and stable isotope approaches to the earliest human remains so far excavated on the islands of Near and Remote Oceania to explore the chronology and diets of the first preserved human individuals to step across these Pacific frontiers. We demonstrate that the oldest human (or indeed hominin) fossil outside of the mainland New Guinea-Aru area dates to approximately 11 800 years ago. Furthermore, although these early sea-faring populations have been associated with a specialized coastal adaptation, we show that Late Pleistocene-Holocene humans living on islands in the Bismarck Archipelago and in Vanuatu display a persistent reliance on interior tropical forest resources. We argue that local tropical habitats, rather than purely coasts or, later, arriving domesticates, should be emphasized in discussions of human diets and cultural practices from the onset of our species' arrival in this part of the world. This article is part of the theme issue 'Tropical forests in the deep human past'.

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.

Exploitation and utilization of tropical rainforests indicated in dental calculus of ancient Oceanic Lapita culture colonists.

Remote Oceania, which largely consists of islands covered in tropical forests, was the last region on earth to be successfully colonized by humans, beginning 3,000 years ago. We examined human dental calculus from burials in an ancient Lapita culture cemetery to gain insight into the early settlement of this previously untouched tropical environment, specifically on the island of Efate in Vanuatu. Dental calculus is an ideal material to analyse questions of human and plant interactions due to the ingestion of plant-derived microparticles that become incorporated into the calculus as it forms throughout a person's life. Most of the microparticles identified here are from tree and shrub resources, including a ~2,900 calibrated (cal) BP example of banana in Remote Oceania, providing direct evidence for the importance of forests and arboriculture during the settlement of Remote Oceania.

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.

Ancient tortoise hunting in the southwest Pacific.

We report the unprecedented Lapita exploitation and subsequent extinction of large megafauna tortoises (?Meiolania damelipi) on tropical islands during the late Holocene over a 281,000 km2 region of the southwest Pacific spanning from the Vanuatu archipelago to Viti Levu in Fiji. Zooarchaeological analyses have identified seven early archaeological sites with the remains of this distinctive hornless tortoise, unlike the Gondwanan horned meiolaniid radiation to the southwest. These large tortoise radiations in the Pacific may have contributed to the rapid dispersal of early mobile Neolithic hunters throughout southwest Melanesia and on to western Polynesia. Subsequent rapid extinctions of these terrestrial herbivorous megafauna are likely to have led to significant changes in ecosystems that help explain changes in current archaeological patterns from Post-Lapita contexts in the region.

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.

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.

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.

Scurvy in a tropical paradise? Evaluating the possibility of infant and adult vitamin C deficiency in the Lapita skeletal sample of Teouma, Vanuatu, Pacific islands.

The Neolithic colonisation of the Pacific islands was one of the most challenging migration events in human history. The regions east of the Solomon Islands were colonised relatively recently by a people known as the Lapita. The Lapita brought with them a 'transported landscape' of domesticated plants and animals that had to be established upon arrival for the survival of these fledgling communities. Colonisation of these previously uninhabited islands was potentially perilous, and could leave colonisers vulnerable to periods of resource stress. The largest cemetery sample of Lapita people from the site of Teouma in Vanuatu offers a unique opportunity to assess the impact of colonisation on the health of pioneering populations. This paper explores the possibility that Teouma people experienced vitamin C deficiency as one of the consequences of the agricultural subsistence practices during the initial phases of island colonisation. Skeletal lesions in infants and adults indicative of scurvy suggest that initial colonisation phases in the Pacific islands involved precarious times involving deficiencies of key nutrients. Colonisation of the Pacific islands may share similar frameworks and problems as periods of subsistence transition in other parts of the world.

Megafaunal meiolaniid horned turtles survived until early human settlement in Vanuatu, Southwest Pacific.

Meiolaniid or horned turtles are members of the extinct Pleistocene megafauna of Australia and the southwest Pacific. The timing and causes of their extinction have remained elusive. Here we report the remains of meiolaniid turtles from cemetery and midden layers dating 3,100/3,000 calibrated years before present to approximately 2,900/2,800 calibrated years before present in the Teouma Lapita archaeological site on Efate in Vanuatu. The remains are mainly leg bones; shell fragments are scant and there are no cranial or caudal elements, attesting to off-site butchering of the turtles. The new taxon differs markedly from other named insular terrestrial horned turtles. It is the only member of the family demonstrated to have survived into the Holocene and the first known to have become extinct after encountering humans.

Phylogeny and ancient DNA of Sus provides insights into neolithic expansion in Island Southeast Asia and Oceania.

Human settlement of Oceania marked the culmination of a global colonization process that began when humans first left Africa at least 90,000 years ago. The precise origins and dispersal routes of the Austronesian peoples and the associated Lapita culture remain contentious, and numerous disparate models of dispersal (based primarily on linguistic, genetic, and archeological data) have been proposed. Here, through the use of mtDNA from 781 modern and ancient Sus specimens, we provide evidence for an early human-mediated translocation of the Sulawesi warty pig (Sus celebensis) to Flores and Timor and two later separate human-mediated dispersals of domestic pig (Sus scrofa) through Island Southeast Asia into Oceania. Of the later dispersal routes, one is unequivocally associated with the Neolithic (Lapita) and later Polynesian migrations and links modern and archeological Javan, Sumatran, Wallacean, and Oceanic pigs with mainland Southeast Asian S. scrofa. Archeological and genetic evidence shows these pigs were certainly introduced to islands east of the Wallace Line, including New Guinea, and that so-called "wild" pigs within this region are most likely feral descendants of domestic pigs introduced by early agriculturalists. The other later pig dispersal links mainland East Asian pigs to western Micronesia, Taiwan, and the Philippines. These results provide important data with which to test current models for human dispersal in the region.