Why do polynesians look like africans

Inland group locations are dark orange dots; shore group locations are light orange dots. Baining Mali and Baining Kaket are two dialects; elsewhere, the two Kaket-speaking locales are identified Rangulit and Malasait , as is Marabu Mali-speakers.

The Micronesians had low levels of inferred ancestry shared with populations in New Guinea, which is not far from Belau, where most of the Micronesian samples are from. This relationship is echoed in mtDNA results as well [ 35 ]. The typical ancestral proportions by population for a majority rule run are given in Table S6. Neighboring groups tended to share similar profiles. Bougainville groups had two common cluster assignments, while there was only one common cluster in New Ireland. Figure 9 shows the unrooted neighbor-joining tree for the East Asia—Pacific populations from a pairwise F ST coancestry distance matrix for microsatellites the pairwise F ST values are in Table S7.

Bootstrap values for the branches, generated with the PHYLIP program from population allele frequencies for different trees, are indicated by branch thicknesses. As shown, most of the trunk elements had high bootstrap values, as did a number of branches within Northern Island Melanesian groups.

By contrast, the mainland East Asian group relationships were considerably more ambiguous, their branches were shorter, and only the Taiwan Aborigines had a strong internal branch. These populations tend to be Papuan-speaking groups in island interiors. Neighbor-joining F ST -based tree for microsatellites from the Pacific, East Asia, and French populations, with the range of bootstrap values indicated by branch thicknesses.

New Britain populations are circled. Papuan-speaking groups are in bold italics; inland groups in Melanesia have asterisks. With our enlarged dataset and microsatellite coverage, we also compared patterns of private alleles and allele sharing between regions Table 3.

We recovered Melanesian-specific alleles, which in raw numbers actually exceeded those for Africa. Correcting for sample sizes, the rate of Melanesian-specific alleles was at the high end of the range for the major regions except for Africa. The number of alleles missing from only one continent, also given in Table 3 , shows the dramatic effect of genetic drift on the American populations. The number of shared alleles between pairs of regions is shown in Table 4 , with the correction for sample sizes in Table 5.

All non-African regions including Melanesia shared the most alleles with Africa, indicating they were primarily subsets of African diversity. Melanesia shared more alleles with East Asia than with any other non-African region, but they cannot simply be viewed as an extension or subset of East Asian diversity. When Papuan and Oceanic speaking groups in Melanesia were analyzed separately, the Papuan-speaking groups showed greater isolation, as they shared fewer alleles with all other regions than did Oceanic speaking groups unpublished data.

Our study suggests that in the Pacific, and specifically in Near Oceania, there is only a modest association between language and genetic affiliation. The result suggests that Oceanic languages were adopted by many formerly Papuan-speaking groups, while at the same time there was little genetic influence or marital exchange. At least in Near Oceania, rates of language borrowing and language adoption have been faster and more pervasive than rates of genetic admixture.

The size of the differences among the populations would appear to equal or surpass those among populations across East Asia, Europe, or even Africa. However, the large Melanesian population distinctions are a direct consequence of their very low levels of internal variation or heterozygosity. These low levels will directly inflate both the proportion of among group variation in AMOVA and also pairwise F ST genetic distances for a full discussion of this point, see especially [ 27 ] and also [ 26 , 37 ].

As population heterozygosities decrease, pairwise F ST s should increase because of this intrinsic mathematical relationship. The genetic distances used were the set of pairwise F ST s involving Bantu South the population with the highest heterozygosity , highlighted in Table S4. Our Structure and tree analyses of the combined microsatellite datasets indicate that Melanesians are quite far removed from Africans, in spite of their superficial similarities in hair form and skin pigmentation [ 38 ].

There, they split from Eurasia before Asians and Native Americans [ 39 ]. This also differed from the result of a genome-wide SNP study [ 40 ] on a very small world-wide dataset. The extreme positioning of Melanesians in our tree was not due to our over sampling.

Rather, our extensive coverage of Melanesian variation has enabled a clearer resolution of their relationships with populations outside the region. The pattern of Near Oceanic diversity has been made clear. The AMOVA analysis of the microsatellites showed that the larger and more rugged the island, the greater the differentiation among populations. Genetic variation from one large Near Oceanic island to the next was also significant. While our coverage of microsatellite variation elsewhere in the Pacific was admittedly spotty, our data as well as other smaller scale microsatellite analyses [ 21 , 41 ] suggest that, excluding the large islands of Near Oceania, there is a gradual decline in variation as one moves from Asia eastward, and variation among populations in the Pacific otherwise is not nearly as great as that in the large islands of Near Oceania.

Our sample coverage and definition was less rigorous there, and we expect equivalent coverage in New Guinea would equal or surpass the divergence of our New Britain series. The biogeographic pattern of population divergences in Near Oceania is most likely attributable to the restricted marital migration distances that have been documented most clearly for inland Bougainville groups [ 42 ], as well as for some New Guinea highlands populations [ 43 ].

Few people in small inland communities traditionally married and established households more than 1—2 kilometers from their birthplaces, while marital migration distances tended to be longer among shoreline communities. Nettle has argued that in ecologically rich tropical regions such as Near Oceania, small populations easily became self-sufficient, which in turn encouraged isolation and discouraged exchange [ 44 , 45 ], causing the development of extreme diversity among populations in both language and genetics.

We suggest this was the underlying cause of the short marital migration distances among inland groups in Near Oceania, which in turn was responsible for the low population heterozygosities and resulting large genetic distinctions among groups [ 42 ]. Because they arrived first and came to occupy large island interiors, the Papuan-speaking groups are considerably more diverse than Oceanic-speaking groups, which tend, in large islands, to be arranged along the shorelines.

The prehistoric record suggests there was a gradual reduction after initial settlement in the size of foraging zones of formerly mobile groups, associated with the filling up of the landscape [3, p. In many ways, these patterns and dynamics parallel the biogeography of birds and ants in the same region, where dispersal abilities of different species have dictated their patterns of diversity, and dispersal tendencies have, in many cases, contracted in island interiors over time [ 46 , 47 ].

The Tolai of East New Britain, with an assignment profile similar to New Ireland groups, are known to have migrated from southern New Ireland over 1, years ago [ 42 ]. Although the two Baining groups of east New Britain formed a cluster of their own, it has been suggested from the mtDNA, Y, and X chromosome analyses that they have been separated by thousands of years [ 48 ] see their long branch lengths in Figure 9. Also, the clustering of the Polynesians, Taiwan Aboriginals, and East Asians reflects ties older than 3, years.

In the Pacific, the change in genetic clustering apparently has evolved over thousands of years, and in many cases tens of thousands.

There were indications from the mtDNA, NRY, and certain autosomal microsatellites that in Remote Oceania, where islands are generally smaller in size, genetic variation among human groups is comparatively reduced, which is a contrast to Near Oceania [ 17 , 19 — 21 , 49 ].

At some point, prehistoric Oceanic mariners apparently became so accomplished that the inter-island water crossings in the central Pacific were often no more of an impediment to travel than the already occupied rugged terrain of the larger island interiors in the western Pacific.

In many areas, the ocean was transformed from a formidable barrier into a highway [ 50 , 51 ]. However, exactly where the relatively homogeneous Polynesians came from has remained controversial, and the number of proposed explanatory models for their origin form a continuum [ 49 , 52 ]. It suggests that, although there certainly must have been a series of introductions and influences from Asia into the Pacific over the millennia, no decipherable signal has survived that can be identified as specifically ancestral to Polynesians, because of the complexities of human interactions from the outset [ 54 ].

Proponents argue that tree-like representations of population or linguistic relationships cannot be expected to develop regularly and are likely to be entirely inappropriate representations of population relationships in many, if not all, instances, since they so often ignore interactions between neighboring groups. Models at the other end of the continuum assume contemporary genetic as well as cultural similarities can carry a clear signal of past population relationships.

It proposes a rapid movement of the ancestors of the Polynesians from the vicinity of Taiwan to the Central Pacific, without extensive contact with indigenous Near Oceanic populations along the way.

At present, the state of the evidence for this association is as follows: a the precursor haplotype to B4a1a1 has been identified in Taiwan aboriginal populations [ 56 ]; b the final development of B4a1a1 with the key mutation at nucleotide site seems to have occurred in eastern Indonesia or Near Oceania [ 17 ]; c its frequency varies widely over Near and Remote Oceania before becoming ubiquitous in Central and Eastern Polynesian populations; d in Near Oceania, it is common along many Oceanic-speaking coastal groups, as well as a number of Papuan-speaking groups, especially in New Ireland and Bougainville [ 17 ]; and e its expansion dates are relatively recent, although old enough to suggest to some observers that it cannot be easily tied to the Polynesian expansion [ 17 , 56 ].

Because of their comprehensive nature, we believe the results of our autosomal microsatellite survey present a resolution to this issue with regard to human genetic relationships. The fact that the STRUCTURE cluster containing Micronesians, Samoans, and Maoris has a detectable signature only in Oceanic-speaking Melanesians and Taiwan Aborigines supports the position that an expansion of peoples from the general vicinity of Taiwan is primarily responsible for the ancestry of Remote Oceania, and that these people left a small but still identifiable signature in some Oceanic-speaking populations of Near Oceania.

Scenarios for different male and female dispersals have been proposed to reconcile the divergent mtDNA and NRY patterns in Oceania [ 35 , 59 ], but the autosomal microsatellite results should now serve as the primary reference. Although the Polynesians in our analysis were similar to Taiwan Aborigines and East Asians, they might be even closer to other populations not covered in our study, from Indonesia, the Philippines, or Southeast Asia.

While there is a substantial body of evidence that indicates Taiwan is the primary point of Austronesian dispersal [ 60 , 61 ], there are now also suggestions of the importance of Island Southeast Asia as well [ 62 , 63 ]. The ties of particular Near Oceanic populations to those regions also remain poorly understood, but should be resolved with additional sampling from these regions and similar analyses. To revisit the questions posed at the beginning, we can provide answers as follows.

Outside the Pacific, East Asian populations are apparently the closest but still very distant relatives of Melanesians. Africans and Europeans are the most distant. The within-group diversity in Melanesian populations is consistently very low, which acts to exaggerate the considerable among-group distinctions there.

This great diversity in such a small region makes comparisons of human population structure from continent to continent problematic. The diversity among groups is primarily organized by island size and topographic complexity, with the inland Papuan-speaking groups the most isolated and differentiated. Shore-dwelling Oceanic-speaking groups are more intermixed dispersal along the shorelines was easier.

The sailing capabilities of the ancestors of the Polynesians transformed the nature of their Diaspora and kept them relatively homogeneous. Our Asia—Pacific sample set came from a variety of sources. The objective was to include between 15 and 25 unrelated individuals minimally excluding reported first-degree relatives from locales where individuals and their parents had all lived.

These criteria were achieved in most instances. Details are given below. Samples from Northern Island Melanesia were collected in three field seasons , , and in collaboration with the Institute for Medical Research of Papua New Guinea.

Besides a 10 ml blood sample, a simple genealogy and residency questionnaire was taken, including in most instances parent and grandparent names, residences, and native languages.

Among over 1, samples collected, were chosen for submission to the Marshfield Clinic for microsatellite and indel analysis. We included multiple locales in larger language groups where feasible; and picked samples from individuals whose family's residence histories suggested close identification with the sampling locale. People of mixed parentage especially with one grandparent from a different language group or island could not always be excluded if the minimum required sample size was to be achieved.

DNA was extracted as previously described [ 17 ]. All individuals gave their informed consent for participation. DNA was extracted from blood using Qiagen kits. Taiwan Aboriginal samples comprise the Northeastern Taroko tribe from Hsiulin, part of the Atayal language group, and the Amis tribe living on the east coast of Taiwan and speaking Amis. All individuals were unrelated and had both parents belonging to the same tribe.

Each individual gave informed consent to participation in population genetics studies and the project was approved by the Ethics Committee of the Hospital and the Department of Health of Taiwan.

Blood samples were collected in acid citrate dextrose tubes. The microsatellites were drawn from Marshfield Screening Sets 16 and 54, and the indel markers were drawn from Marshfield Screening Set Where the primer changed, allele sizes from one of the two data sets were adjusted Table S9.

The expected heterozygosity and average number of alleles per locus were computed on the microsatellites with the GDA software [ 65 ], using the sample-size corrected estimator, as in [ 66 ]. F ST was estimated on the microsatellites as in Equation 5. Cluster analysis of genotypes utilized the Structure versions 2. Results using Structure 2. When multiple runs at the same values of K produced discrepant results, we relied on majority rule i.

Details are provided in the Tables S3 and S5. Individual similarity coefficients for pairs of runs were calculated as in [ 24 ] and Methods. Great circle geographic distances were calculated with the Haversine method as described in [ 26 ]. After Blust [ 10 , 71 , 72 ] and Pawley personal communication. We are greatly indebted to the people from the different parts of Oceania who collaborated so willingly with us in this project. We hope this paper will help to illuminate their population histories and relationships, as we promised them at the outset.

We thank Jeff Long and two anonymous reviewers for their suggestions, which have considerably strengthened the paper. We thank Andrew Pawley, Glenn Summerhayes, and Peter Bellwood for suggestions on the historical linguistics and prehistory. Sarah Tishkoff suggested the genotyping of these samples at the Marshfield Clinic and was involved in a number of helpful subsequent discussions.

Author contributions. JLW supervised genotyping at the Marshfield Clinic. Competing interests. The authors have declared that no competing interests exist. National Center for Biotechnology Information , U. PLoS Genet. Thank you for visiting nature. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser or turn off compatibility mode in Internet Explorer. In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

The debate concerning the origin of the Polynesian speaking peoples has been recently reinvigorated by genetic evidence for secondary migrations to western Polynesia from the New Guinea region during the 2nd millennium BP.

These long-distance migrants into Polynesia experienced additional admixture with northern Melanesians prior to the secondary migrations of the 2nd millennium BP. Moreover, the genetic diversity of mtDNA and Y chromosome lineages in the Leeward Society Islands is consistent with linguistic evidence for settlement of eastern Polynesia proceeding from the central northern Polynesian outliers in the Solomon Islands. These results stress the complex demographic history of the Leeward Society Islands and challenge phylogenetic models of cultural evolution predicated on eastern Polynesia being settled from Samoa.

He suggested that the similarity of the languages spoken there, now known as Polynesian, reflected a comparatively shallow time-depth since their dispersal 1. This appeared to be the only explanation for the striking difference in phenotype that he observed between the peoples of the central Pacific and those of the intervening region, which is now known as Melanesia.

Herein, the terms Melanesia and Micronesia are used in their geographical sense. We use the term Polynesia to include all islands and atolls whose inhabitants speak Polynesian languages, including 23 found throughout Melanesia and Micronesia, referred to as outlier Polynesia Fig. Sampling locations and overview of genomic diversity. Colours indicate regional affiliation of populations used for analysis of autosomal DNA: orange — mainland Southeast Asia and East Asia; dark blue — Taiwan; brown — Philippines Aeta, Agta and Batak negritos; light blue — Philippines non-negritos; red — western Indonesia; pink — eastern Indonesia; purple — northern Melanesia and New Guinea; black — Australia; green —Polynesia.

The usage of populations varies with the type of analysis employed Supplementary Table S1. The red circles within Micronesia and Melanesia represent 20 of the atolls and islands referred to collectively as outlier Polynesia. The red stars denote the three additional Polynesian outlier populations Rennell and Bellona, Tikopia , which together with Tonga, were used in analysis of ancient admixture by Skoglund, et al.

Detailed sample information is given in Supplementary Table S1. The map was created using R v. R: A language and environment for statistical computing.

The critical differences are the position of the East Polynesian languages relative to the rest of nuclear Polynesian, and their relationship to the Central Northern Outlier languages. In the sub-grouping according to Pawley 31 all the Polynesian Outlier languages group within Samoic implying an early separation of Proto-East Polynesian from the rest of the Nuclear Polynesian languages.

In the alternative sub-grouping proposed by Wilson 32 the Central Northern Outlier languages group with the languages of East Polynesia, within a larger clade containing the other Northern Outlier languages.

Separating the demographic histories of Polynesia and Melanesia became difficult to sustain with developments in archaeology during the second half of the 20th century. At the same time, the study of comparative linguistics has shown that the Oceanic branch of the Austronesian phylum of languages, of which Polynesian is a member, is spoken throughout most of Melanesia and parts of coastal New Guinea, and appears to be a recent intrusion from ISEA 5.

So while there is considerable overlap between the distributions of the LCC and the Oceanic languages, there remains a phenotypic divide between southern Melanesia and western Polynesia, which is observed between Fiji and Tonga 6 , 7.

A central theme in this debate is the extent to which the development of the LCC involved local people in the Bismarck Archipelago of northern Melanesia 8 , 9 , An alternative is that the LCC represents the arrival of a largely pre-formed cultural package carried by speakers of proto-Oceanic languages from Taiwan, via the Philippines, in ISEA Hypotheses are placed on a continuum from a dendritic, radiating, phylogenetic model of cultural evolution that relies on the relative isolation of populations 12 , to one based on complex ongoing biological and cultural interaction between groups, leading to reticulated networks of genes and culture 9.

Genetic evidence for this intermediate model is provided by the presence of members of Y chromosome haplogroup hg C2a-M, together with its daughter lineage C2a1-P33, among Polynesian speakers 16 , This is seen as a proxy for male-mediated admixture from northern Melanesian and New Guinean sources into the gene pool of migrants from ISEA during the formative period of the LCC in the Bismarck Archipelago, prior to the settlement of southern Melanesia and western Polynesia 13 , In contrast, the near fixation in Polynesian speaking groups of the mitochondrial lineage B4a1a1 is seen as evidence of a predominantly ISEA maternal heritage 13 , Subsequent research, however, has shown that B4a1a1 is widespread throughout northern Melanesia 20 , including regions that show no evidence of autosomal admixture with people from ISEA Alternatively, therefore, hg B4a1a1 might also have been present in northern Melanesia before the emergence of the LCC 22 , Similar ambiguity now exists over the origins of paternal lineage C2a-M, due to its presence in ISEA 24 and rather low overall frequencies in the Bismarck Archipelago and coastal New Guinea These data show that some settlers of the LCC period appear to have transited northern Melanesia and New Guinea from ISEA without receiving any significant amounts of genetic admixture.

This result is consistent with post-LCC movements of people into southern Melanesia and western Polynesia, in a process of polygenesis, being responsible for the differences in phenotype observed between the two regions 6. The potential significance of this proposed post-LCC migration for the phylogenetic approach to cultural evolution cannot be overstated.

The settlement of eastern Polynesia, however, has witnessed significant reductions in the earliest secure radiometric dates in recent years. A central component of the original phylogenetic model is the long-standing sub-grouping of the Polynesian languages.

The initial divergence of Nuclear Polynesian from the Tongic languages is followed by a second-order split, between Proto-East Polynesian Rapa Nui, Marquesan and Tahitic and the rest of the Nuclear Polynesian languages Samoic and all the Polynesian outlier languages 31 Fig.

An alternative linguistic sub-grouping that places the East Polynesian languages together with those of the central northern outliers east coast of the northern Solomon Islands provides a potential solution for the apparent discordance between archaeology and language 32 , 33 Fig. This also challenges the orthodoxy within Polynesian studies that eastern Polynesia was settled directly from Samoa 11 , 12 , The Leeward Society Isles, therefore, are of central importance to understanding the reasons for these conflicting signals from archaeology and language.

If the ancestors of the Leeward Society Islanders experienced the same episode of ancient admixture as people in western Polynesia and outlier Polynesia during the mid 2 nd millennium BP 25 , this would support the late settlement chronology. We use the analysis of genotype and haplotype data to ascertain whether the signals of admixture present in these eastern Polynesian populations are similar to those from western and outlier Polynesia and identify potential donors to the ancestors of the Leeward Society Islanders.

Further insights into the demographic history of eastern Polynesia is provided by the first deep re-sequencing of Polynesian Y chromosomes, complemented by high-resolution genotyping of key paternal and maternal lineages from the Leeward Society Isles and New Zealand.

Here we present a new genomic dataset sampled from the Leeward Society Islands, eastern Polynesia. For detailed information about samples used in the present study, please refer to the Materials and Methods section.

The close overlap between eastern Polynesians and Samoans on the PC1 axis suggests similar amounts of genetic ancestry shared with New Guinea and northern Melanesia. However, the latter is unlikely due to the lack of close relatives up to third-degree, inclusive in four Polynesian groups, and normal range of inbreeding coefficients when comparing to other human populations F IS , Supplementary Table S6. Details of the populations are provided in Supplementary Table S1B.

The colors used have been selected to be equivalent to those used in Fig. Populations from the Philippines can be generally divided into Negritos Aeta, Agta and Batak , Kankanaey of northwestern Luzon, and all others representing an amalgamation of groups from Luzon, Palawan and Visayas see Fig.

S6 , indicating the heterogeneous distribution of East Asian and European ancestry among the Leeward Society Islanders. The distinctive and uniform profiles of additional ISEA, Melanesian, and East Asian ancestries in two Tonga and Samoa out of four, otherwise very closely related, Polynesian groups hint that these may be the result of an old admixture process, rather than genetic drift, extreme bottlenecks or algorithmic artifacts.

In contrast, the noticeably uneven distribution of the East Asian yellow and western European grey ancestry components within the profiles of the Leeward Society individuals Fig. The outgroup f 3 36 allele-sharing plot shows the length of a phylogenetic branch shared between two study populations and African Yoruba.

S3 , Supplementary Table S7 , the f3 allele-sharing results are consistent with a most recent evolutionary history shared with Samoa, Tahiti, and Tonga.

It also suggests that the Kankanaey of the Philippines and Taiwanese aborigines are the next closest populations to all four Polynesian groups. In contrast, the f 3 admixture plots Supplementary Fig. S5 , Supplementary Table S7 , which detect the presence of admixture in a study population from two reference groups, display different results for western and eastern Polynesia.

These differences could be explained by a reduced effective population size for eastern Polynesians, caused by bottlenecks during the initial settlement process, or because Tonga and Samoa have experienced additional admixture since they last shared a common ancestral gene pool with Tahiti and the Leeward Society Isles.

Population admixture events and inferred contact dates. Each color represents a separate genetic cluster estimated with FS that acts as a donor to the recipient cluster Leeward Society Isles in the GT analysis.

There is strong statistical support for two episodes of admixture; the ancient and recent events are represented by the left- and right-hand bar plots, respectively. Each episode involves two pairs of sources minor and major ; bar plots depict the inferred composition of the mixing sources for each, with admixture dates calculated using a generation time of 28 years. Detailed information about the inferred admixture episodes and composition of mixing sources is given in Supplementary Table S8.

In order to investigate the presence of the northern Melanesian contributions further, we performed a GT analysis on different subsets of Leeward Society Islanders as recipient groups Supplementary Fig.

S7 , Supplementary Table S8. Some of the variability in the dating may be due to the heterogeneous distribution of what appears to be recent admixture with people of East Asian ancestry Figs 2B , S7 , Supplementary Table S8 , but the result is robust to variations in the makeup of the recipient group. A further insight from the FS and GT analyses of haplotypes is the clear delineation between possible donor groups within the Philippine palette of populations.

The apparent discrepancy with the analysis of unlinked SNPs Supplementary Figs S3 and S4 , which indicates the Kankanaey as being closest to the Leeward Society Isles, may be caused by the two methods measuring different aspects of the underlying genetic structure. In addition, ascertainment bias inherent to genotyping arrays data can affect the allele-sharing statistics.

The GT analysis, in contrast, is based on combinations of linked markers, and is consequently more powerful and robust for identification of complex admixture events Ninety-six percent of Leeward Society Isles mitochondrial lineages belong to the haplogroup hg B4a1a1 typical of Polynesian speaking populations. The Bayesian estimate of the time to the most recent common ancestor MRCA for well-supported clades of mitochondrial hg B4a1a1 Supplementary Table S10A is consistent with more than a third being significantly older than the first settlement of southern Melanesia and western Polynesia.

S11 , Supplementary Table S10B. Seven of the eight individuals belong to the O3i-B clade, which so far has only been identified among Austronesian speakers in ISEA 39 , S11 , Supplementary Table S3. These individuals, therefore, likely belong to hg O3i. S11 and its legend online, Supplementary Table S9B. How this came about is the subject of considerable debate 9 , 11 , 12 , These genetic dates are nearly identical to those of a previous analysis that used a different method and amalgamated haplotype data from western Tonga and outlier Rennell, Bellona and Tikopia Polynesia The method used here has been demonstrated to accurately identify known historical admixture events during the past 2, years 38 , 44 , but it is also possible that other analytical approaches may provide insights into a different part of the genealogical process.

The substantial body of linguistic evidence supporting this sub-grouping includes over lexical and grammatical innovations that are shared between the languages of eastern Polynesia and the central northern outliers Luanguia, spoken on Ontong Java, Takuu, Nukumanu and Nuguria.

Moreover, these innovations are stepwise and directional in nature, a pattern that is only consistent with a west-to-east movement of people, tracing the origins of eastern Polynesians to central northern outlier Polynesia, rather than Samoa 32 , The principal component analysis and phylogenetic reconstruction of the Polynesian mtDNA B4a1a1 sub-groups and C2a1-P33 paternal lineages Supplementary Figs S8 — S10 , S12 , are consistent with this linguistic evidence for the recent settlement of eastern Polynesia from the central northern outliers.

This result is robust to analysis by subsets of the data Supplementary Fig. S7 , but it is not possible to determine how and when this northern Melanesian ancestry entered into the ancestral gene-pool of the Leeward Society Islanders. It is arguable that one or other result is misleading as an effect of severe genetic drift. However, this hypothesis requires the concurrent excess retention of either SNPs should f 3 results be taken at face value , or haplotypes should we trust only GT , typical of those found in the Leeward Society Islands today, which is statistically unlikely.

This hypothesis is preferred because the GT approach models the recipient population using donors who are reconstructed rather than observed, allowing for subsequent admixture in the donor groups Within the geographical context of the Philippines, the GT results make sense because the populations making up the other three Philippines clusters are all located in mountainous regions and have languages that are either relics or indicate long-term isolation 46 , In contrast, the ancestors of the demographic expansion that led to the settlement of Polynesia are anticipated to be part of a recent seafaring tradition.

This necessarily would have been based in the coastal regions and could be related to pre-existing trading networks within ISEA that already had links to Melanesia see Donohue and Denham 48 and comments for a discussion of this subject. This sex bias holds across Polynesia and is observed as far back as Island Southeast Asia 49 , and may have resulted from the practice of exogamy and matrilocal post-marital residence among early Austronesian speaking groups A sex bias is also reflected in the nuclear genomes of Austronesian speakers and appears to be a characteristic of the Pacific region as a whole 25 , In conclusion, the picture of Polynesian origins emerging from the present study is one of a more complex demographic history than that originally envisioned in the phylogenetic model of cultural evolution The results presented here provide support for models based on inter-connectivity among, and within, the different parts of the Pacific, rather than their relative isolation 8 , 9.

The new data concur with a late chronology for the settlement of eastern Polynesia, which fits better with the linguistic arguments and haploid data linking this region to the northern central Polynesian outliers. With respect to the ultimate origin of the Island Southeast Asian ancestry found in the Leeward Society Isles, the results indicate a significant role for the lowland region of the Philippines, as predicted by Johann Reinhold Forster in his seminal comparative study of languages conducted more than two hundred and forty years ago.

All samples were collected with informed consent and with the approval of the institutional review boards at the University of Colorado, U. All experiments were performed in accordance with the relevant guidelines and regulations of the collaborating institutions.

Nomenclature followed that of Phylotree. This resulted in 78 individuals allocated to the hg B4a1a1 and three individuals to hg Q. The control region nps 57— and nps — was sequenced for 36 samples that could not be allocated to the known sub-clades of these two haplogroups, and 25 samples were further selected for complete mitochondrial genome sequencing. Using information from the full sequences, additional nucleotides were typed by Sanger sequencing to complete the haplogroup assignment.

The data sets were partitioned into the D-loop, rRNA genes, and first, second, and third, codon positions of the 13 protein-coding genes. Each data subset was assigned an independent model of nucleotide substitution, selected using the Bayesian information criterion in PartitionFinder Four demographic models for the tree prior were compared: constant size, exponential growth, logistic growth, and Bayesian skyride coalescent 59 , together with two models of rate variation across lineages: strict clock and uncorrelated lognormal relaxed clock For the B4a1a1 analysis, the best combination was a strict clock with a logistic growth coalescent model.

For the MQ analysis, the combination of strict clock and Skyride model is reported because the demographic model showed a clear change in population size.

To calibrate the estimate of the timescale, a normal prior for the mutation rate was specified mean 2. The posterior distributions of parameters, including the genealogy, were estimated using MCMC sampling. To check for convergence, each analysis was run in duplicate. After checking for acceptable MCMC mixing, the samples from the two runs were combined.

Sufficient sampling was checked by computing the effective sample sizes of all parameters, which were found to be greater than Haplotypes were assigned to sub-clades of the hg B4a1a by the HaploGrep2 software 62 and manual inspection of sequences. Unless otherwise stated, all nomenclature follows that of Karmin, et al.

The Y chromosomes of nine individuals belonged to haplogroups typical of Europeans G, J, and R and were not subject to further analysis. These samples were also hierarchically tested to a level of phylogenetic resolution equivalent to the main haplogroup level in the Leeward Society Islanders Supplementary Table S3.

The same 15 microsatellites occurring on the C2a-M background were used to perform PCA in R 63 , in order to examine the relationship of eastern, western and outlier Polynesia populations for this key haplogroup within Polynesian Y chromosome diversity Supplementary Fig. The paired-end reads were mapped to the GRCh37 human reference sequence.

The reconstruction and rooting of the phylogeny of the seven samples from the Leeward Society Islands used 56 sequences published in Karmin, et al. This resulted in 6. To correct for ascertainment bias, we added constant sites corresponding to the nucleotide composition across the remainder of the chromosome. Gmail is finally back up following a five HOUR outage that left frustrated users unable to send or receive Israel shows off new electronic warfare system that uses BEAMS instead of missiles or bullets to go after Unsuspecting Colorado homeowner finds black-footed ferret, the 'rarest mammal in North America,' scampering Mass grave with more than 25 skeletons of men, women and children dating back to the 15th century is found Catch of a lifetime!

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Ad Feature 'My job is done now! Apple's new iPad is blazingly fast, gorgeous to look at, and quite simply the best tablet out there - and for a lot of people, probably the best computer out there. The small smart display with big potential: Google Home Hub review. Google is late to the game with its Home Hub, but the low price and AI features make it a great choice for controlling your home, showing pictures and even helping run your life.

On one hand, the XR lacks the high-resolution screen and dual-lens camera on the XS. AI seems to permeate every part of its software, from the ability to answer calls for you to being able to almost perfectly predict your morning commute. Apple's new iPhone XS and XS Max go on sale on Friday - and the biggest handset Apple has ever made is also its best and possibly unsurprisingly, its most expensive.

Israeli beauty-tech firm Pollogen has launched its Geneo Personal device, which stimulates oxygen from beneath the skin's surface to give you a clearer, fresher face within minutes. Rather than cram in a plethora of new features, Apple's latest update is about boosting stability, with improvements in everything from FaceID and battery life. Naim Atom: The hifi that will change the way you listen to music. Naim's incredible Mu-So Qb takes you back to the good old days - where the music captivates and enthralls, rather that simply being something in the background.

Peloton's hi-tech bike lets you stream live and on demand rides to your home - and it's one of the best examples of fitness technology out there - at a price. The best all in one wireless speaker you'll ever hear: Naim Mu-so review. Discover deals on home essentials and electricals. Apply AO. Very - Very deals. Keep yourselves entertained with these electrical offers.

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