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Author contributions: Physical resemblance between Polynesians and Asian populations was detected, but in between were the islands of Melanesia, inhabited largely by people of different phenotype. However, the Lapita culture bridged this geographical divide 3, y ago. Morphological studies of early Lapita colonists from Teouma, Vanuatu align them with present-day Polynesian and Asian populations, whereas skeletal remains of later generations show a more Melanesian phenotype predominating.

We suggest that migration streams from already-inhabited parts of Melanesia dating from the late-Lapita phase ultimately dominated the original Polynesian phenotype in eastern Melanesia, but not in Polynesia, which became relatively isolated soon after initial settlement. 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 first human settlement of Vanuatu is indicated by the Lapita culture, whose earliest signature appears in the northwestern Melanesian islands toward the end of the interval 3,—3, y B. The Lapita culture is defined by a set of artifacts including highly decorated pottery displaying a distinctive design system, long-distance exchanges of raw material and finished items, translocations of plants and animals, and the initial incursion of humans into the pristine island environments of Remote Oceania to the east of the main Solomon chain between 3, and 2, y B.

In Vanuatu, as in the rest of Remote Oceania, Lapita quickly evolved, within — y, into distinctive local cultures in conjunction with increased population size and sedentism by the end of the Lapita period 3. More generally, the results indicate two contrasting divisions, an Australo-Melanesian pole comprising groups from the western part of Remote Oceania Island Melanesia and an Asian pole including groups from the far eastern part of Remote Oceania Polynesia.

This pattern suggests separate origins for the indigenous inhabitants of these two regions. Evidence from inherited genetic markers indicates that the populations living today in Vanuatu and generally in the region first settled by Lapita groups share a common origin in an area that encompasses Island South East Asia, the north coast of New Guinea, and the Bismarck Archipelago 7 — Geographical variations in haplotype frequencies distinguish the western part of the initial Lapita region from the eastern part, with a smaller diversity in the eastern populations in what is today Western Polynesia.

In a recent biodistance study of mandibles from Watom New Britain , Pietrusewsky et al. Past haplotype distribution reconstructed with ancient DNA aDNA data obtained from skeletal remains representing early human groups may theoretically be a means to investigate the issue. However, published data on prehistoric Pacific Islanders are sparse and mainly centered on Polynesia Regarding the Lapita question directly, the current aDNA results include only one individual representing the first thousand years of settlement in Vanuatu [Efate, Taplins, midlate third millennium B.

Although consistent with the morphological evidence 15 , 16 , 27 , these pioneering results are still regarded as uncertain Large-scale excavations at the Teouma site have revealed a Lapita cemetery with 68 burial features where adults were preferentially treated by inhumation However, inhumation was temporary; bones, including skulls and mandibles, were removed from burials postdecomposition and redeposited at the site but in a much smaller number than the incomplete inhumations recovered to date The extant cranial elements consist of seven skulls in secondary deposits B10 cache, B17, and B30 Fig.

Although disconnected from the infracranial skeletons these cranial remains seem to have belonged to individuals of the same group, as shown by similarities in isotopic values measured in bone collagen 29 and in dental enamel In the current study, we use five of the seven skulls labeled TEO 10a, 10b, 10c, 17, and 30a that, after reconstruction, are almost complete and suitable for metric study Fig.

To assess the biological affinities of these early Vanuatu settlers, we have generated two distinct comparison datasets SI Materials and Methods. The first is a composite sample of 12 archaeological adult specimens that were chronologically selected to help in evaluating Lapita-associated population origins. Craniofacial shape affinities were assessed with series of multivariate analyses, including principal component analyses PCA on size-corrected measurements of the neurocranium and upper face and linear discriminant analyses LDA , which have been used to predict the classification of the archaeological specimens, including the five Teouma individuals SI Materials and Methods and Table S4.

For this study, we have generated two distinct comparison datasets. Four of the seven Australian specimens from Coobool Creek included in this study have been described as being altered—or possibly altered—by artificial cranial modification by head binding 34 , Although archaeological specimens are not taken into account in computing the discriminant functions, particular attention was paid to the results obtained on these individuals. Other Lapita and immediately post-Lapita specimens from Near and Remote Oceania, especially from Vanuatu, were not included due to limited preservation of the cranium precluding a similar set of measurements Table S1.

The second dataset consists of nine samples selected from the publicly available database of Howells 33 , representing five geographically distinct modern and near modern populations Table S3. This dataset includes adult individuals [detailed descriptions of the samples are provided by Howells 63 ]. Previous craniometric studies have indeed repeatedly shown that samples from geographical Melanesia systematically clustered with Tasmanian and Australian samples whereas East and West Polynesian samples clustered with Southeast Asian and East Asian samples 5.

Most of the specimens are devoid of archaeological context with the exception of two series. Sex was generally diagnosed by Howells himself by inspection of cranial criteria, cross-checked by observations of the postcranial skeleton where available The original geographical subsamples are not all balanced between males and females, and the smallest subsample Micronesian females is limited to 27 specimens Table S3.

Because the sex of the Teouma specimens is not cross-checkable using pelvic elements individuals with a pelvis have no skull and skulls are not associated with a pelvis and the cranial sexual dimorphism of early Lapita population is unknown, the present study includes morphometric comparisons with pooled-sex samples. To undertake the morphometric analyses on well-balanced comparison samples for sex and geography , subsamples of 54 specimens 27 males and 27 females without missing values were randomly generated from the original samples of Howells for the five geographical groups.

This is especially important for linear discriminant analysis that is usually highly sensitive to unbalanced reference samples In the analyses, missing values were not statistically estimated and specimens with missing values are thus excluded. Statistical exploration of the whole dataset guided the selection of measurements that best allow investigation of the affinities of the Vanuatu specimens. This was done under the double perspective of i having the smallest number of missing values for the Teouma specimens and ii selecting measurements that make sense in terms of craniofacial shape variation.

Eleven measurements were selected for the analyses presented here Table S4. Log shape ratios LSRs were used instead of raw linear measurements to better distinguish shape differences from size differences Indeed, it is well known that size variations are always dominant over shape variations in linear morphometrics, whereas the latter are often more informative in terms of biological affinities. It is also generally accepted that controlling the dominance of size variations in the analyses should reduce the effects of form differences related to sexual dimorphism, even though potential allometric relationships are not eliminated per se by the LSR transformation [for a comprehensive review and discussion of size-adjustment procedures in morphometrics see Jungers et al.

With LSR, one degree of freedom is lost due to the scaling of measurements. To reduce the dimensionality of the original dataset, PCA was first conducted on recent specimens, the five specimens from Teouma and the 12 other archaeological specimens using the 11 LSRs. Then LDA was conducted with geographical origin as group factor on the recent specimens, using the individual scores on the first 10 PCs. The percentage of correctly assigned specimens was computed using a leave-out-one cross-validation procedure.

Finally, the resulting discriminant functions were used to predict the classification of the 17 archaeological specimens, including the five Teouma individuals. LSRs, statistical analyses, and graphics were computed with R www. The scatter plot of specimens on PC1 vs. However, the variation along PC1 distinguishes Australian and Melanesian specimens that mainly plot toward negative values from Chinese, Western Micronesian, and Polynesian specimens that mainly plot toward positive values.

Australian and Melanesian individuals thus tend to display relatively long skulls especially the parietal cord and upper faces with relatively short and wide nasal apertures and low orbits. Chinese, Western Micronesian, and Polynesian individuals present the opposite morphological characteristics. The post-Lapita Vanuatu individual from Mangaliliu presents rather different craniofacial shape affinities because it plots closer to recent Australo-Melanesian groups and archaeological Australian and pre-Neolithic Southeast Asian individuals.

PC2; however, neither of these two individuals is described as artificially modified by head binding PCA on the log-shape ratios of 11 craniofacial variables for recent and 17 archaeological specimens: The LDA is able to correctly discriminate the five geographical groups of the recent specimens. The scatter plot on LD1 vs. LD2 mainly distinguishes two major geographical groups along LD1 Fig.

Australian and Melanesian individuals plot toward negative values of LD1, whereas Chinese, Micronesian, and Polynesian individuals plot toward the positive. Discriminations inside each of these two major geographical groups are mainly expressed along LD2, but at lower magnitudes. Australians are separated with some overlap from Melanesians that tend to have positive values for LD2. The same trend is observed for Polynesians that have generally higher values for LD2 than Chinese and Micronesians.

These two last groups are largely overlapping on LD1 vs. The predicted values on LD1 and LD2 of the 17 archaeological specimens with no a priori geographical assignations fall well inside the overall variation of the recent specimens. LD1 vs. The cross-validation procedure indicates that The results indicate If one takes into account the two larger geographical groups that are distinguished in the PCA and the LDA scatter plots i. This implies that despite a significant percentage of incorrect assignations the discriminant functions computed with the 11 craniofacial shape variables for recent specimens divided into five geographical groups provide a good discrimination between all of the groups, and, at a higher geographical level, a very good discrimination between an Australian and Melanesian pole and a Chinese, Micronesian, and Polynesian pole.

Discriminant analysis: Assignations of specimens to geographical groups after cross-validation. The results of the predicted classifications of the 17 archaeological specimens computed with the discriminant functions are presented in Figs. The eighth Australian archaeological specimen, Coobool Creek 82, presents a completely different profile of its predicted assignation. This particular specimen has been systematically identified as artificially modified in previous work 34 , 35 , which could explain its particular craniofacial shape.

Bar plot of the predicted probabilities of posterior assignations to the five geographical groups: A Australian archaeological specimens; B Javanese archaeological specimens; C Pacific archaeological specimens. Teouma archaeological specimens: The two archaeological specimens Song Keplek 4 and Wajak 1 Eastern Java, Indonesia present different profiles for their respective predicted assignations Fig.

The probabilities for Wajak 1 are largely distributed among the Melanesian, Chinese, Micronesian, and Polynesian groups, with higher probabilities for the Polynesian group This points to shared affinities for the craniofacial shape variables of Wajak 1 with the Australian and Melanesian pole as well as with the Chinese, Micronesian, and Polynesian pole. The two non-Teouma archaeological comparison specimens from the Pacific present two opposite profiles for their predicted assignations Fig.

The Terminal Lapita or immediately post-Lapita individual Y from Waya Yasawa, Fiji presents dominant affinities with recent Micronesians and Chinese respectively The five Lapita specimens from the Teouma cemetery show craniofacial shape affinities with the Chinese, Western Micronesian, and Polynesian geographical pole, with only minor or null percentages of predicted assignations to recent Australian and Melanesian groups Fig. However, the detailed results of individual predicted assignations distinguish two groups.

Teouma B10A, B10B, and B30A present largely dominant percentages of predicted assignations to the recent Polynesian group respectively These contrasted craniofacial shape affinities at a finer scale among the Teouma specimens are also found in the scatter plot on LD1 vs. LD2 Fig. The five Teouma individuals, as well as the Terminal Lapita or immediately post-Lapita specimen from Waya Y , are clearly aligned with the recent specimens on the Asian pole toward positive values on LD1, but they tend to be distinguished into two subgroups along LD2.

Our analyses, involving five ca. The results show little or no affinities between the Teouma cranial series and the modern Australian and Melanesian cranial series but strong similarities to Chinese, Western Micronesian, and especially Polynesian cranial series; in other words, the Teouma individuals fit into the Asian pole and not into the Australo-Melanesian pole 5 , 6. Our results also confirm the East Asian affinity previously identified in the Terminal Lapita or immediately post-Lapita individual Y from Waya Yasawa in the Fiji group 19 and the resemblances with modern Melanesians already observed for the post-Lapita specimen recovered at Mangaliliu Efate, Vanuatu Our results also highlight a lack of morphological resemblance between the individuals uncovered at Teouma and the late Pleistocene—early Holocene individuals from Australia and Island Southeast Asia, which in turn display similarities with modern-day inhabitants of Australia and northern Melanesia.

The only notable exceptions to this pattern are Coobool Creek 82, whose proximity with recent Polynesians could be a side effect of artificial cranial modification by head binding 34 , 35 , and the Pleistocene Javanese specimen Wajak 1 recently redated to 37,—28, y B. This individual has been described in previous studies as a very large specimen with some resemblances to early anatomically modern humans

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Author contributions: Physical resemblance between Polynesians and Asian populations was detected, but in between were the islands of Melanesia, inhabited largely by people of different phenotype. However, the Lapita culture bridged this geographical divide 3, y ago. Morphological studies of early Lapita colonists from Teouma, Vanuatu align them with present-day Polynesian and Asian populations, whereas skeletal remains of later generations show a more Melanesian phenotype predominating.

Geneticists have begun using old bones to make sweeping claims about the distant past. But their revisions to the human story are making some scholars of prehistory uneasy.

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The many islands of Vanuatu share an ancient Pacific tribal culture dating back more than years. The more recent past of the islands formerly known as New Hebrides named by one James Cook in , has been one of explorers, blackbirders, missionaries, colonial settlement and, finally, independence. The history of Vanuatu begins with the Lapita people. They also made long canoe voyages, leaving their lovely pottery at sites from northeast Papua New Guinea to Samoa. There is evidence of their occupation of Vanuatu in many places. Pottery found on Malo, off Santo, showed they settled there about BC.

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