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

Frederique Valentin, Florent Detroit, Matthew Spriggs, Stuart Bedford

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Frederique Valentin, Florent Detroit, Matthew Spriggs, Stuart Bedford. Early Lapita skeletons from Vanuatu show Polynesian craniofacial shape: Implications for Remote Oceanic settlement and Lapita origins. Proceedings of the National Academy of Sciences of the United States of America, 2016, 113 (2), pp.292 - 297. 10.1073/pnas.1516186113. halshs-01730862

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

Frédérique Valentin<sup>a,1</sup>, Florent Détroit<sup>b</sup>, Matthew J. T. Spriggs<sup>c</sup>, and Stuart Bedford<sup>d</sup>

Edited by Jim Allen, La Trobe University, Broulee, NSW, Australia, and approved November 24, 2015 (received for review August 14, 2015)

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.

Pacific Islands peopling | Polynesian origin | early Lapita | skull | morphometric analysis

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 3,470–3,250 y B.P. or slightly later (1). 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 3,000 and 2,800 y B.P. (1, 2). In Vanuatu, as in the rest of Remote Oceania, Lapita quickly evolved, within 200–300 y, into distinctive local cultures in conjunction with increased population size and sedentism by the end of the Lapita period (3).

The question of the biological nature of the Lapita populations is routinely approached with data collected from protohistoric/historic or extant populations used as proxies. Analysis of skull morphology and morphometrics of protohistoric/historic populations from Oceania shows a geographical pattern of variation, separating northern and southern Melanesia from western and eastern Polynesia (4–6). 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 (7–13). These populations display haplogroups attributed both to the Pleistocene settlement of the northern Melanesian/Near Oceanic region and to the Lapita diaspora, with chronological estimates based on genetic data. 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.

Studies on Lapita skeletal morphology (Table S1) have identified diversity in the physical makeup along with some level of local homogeneity, similarities with pre-Neolithic specimens from Island Southeast Asia and protohistoric/historic populations from southeastern Melanesia, and features not represented in any later Pacific Island groups (14–20). In a recent biodistance study of mandibles from Watom (New Britain), Pietrusewsky et al. (16) conclude that “expectation that skeletons associated with the Lapita Cultural Complex, Early or Late Lapita, biologically resemble the modern-day inhabitants of Remote Oceania is not supported” and challenge “the prevailing orthodox view that the origin of Polynesians is associated with Lapita culture.” However, whether the few analyzed individuals represent initial “Lapita people” is open to question. Because they postdate the initial appearance of the Lapita culture in the region (20), they may actually reflect subsequent gene flow and migratory events within the Melanesian region, saying more “about the contemporary indigenous inhabitants of eastern Melanesia than . . . about the ancestors of the Polynesians,” as noted by Pietrusewsky et al. (18). Alternatively, the possibility that these late Lapita and (immediately) post-Lapita individuals derive directly from the initial “Lapita population” is not excluded, because heterogeneity among the early populations of the region and among the Lapita groups themselves might be expected (21–23).

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

## Significance

The question of the origin of Pacific Islanders has exercised scientists since the first explorers' voyages of the 16th century. 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 3,000 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.

Author contributions: F.V. and F.D. designed research; F.V. and F.D. performed research; F.V. and F.D. collated the data; F.V. and F.D. analyzed data; and F.V., F.D., M.J.T.S., and S.B. wrote the paper.

This article is a PNAS Direct Submission.

¹To whom correspondence should be addressed. Email: frederique.valentin@mae.u-paris10.fr.

This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1516186113/-/DCSupplemental.

<footnote><sup>a</sup>Maison de l'Archéologie et de l'Ethnologie, CNRS, UMR 7041, 92023 Nanterre, France; <sup>b</sup>Département de Préhistoire, Muséum National d'Histoire Naturelle, Musée de l'Homme, CNRS, UMR7194, 75116 Paris, France; <sup>c</sup>School of Archaeology and Anthropology, College of Arts and Social Sciences, The Australian National University, Canberra, ACT 2601, Australia; and <sup>d</sup>School of Culture, History and Language, College of Asia & the Pacific, The Australian National University, Canberra, ACT 2601, Australia</footnote>
issue. However, published data on prehistoric Pacific Islanders are sparse and mainly centered on Polynesia (24). 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.P. (25)] and four individuals representing the late Lapita (or immediately post-Lapita) human group from Watom (New Britain), all appearing in a separate mtDNA lineage to modern Polynesians (26). Although consistent with the morphological evidence (15, 16, 27), these pioneering results are still regarded as uncertain (24). The lack of preservation of aDNA is a major complication, as attested in early specimens from New Caledonia (WKO013B) and Mussau (Bismarck Archipelago) (18, 28) as well as in the Teouma human samples tested so far.

Here, we provide for the first time to our knowledge insights into the biological diversity and affinities of the human population in Vanuatu at the time of the settlement of Remote Oceania, using craniometric evidence recorded on a sample of the initial “Lapita population” dated to *ca.* 3,000–2,850 y B.P. (29) and comparisons with prehistoric and modern populations from the Asia-Pacific region. Large-scale excavations at the Teouma site have revealed a Lapita cemetery with 68 burial features where adults were preferentially treated by inhumation (30). 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 (31). The extant cranial elements consist of seven skulls in secondary deposits (B10 cache, B17, and B30) (Fig. 1), two partial skulls in a disturbed context (Quarry Area), single cranial fragments associated either with incomplete inhumation or secondary deposits (B10, B12, B29, and B45), six mandibles and a fragment of a seventh (B10, B17, B30, and B29), and 98 associated teeth. Although disconnected from the infra-cranial 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 (32). 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. 1).

## Materials and Methods

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. It comprises Late Pleistocene and Holocene individuals from the Sahul region, pre-Neolithic individuals from Island Southeast Asia (ISEA), and immediately post-Lapita and more recent prehistoric individuals from Remote Oceania (Table S2). The second dataset consists of nine samples selected from the database of Howells (33), representing five geographically distinct human populations from East Asia (China), Western Micronesia (Guam), Island Melanesia (mainly Tolai from New Britain), Australia, and Eastern Polynesia (Table S3). This dataset originally included a total of 707 adult individuals out of which subsamples of 54 specimens (27 male and 27 female) were sampled in each of the five geographical groups (n = 270). 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).

## Results

The scatter plot of specimens on PC1 vs. PC2, which accounts for almost 48% of the total shape variation, shows rather large overlaps between the recent specimens of the five geographical groups (Fig. 2). 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. Contributions of variables indicate PC1 mainly opposes the variables NLH and OBH that are positively correlated with PC1, with the variables NLB and PAC that are negatively correlated with PC1

<figure><img src="image_3.png" /><figcaption>**Fig. 1.** Teouma Lapita skulls TEO 10a, 10b, 10c, 17, and 30a [images of the skull courtesy of Chris Smith (Anatomy Museum Curator, University of Otago) and burial features B17 and B30 (bone collection B30 lies on the lower limbs of without-skull burial B44) [computer-assisted design courtesy of Michèle Ballinger (CNRS, UMR 7041)]].</figcaption></figure>

(Fig. 2). 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 five Teouma specimens, as well as the Fijian Terminal Lapita or immediately Post-Lapita specimen Waya Y2-25-1, plot clearly among the variation range of these three recent “Asian-like” groups on PC1 vs. PC2. 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. Two archaeological Australian specimens from the Cooolool Creek series (CC28 and CC36) plot at the margin of the variation of recent Australians on PC1 vs. PC2; however, neither of these two individuals is described as artificially modified by head binding (34).

The LDA is able to correctly discriminate the five geographical groups of the 270 recent specimens. The scatter plot on LD1 vs. LD2 mainly distinguishes two major geographical groups along LD1 (Fig. 3). 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. LD2 and their 95% ellipses are only slightly discriminated.
<figure><img src="image_4.png" /><figcaption>**Fig. 2.** PCA on the log-shape ratios of 11 craniofacial variables for 270 recent and 17 archaeological specimens: scatter plot of specimens on PC1 vs. PC2, with 95% ellipses for the five geographical groups of recent specimens and plot of the 11 craniofacial variables (log shape ratio) scores for PC1 vs. PC2.</figcaption></figure>

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 270 recent specimens.

The cross-validation procedure indicates that 72.6% of the 270 recent specimens are correctly assigned to their original group (Table 1). This percentage is significantly higher than the 20% probability of a correct assignation by chance only for five geographical groups. The results indicate 77.8% of correct assignments for Australians, 75.9% for Melanesians, 75.9% for Chinese, 70.4% for Polynesians, and 63% for the Western Micronesian group that provides the highest number of incorrectly assigned specimens. If one takes into account the two larger geographical groups that are distinguished in the PCA and the LDA scatter plots (i.e., Australian and Melanesian on the one hand and Chinese, Micronesian and Polynesian on the other hand), percentages of correctly assigned specimens significantly increase to, respectively, 88.9% and 95.7%. This implies that despite a significant percentage of incorrect assignations the discriminant functions computed with the 11 craniofacial shape variables for 270 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.

The results of the predicted classifications of the 17 archaeological specimens computed with the discriminant functions are presented in Figs. 4 and 5. Seven of the eight Australian archaeological individuals present high assignation probabilities to Australian and/or Melanesian groups (Fig. 4A). The eighth Australian archaeological specimen, Cooolool Creek 82, presents a completely different profile of its predicted assignation. The probabilities indicate more than 84% for an assignation to the Polynesian group and negligible probabilities for assignations to the Australian and Melanesian groups. This particular specimen has been systematically identified as artificially modified in previous work (34, 35), which could explain its particular craniofacial shape.

The two archaeological specimens Song Keplek 4 and Wajak 1 (Eastern Java, Indonesia) present different profiles for their respective predicted assignations (Fig. 4B). Song Keplek 4 presents very clear craniofacial shape affinities with the recent Melanesian group, with probabilities of more than 83% for an assignation to this group. The probabilities for Wajak 1 are largely distributed among the Melanesian, Chinese, Micronesian, and Polynesian groups, with higher probabilities for the Polynesian group (53.5%). 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. 4C). The Terminal Lapita or immediately post-Lapita individual Y2-25 from Waya (Yasawa, Fiji) presents dominant affinities with recent Micronesians and Chinese (respectively

68.6% and 29.9%), whereas the post-Lapita specimen from Mangaliliu (Efate, Vanuatu) shows almost exclusive craniofacial shape affinities with recent Australian and Melanesian specimens, with probabilities of predicted assignations to these two groups of, respectively, 71.5% and 27.5%.

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. 5). 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 70.3%, 94.0%, and 88.8%), whereas Teouma B10C and B17 present dominant affinities with the recent Chinese sample (respectively 78.2% and 72%). 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. 3). The five Teouma individuals, as well as the Terminal Lapita or immediately post-Lapita specimen from Waya (Y2-25-1), are clearly aligned with the recent specimens on the Asian pole toward

<figure><img src="image_5.png" /><figcaption>**Fig. 3.** LD1 vs. LD2: scatter plot of the 270 recent specimens (with 95% ellipses for the five geographical groups) used to compute the discriminant functions and 17 archaeological specimens (predicted values).</figcaption></figure>
<table><caption>Table 1. Discriminant analysis: Assignations of specimens to geographical groups after cross-validation</caption><thead><tr><th>Geographical groups</th><th>Australia</th><th>Melanesia</th><th>China</th><th>Micronesia</th><th>Polynesia</th><th>Total (original)</th><th>Percent correct, %</th><th>Pooled geographical groups (original)</th><th>Percent correct (pooled geographical groups), %</th></tr></thead><tbody><tr><td>Australia</td><td>42</td><td>9</td><td>0</td><td>1</td><td>2</td><td>54</td><td>77.8</td><td>108</td><td>88.9</td></tr><tr><td>Melanesia</td><td>4</td><td>41</td><td>2</td><td>4</td><td>3</td><td>54</td><td>75.9</td><td></td><td></td></tr><tr><td>China</td><td>0</td><td>1</td><td>41</td><td>6</td><td>6</td><td>54</td><td>75.9</td><td>162</td><td>95.7</td></tr><tr><td>Micronesia</td><td>0</td><td>3</td><td>11</td><td>34</td><td>6</td><td>54</td><td>63.0</td><td></td><td></td></tr><tr><td>Polynesia</td><td>2</td><td>1</td><td>5</td><td>8</td><td>38</td><td>54</td><td>70.4</td><td></td><td></td></tr><tr><td>Total</td><td>48</td><td>55</td><td>59</td><td>53</td><td>55</td><td>270</td><td>72.6</td><td>270</td><td>92.3</td></tr></tbody></table>

positive values on LD1, but they tend to be distinguished into two subgroups along LD2.

Discussion

**A Lapita-Asian Connection.** Our analyses, involving five *ca.* 3,000- to 2,850-y-old skulls associated with the Lapita culture uncovered from the Teouma site (Efate, Vanuatu) and 270 recent skulls from the five geographic regions of Australia, Melanesia, Western Micronesia, Polynesia, and China clearly point to a Lapita-Asian biological connection. 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 Y2-25 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) (36).

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 37,400-28,500 y B.P. (37). This individual has been described in previous studies as a very large specimen with some resemblances to early anatomically modern humans (38). However, several studies also pointed out its “Asian characteristics,” especially in its facial features (37, 39, 40). According to our analysis, those attributes are shared with the Lapita individuals. Interestingly, Early Holocene (11,000-8,000 y B.P.) and Mid-Late Holocene (4,000/3,000-2,200 y B.P.) individuals from Niah West Mouth Cave (41) have been shown to share similar cranial and facial morphology (42). These observations suggest that the identified Lapita-Asian biological connection, regardless of the

<figure><img src="image_6.png" /><figcaption>Fig. 4. Bar plot of the predicted probabilities of posterior assignments to the five geographical groups: (A) Australian archaeological specimens; (B) Javanese archaeological specimens; (C) Pacific archaeological specimens. A, Australia; ME, Melanesia; C, Chinese; MI, Micronesia; P, Polynesia.</figcaption></figure>
<figure><img src="image_7.png" /><figcaption>**Fig. 5.** Teouma archaeological specimens: bar plot of the predicted probabilities of posterior assignations to the five geographical groups (AUST, Australia; MELA, Melanesia; CHIN, Chinese; MICR, Micronesia; POLY, Polynesia.</figcaption></figure>

time period it was transferred to Remote Oceania, could have an antiquity predating “neolithization” in ISEA and roots in the late Pleistocene/early Holocene populations already established in the greater Southeast Asian region. Even if phenotypic and genetic markers are not necessarily connected, it is worth noting that the observed pattern finds parallels in genetic evidence pointing to the presence of Asian features as early as the Early-Mid Holocene in Island Southeast Asia and Near Oceania and indicating multiple phases of dispersals within this region (11, 43, 44). The population movement associated with the Lapita culture could therefore have been of much shorter distance than usually thought (45).

**The Question of Admixture.** Our results otherwise echo earlier morphometric conclusions formulated by Howells (46), affirming no important gene exchange between Melanesians and pre-Polynesians. Indeed, the observed lack of similarity between the five individuals recovered at Teouma and the modern-day inhabitants of Australia and northern Melanesia suggests little or no admixture between individuals, either male or female, associated with the Lapita culture who colonized Central Vanuatu ca. 3,000 y ago and the population already established/resident in Near Oceania. This interpretation, differing from that of genetic studies that indicate a mixed (varying with sex) Asian/Near Oceanian ancestry of Remote Oceanians (8–10, 12, 47), could be related to the time of admixture, with estimates calculated via genomewide data analyses being imprecise: around 3,000 y ago [ABC simulation analysis (10)] and even 2,700 y ago (wavelet transform analysis, 95% confidence interval: 2,300–3,900 y (48)]. Conflicting with what is implied in the Triple I model for Lapita (21, 49), incorporation of Near Oceanian biological features and the spread of Lapita culture could therefore have been time-dissociated processes. Nevertheless, absence of evidence of Near Oceanian admixture in our sample does not rule out previous expectations of heterogeneity among the early populations of the region and among Lapita-associated groups themselves (21–23). In fact, heterogeneity does exist in the Teouma group, a sample representing a single community. Our results point to morphological diversity within the Asian range of variation (Figs. 2, 3, and 5), suggesting a composite group with multifocal origins. Results of isotopic studies, showing diversity in geographical origins of the individuals buried at Teouma (32), support this view as well as long-distance exchanges of obsidian (50) and pots (51) during the earliest phases of site use.

**The Lapita/Polynesian Connection.** Contrasting with previous morphometric studies of Lapita cranial elements (14–16, 18, 19), evidence presented in this study definitively supports the consensual view that the origin of the Polynesians is associated with the Lapita culture (28, 52–54). Our results, illustrating strong similarities between ca. 3,000-

y-old cranial remains from Teouma in Vanuatu and the Polynesian cranial series in this study, demonstrate a common ancestry for at least some of the people associated with the early Lapita cultural complex in Vanuatu and the Polynesian peoples, and a Lapita foundational contribution to the Polynesian gene pool. This common ancestry seems to be shared with early people established in Fiji. The late Lapita individual from Naitabale (Moturiki) is reported to display Polynesian features (20), and the Terminal Lapita or immediately post-Lapita Y2-25 individual from Waya (Yasawa) presents East Asian affinities (19). These data give little support to the ideas of distinctiveness between Fijian and Tongan founding populations and of an emergence of a distinctive Polynesian phenotype in West Polynesia (55). The Polynesian physical makeup seems to predate the development of Ancestral Polynesian society and its spread to East Polynesia (56).

**Secondary Population Movement into Remote Oceania.** In this context, the recurring observations of similarities between the skeletons associated with the late Lapita and immediately post-Lapita periods in Near and Remote Oceania and protohistoric/historic skeletons from island Melanesia reported by previous studies need explanation. Affinities with modern inhabitants of Melanesia are recognized for the skeletons associated with the late Lapita period from Watom Island (New Britain) (15, 16). Similarities with East Polynesian populations, including New Caledonia, are diagnosed in the WKO013B (18) and in WKO013C (17) skeletons, both excavated from the site of Lapita in Koné (New Caledonia) and dated to the immediately post-Lapita period. Polynesian morphological features are identified (27) in skeletons uncovered at the Taplins site in Mele (Efate, Vanuatu) also dated to the immediately post-Lapita period (25). We suggest that impact from a northern Polynesian gene flow, in at least some regions of Remote Oceania such as Vanuatu and New Caledonia, could account for this pattern of morphological variation over time (57, 58). Twenty years ago, Green (59) proposed that this movement started by 2,500 y B.P. We propose here that it occurred earlier, shortly after the initial colonization and before the end of the Lapita period. This suggestion, biologically plausible considering the short duration of the Lapita period in Remote Oceania (200–300 y, 12–15 human generations), is consistent with the results of genetic studies. There is a larger Near Oceanian contribution to the gene pool in populations living today in the western part of Remote Oceania (10, 12) with the possibility of an admixture time at about 2,700 y ago (48).

This interpretation is also supported by cultural evidence. Earlier there were suggestions of a post-Lapita secondary migration from Near Oceania into western parts of Remote Oceania, supposedly associated with the spread of incised and applied relief pottery of the Mangaasi style (54). However, the end of the Lapita period is marked by a major decrease in mobility indicated by cessation of obsidian long-distance exchange (22, 50), by the divergence and ultimate end of production of Lapita pottery itself, and by radical changes in dietary and FUNERARY PRACTICES as observed at the Teouma site (31).

This alternative prospect, considering a modification/alteration of the initial morphology of the population associated with the Lapita culture as occurring during the Lapita period rather than immediately afterward, reconciles the contradictory archaeological and bioarchaeological views on the origins of Polynesians better than other explanations such as selection, adaptation, secular change, bottleneck, and founder effect (60, 61), or the addition of a significant later Asiatic migration via Micronesia (16, 62). The late Lapita individuals that have been the subject of study before the discovery of the Teouma site are simply not representative of the biology of the initial Lapita dispersal through Remote Oceania into Polynesia. In Island Melanesia this original migratory push was overlain by further migration from Near Oceania that did not significantly affect Tonga, Samoa and adjacent regions of Western Polynesia.
1. Sheppard PJ, Chiu S, Walter R (2015) Re-dating Lapita movement into Remote Oceania. *J Pacific Archaeol* 6(1):26-36.

2. Bedford S (2006) *Pieces of the Vanuatu Puzzle: Archaeology of the North, South and Centre, Terra Australis 23* (Pandanus, Canberra, Australia).

3. Bedford S, Clark G (2001) The rise and rise of the incised and applied relief tradition: A review and reassessment. *The Archaeology of Lapita Dispersal in Oceania, Terra Australis 17*, eds Clark G, Anderson A, Sorovi-Vunidilo T (Pandanus, Canberra, Australia), pp 61-74.

4. Hanihara T (1994) Craniofacial continuity and discontinuity of Far Easterners in the Late Pleistocene and Holocene. *J Hum Evol* 27(5):417-441.

5. Pietrusewsky M (2008) The modern inhabitants of Island Southeast Asia: A craniometric perspective. *Recent Advances on Southeast Asian Paleoanthropology and Archaeology*, ed Indriani E (Faculty of Medicine, Gadjah Mada University, Yogyakarta, Java), pp 185-201.

6. Buck T, Vidarsdóttir US (2012) Craniofacial evolution in Polynesia: A geometric morphometric study of population diversity. *Am J Hum Biol* 24(6):776-785.

7. Cox MP, et al. (2007) A Polynesian motif on the Y chromosome: Population structure in remote Oceania. *Hum Biol* 79(5):525-535.

8. Kayser M, et al. (2008) Genome-wide analysis indicates more Asian than Melanesian ancestry of Polynesians. *Am J Hum Genet* 82(1):194-198.

9. Friedlaender JS, et al. (2008) The genetic structure of Pacific Islanders. *PLoS Genet* 4(1):e19.

10. Wollstein A, et al. (2010) Demographic history of Oceania inferred from genome-wide data. *Curr Biol* 20(22):1983-1992.

11. Soares P, et al. (2011) Ancient voyaging and Polynesian origins. *Am J Hum Genet* 88(2):239-247.

12. Delfin F, et al. (2012) Bridging near and remote Oceania: mtDNA and NRY variation in the Solomon Islands. *Mol Biol Evol* 29(2):545-564.

13. Duggan AT, et al. (2014) Maternal history of Oceania from complete mtDNA genomes: Contrasting ancient diversity with recent homogenization due to the Austronesian expansion. *Am J Hum Genet* 94(5):721-733.

14. Kirch PV, Swindler DR, Turner CG, 2nd (1989) Human skeletal and dental remains from Lapita sites (1600-500 B.C.) in the Mussau Islands, Melanesia. *Am J Phys Anthropol* 79(1):63-76.

15. Pietrusewsky M (1989) A study of skeletal and dental remains from Watom Island and comparisons with other Lapita people. *Rec Aust Mus* 41(3):235-292.

16. Pietrusewsky M, Buckley H, Anson D, Toomay Douglas M (2014) Polynesian origins: A biodiversity study of mandibles from the Late Lapita site of Reber-Rakival (SAC), Watom Island, Bismarck Archipelago. *J Pacific Archaeol* 5(1):1-20.

17. Valentin F (2003) Human skeletons recovered in the site of Lapita at Koné (New Caledonia). *Pacific Archaeology: Assessments and Prospects*, Les Cahiers de l'Archéologie en Nouvelle-Calédonie 15, ed Sand C (Département Archéologie, Service des Musées et du Patrimoine de Nouvelle-Calédonie, New Caledonia), pp 285-293.

18. Pietrusewsky M, Galipaud JC, Leach F (1998) A skeleton from the Lapita site at Koné, Foué Peninsula, New Caledonia. *N Z J Archaeol* 18:25-74.

19. Pietrusewsky M, Hunt TL, Ikehara-Quebral RM (1997) A Lapita skeleton from Waya Island, Fiji. *Micronesica* 3(2):355-388.

20. Katayama K, et al. (2007) Osteological description of the Lapita-associated human skeleton discovered on Moturiki Island, Fiji. *People Culture Oceania* 23:73-98.

21. Green RC (2003) The Lapita horizon and traditions—Signature for one set of Oceanic migrations. *Pacific Archaeology: Assessments and Prospects*, Les Cahiers de l'Archéologie en Nouvelle-Calédonie 15, ed Sand C (Département Archéologie, Service des Musées et du Patrimoine de Nouvelle-Calédonie, New Caledonia), pp 95-120.

22. Summerhayes GR (2010) *Lapita Obsidian Sources and Distribution* / *Les Sources et la Répartition de l'obsidienne Lapita, Lapita: Ancêtres Océaniens*, eds Sand C, Bedford S (Somogy Edition/Musée du Quai Branly, Paris), pp 92-101.

23. Clark GR (2012) Responses to Intrusion, integration and innovation on small and not-so-small islands with particular reference to Samoa. *Archaeol Ocean* 47(1):11-12.

24. Matiso-Smith E (2015) Ancient DNA and the human settlement of the Pacific: A review. *J Hum Evol* 79:93-104.

25. Valentin F, Spriggs M, Bedford S, Buckley H (2011) Vanuatu mortuary practices over three millennia: Lapita to the early European contact period. *J Pacific Archaeol* 2(2): 49-65.

26. Hagelberg E, Clegg JB (1993) Genetic polymorphisms in prehistoric Pacific islanders determined by analysis of ancient bone DNA. *Proc Biol Sci* 252(1334):163-170.

27. Ward G, Houghton P (1991) The Mele burials (Vanuatu): Salvage excavations and biological relationships. *Indo-Pacific Prehistory Assoc Bull* 11:229-235.

28. Kirch PV (1997) *The Lapita Peoples: Ancestors of the Oceanic World* (Blackwell, Oxford).

29. Petchey F, Spriggs M, Bedford S, Valentin F, Buckley H (2014) Radiocarbon dating of burials from the Teouma Lapita cemetery, Efate, Vanuatu. *J Archaeol Sci* 50:227-242.

30. Bedford S, Spriggs M, Regenvanu R (2006) The Teouma Lapita site and the early settlement of the Pacific islands. *Antiquity* 80(310):812-828.

31. Valentin F, Herrscher E, Bedford S, Spriggs M, Buckley H (2014) Evidence for social and cultural change in Central Vanuatu between 3000 and 2000 BP: Comparing funerary and dietary patterns of the first and later generations at Teouma, Efate. *J Island Coast Archaeol* 9(3):381-399.

32. Bentley AR, et al. (2007) Lapita migrants in the Pacific's oldest cemetery: Isotopic analysis at Teouma, Vanuatu. *Am Antiq* 72(4):645-656.

33. Howells WW (1996) Howells' craniometric data on the Internet. *Am J Phys Anthropol* 101(3):441-442.

34. Durband AC (2008) Artificial cranial deformation in Pleistocene Australians: The Coobool Creek sample. *J Hum Evol* 54(6):795-813.

35. Howells WW (1989) *Coobool Creek: A Morphological and Metric Analysis of the Crania, Mandibles and Dentitions of a Prehistoric Australian Human Population*, Terra Australis 13 (Pandanus, Canberra, Australia).

36. Valentin F, Shing R, Spriggs M (2005) Des restes humains datés du début de la période de Mangaasi (2400-1800 BP) découverts à Mangalili (Efate, Vanuatu). *C R Palevol* 4:420-427.

37. Storm P, et al. (2013) U-series and radiocarbon analyses of human and faunal remains from Wajak, Indonesia. *J Hum Evol* 64(5):356-365.

38. Stringer CB (1992) Reconstructing recent human evolution. *Philos Trans R Soc Lond B Biol Sci* 337(1280):217-224.

39. Storm P (1995) The evolutionary significance of the Wajak skulls. *Scripta Geologica* 110:1-247.

40. Jacob T (1967) *Some Problems Pertaining to the Racial History of the Indonesian Region* (Drukkerij Nederland, Utrecht, The Netherlands).

41. Barker G, Richards MB (2013) Foraging-farming transitions in Island Southeast Asia. *J Archaeol Method Theory* 20(2):256-280.

42. Krigbaum J, Manser J (2005) The West Mouth burial series from Niah cave: Past and present. *The Perak Man and the Other Prehistoric Skeletons from Malaysia*, ed Majid Z (Penang Centre for archaeological research, Penerbit Universiti Sains Malaysia, Penang, Malaysia), pp. 175-206.

43. Karafet TM, et al. (2010) Major east-west division underlies Y chromosome stratification across Indonesia. *Mol Biol Evol* 27(8):1833-1844.

44. Tumonggor MK, et al. (2013) The Indonesian archipelago: An ancient genetic highway linking Asia and the Pacific. *J Hum Genet* 58(3):165-173.

45. Terrell J (2013) Polynesians and the seductive power of common sense. *Cult Geogr* 20(2):135-152.

46. Howells WW (1979) Physical anthropology. *The Prehistory of Polynesia*, ed Jennings JD (Australian National Univ Press, Canberra, Australia), pp 271-285.

47. Cox MP, Karafet TM, Lansing JS, Sudoyo H, Hammer MF (2010) Autosomal and X-linked single nucleotide polymorphisms reveal a steep Asian-Melanesian ancestry cline in eastern Indonesia and a sex bias in admixture rates. *Proc Biol Sci* 277(1687): 1589-1596.

48. Pugach I, Matveey R, Wollstein A, Kayser M, Stoneking M (2011) Dating the age of admixture via wavelet transform analysis of genome-wide data. *Genome Biol* 12(2):R19.

49. Green RC (1991) The Lapita cultural complex: Current evidence and proposed models. *Bull Indo-Pacific Prehistory Assoc* 11:295-305.

50. Reepmeyer C, Spriggs M, Bedford S, Ambrose W (2011) Provenance and technology of lithic artifacts from the Teouma Lapita site, Vanuatu. *Asian Perspective* 49(1):205-225.

51. Dickinson WR, Bedford S, Spriggs M (2013) Petrography of temper sands in 112 reconstructed Lapita pottery vessels from Teouma (Efate): Archaeological implications and relations to other Vanuatu tempers. *J Pacific Archaeol* 4(2):1-20.

52. Bellwood P (1978) *Man's Conquest of the Pacific: The Prehistory of Southeast Asia and Oceania* (Collins, Auckland, New Zealand).

53. Bellwood P (2005) *First Farmers: The Origins of Agricultural Societies* (Blackwell, Oxford).

54. Spriggs M (1997) *The Island Melanesians* (Blackwell, Oxford).

55. Burley DV (2013) Fijian polygenesis and the Melanesian/Polynesian divide. *Curr Anthropol* 54(4):436-462.

56. Wilmshurst JM, Hunt TL, Lipo CP, Anderson AJ (2011) High-precision radiocarbon dating shows recent and rapid initial human colonization of East Polynesia. *Proc Natl Acad Sci USA* 108(5):1815-1820.

57. Green RC (1963) A suggested revision of the Fiji sequence. *J Polyn Soc* 72:235-253.

58. Clark GR (2009) Post-Lapita ceramic change in Fiji. *The Early Prehistory of Fiji, Terra Australis* 31, eds Clark GR, Anderson AJ (Pandanus, Canberra, Australia), pp 387-418.

59. Green RC (1997) Linguistic, biological and cultural origins of the initial inhabitants of Remote Oceania. *N Z J Archaeol* 17:5-27.

60. Houghton P (1991) Selective influences and morphological variation amongst Pacific "Homo sapiens". *J Hum Evol* 21(1):49-59.

61. Katayama K (1996) Polynesians the hypermorphic Asiatics – A scenario on prehistoric Mongoloid dispersals into Oceania. *Anthropol Sci* 104(1):15-30.

62. Addison DJ, Matiso-Smith E (2010) Rethinking Polynesian origins: A West-Polynesia triple-I model. *Archaeol Ocean* 45(1):1-12.

63. Howells WW (1989) *Skull Shapes and the Map: Craniometric Analyses in the Dispersion of Modern Homo* (Peabody Museum, Cambridge, MA).

64. Pietrusewsky M (2013) Biological connections across the Sea of Japan: A multivariate comparison of ancient and more modern crania from Japan, China, Korea, and Southeast Asia. *Bioarchaeology of East Asia: Movement, Contact, Health*, eds Pechenkina E, Oxenham MF (Univ Press of Florida, Gainesville, FL), pp 143-178.

65. Kovarovic K, Aiello LC, Cardini A, Lockwood CA (2011) Discriminant function analyses in archaeology: Are classification rates too good to be true? *J Archaeol Sci* 38(5): 3006-3018.

66. Darroch JN, Mosimann JE (1985) Canonical and principal components of shape. *Biometrika* 72:241-252.

67. Jungers WL, Falsetti AB, Wall CE (1995) Shape, relative size, and size-adjustments in morphometrics. *Am J Phys Anthropol* 38(52):137-161.

68. Claude J (2008) *Morphometrics with R* (Springer, New York).

69. Petchey F, et al. (2011) Testing the human factor: Radiocarbon dating the first peoples of the South Pacific. *J Archaeol Sci* 38(1):29-44.

70. Bedford S, Buckley H, Valentin F, Tayles N, Longga F (2011) Lapita burials, a new Lapita cemetery and post-Lapita burials from Malakula, northern Vanuatu, South-west Pacific. *J Pacific Archaeol* 2(2):26-48.

71. Simanjuntak T (2004) New insight on the prehistoric chronology of Gunung Sewu, Java, Indonesia. *Modern Quaternary Res Southeast Asia* 18:9-30.