By JOHN NOBLE WILFORD
January 18, 2008, NY Times
The ancestral relationships of people living in the widely scattered islands of the Pacific Ocean, long a puzzle to anthropologists, may have been solved by a new genetic study, researchers reported Thursday.
In an analysis of the DNA of 1,000 individuals from 41 Pacific populations, an international team of scientists found strong evidence showing that Polynesians and Micronesians in the central and eastern islands had almost no genetic relationship to Melanesians, in the western islands like Papua New Guinea and the Bismarck and Solomons archipelagos.
The researchers also concluded that the genetic data showed that the Polynesians and Micronesians were most closely related to Taiwan Aborigines and East Asians. They said this supported the view that these migrating seafarers originated in Taiwan and coastal China at least 3,500 years ago.
The findings were described in the online journal Public Library of Science Genetics (www.plosgenetics.org) by researchers led by Jonathan S. Friedlaender, professor emeritus of biological anthropology at Temple University. He was assisted in the data analysis by his wife, Françoise R. Friedlaender, an independent researcher. Other participants included scientists in the islands and at the Marshfield Clinic Research Foundation in Marshfield, Wis.
“Our analysis,” the scientists wrote, “indicates the ancestors of Polynesians moved through Melanesia relatively rapidly and only intermixed to a very modest degree with the indigenous populations there.”
Dr. Friedlaender of Temple said in an interview that the evidence was “substantial” and “solves a number of issues about the migration and settlement of Pacific people.”
In particular, he and other anthropologists not involved in the study said, the genetic research supported the “fast train” hypothesis. Increasing archaeological and linguistic evidence in recent years has suggested that ancestors of Micronesians and Polynesians had moved through Indonesia and Melanesia without having any significant contact there, culturally or genetically.
An alternative argument, the “slow boat” hypothesis, which had some support from male Y chromosome studies, raised the possibility that Polynesians were primarily Melanesians who had ventured on in their outrigger canoes. And a few anthropologists despaired of ever solving the mystery. Theirs was the “entangled bank” hypothesis.
The new genetic research, said Patrick V. Kirch, an anthropologist at the University of California, Berkeley, who is an authority on Pacific cultures, was “overwhelming biological evidence for a clear population movement out of Southeast Asia and Taiwan to Polynesia.”
Dr. Kirch, who did not participate in the genetic study, said that it reinforced research showing that Polynesian speech patterns were unrelated to Melanesian languages, suggesting — along with discoveries of the distinctive Lapita pottery across the Pacific — links to Taiwan and China, not Melanesia. “The combination of evidence shows we really can read this history,” he said.
As Dr. Friedlaender said, “If it wasn’t exactly an express train, it was pretty fast, and very few passengers climbed aboard or got off along the way.”
In the research, scientists examined more than 800 genetic markers known to be useful in distinguishing the ancestry of people. These involved mitochondrial DNA, passed down through females, and the Y chromosomes in males. Previous investigations along these lines had been conducted on a much smaller scale, Dr. Friedlaender said.
The new test results were repeatedly analyzed with a software program recently developed to classify genetic similarities and variations among different populations.
Primary support for the study was provided by the National Science Foundation, the Wenner-Gren Foundation of Anthropological Research, the National Geographic Society and the National Institutes of Health.
Further research to confirm the history of the Pacific diaspora, Dr. Friedlaender said, would require an expansion of genetic tests among people in the Philippines and Indonesia, regions that the migrants presumably passed through after leaving Taiwan more than 3,500 years ago, ultimately reaching as far as Hawaii and Easter Island. The Melanesians, on the other hand, probably arrived on their islands about 35,000 years ago, sometime later than the Aborigines reached Australia.
Years ago, a reporter who visited the Marshall Islands asked an aging Micronesian chief where his people came from long, long ago. “We have always been here,” he replied. Now, if it matters to them, his descendants have been given a more scientific answer.
“Genome Scans Show Polynesians Have Little Genetic Relationship to Melanesians”, Press Release, Temple University, 18 January 2008, accessed 9 March 2013
Melanesian and Asian Origins of Polynesians:mtDNA and Y Gradients across the Pacific
Manfred Kayser et al.
The human settlement of the Pacific Islands represents one of the most recent major migration events of mankind. Polynesians originated in Asia according to linguistic evidence or in Melanesia according to archaeological evidence. To shed light on the genetic origins of Polynesians, we investigated over 400 Polynesians from 8 island groups, in comparison with over 900 individuals from potential parental populations of Melanesia, Southeast and East Asia, and Australia, by means of Y chromosome (NRY) and mitochondrial DNA (mtDNA) markers. Overall, we classified 94.1% of Polynesian Y chromosomes and 99.8% of Polynesian mtDNAs as of either Melanesian (NRY-DNA: 65.8%, mtDNA: 6%) or Asian (NRY-DNA: 28.3%, mtDNA: 93.8%) origin, suggesting a dual genetic origin of Polynesians in agreement with the “Slow Boat” hypothesis. Our data suggest a pronounced admixture bias in Polynesians toward more Melanesian men than women, perhaps as a result of matrilocal residence in the ancestral Polynesian society. Although dating methods are consistent with somewhat similar entries of NRY/mtDNA haplogroups into Polynesia, haplotype sharing suggests an earlier appearance of Melanesian haplogroups than those from Asia. Surprisingly, we identified gradients in the frequency distribution of some NRY/mtDNA haplogroups across Polynesia and a gradual west-to-east decrease of overall NRY/mtDNA diversity, not only providing evidence for a west-to-east direction of Polynesian settlements but also suggesting that Pacific voyaging was regular rather than haphazard. We also demonstrate that Fiji played a pivotal role in the history of Polynesia: humans probably first migrated to Fiji, and subsequent settlement of Polynesia probably came from Fiji.
polynesia Y chromosome mtDNA genetic origins human population history
The colonization of Polynesia which ranges from Hawaii in the north to Easter Islands in the east, Fiji in the west, and New Zealand in the south, is still a matter of debate. According to linguistic evidence, Polynesian languages are closely related to each other and belong to the Austronesian language family that can be traced back to East Asia, in particular to the present-day languages of Taiwanese Aborigines (Blust 1999; Diamond 2000). Furthermore, linguistic evidence (Gray and Jordan 2000) is usually interpreted to support the “Express-train” hypothesis (Diamond 1988), according to which Polynesian ancestors moved rapidly from Eastern Asia into the Pacific without significant admixture with Melanesians (we use the term “Melanesia” in the geographic sense, to include here the mainland of New Guinea and surrounding islands, also referred to as Near Oceania).
Archaeological evidence suggests that western Polynesian islands (Fiji, Futuna, Samoa, Tonga) were settled 2,100–3,200 years ago by people belonging to the so-called Lapita cultural complex that originated 3,000–3,500 years ago in Island Melanesia, in particular the Bismarck Archipelago (Kirch 2000). However, some archaeologists argue that the Lapita cultural complex originated about 6,000 years ago in China and thus associate the spread of Austronesian languages with the Neolithic spread of material culture, including agriculture and Lapita, from East Asia into the Pacific under the Express-train scenario (Bellwood 1978; Diamond and Bellwood 2003), whereas others suggest a strict Melanesian origin of the Lapita cultural complex (White et al. 1988; Terrell 1989; Terrell et al. 2001). Besides the 2 “extreme” models, the “Express train” assuming an Asian origin of Polynesians with no or little admixture of ingenious Melanesians and the “Entangled bank” assuming a long and complex history of human interactions starting from the first occupation of Melanesia in the Pleistocene (Terrell 1988), there are additional “intermediate” models such as the “Triple I” (Green 1991). The Triple I model assumes that components of the Lapita cultural complex are results of Intrusions of nonindigenous Asian components together with the Integration of indigenous Melanesian elements and new Innovations (Green 1991).
In contrast to the clear evidence for an Asian origin of Polynesian languages and a probable Melanesian origin of the Lapita material culture found in Polynesia, the genetic origin of Polynesians is still contentious. Studies of maternally inherited mtDNA markers have favored an Asian origin of Polynesian maternal lineages (Melton et al. 1995; Redd et al. 1995; Sykes et al. 1995; Trejaut et al. 2005) in support of the Express-train hypothesis. In contrast, studies of paternally inherited DNA markers from the nonrecombining portion of the Y chromosome (NRY) have revealed a mostly Melanesian origin of Polynesian paternal lineages (Kayser, Brauer et al. 2000; Capelli et al. 2001; Underhill, Passarino, Lin, Marzuki et al. 2001; Hurles et al. 2002) supporting the “Slow Boat” hypothesis (Kayser, Brauer et al. 2000). The Slow Boat model assumes that Polynesian ancestors originated in Eastern Asia but mixed extensively with indigenous Melanesians before colonizing the Pacific (Kayser, Brauer et al. 2000). Unfortunately, a similar term “Slow boat to Melanesia” was subsequently used to suggest a Southeast Asian genetic origin of Polynesians in the Pleistocene based on mitochondrial DNA (mtDNA) evidence (Diamond 2001; Oppenheimer and Richards 2001). Studies of autosomal DNA markers suggest different scenarios depending on the markers used, for example, a Melanesian origin of Polynesian hemoglobin genes (Hill et al. 1985, 1987) versus an Asian origin of Polynesian human leucocyte antigen (HLA) genes (Mack et al. 2000; Mack and Erlich 2005).
In this study, we have used NRY and mtDNA markers to investigate the paternal and maternal genetic origin of over 400 individuals from 8 different Polynesian island groups by comparing them with over 900 individuals from Melanesia, Southeast and East Asia, and Australia. This significant increase over previous studies, both in populations and markers analyzed, provides new insights into the history of the human colonization of the Pacific.
The mtDNA sequence data together with data from the 9-bp deletion allowed us to infer 31 mtDNA haplogroups, of which 10 are found in Polynesia, 12 in Melanesia, and 26 in Asia (Table S4, Supplementary Material online; fig. 1). Five Polynesian mtDNA haplogroups have an Asian origin: B4, B4a, B4b1, Polynesian motif (PM), and M7c1c (Kivisild et al. 2002) resulting in an overall estimate of 93.8% of Polynesian mtDNAs being of Asian origin, of which 77.6% accounted for by a single haplogroup (the PM). In addition, 4 Polynesian mtDNA haplogroups have a probable Melanesian origin: P1, Q1, Q2 (Friedlaender et al. 2005), and M28 (Merriwether et al. 2005), resulting in an overall estimate of 6% of Polynesian mtDNAs with a Melanesian origin. Altogether, we could classify all but one of the Polynesian mtDNAs analyzed as either Asian (93.8%) or Melanesian (6%) in origin (Table S4, Supplementary Material online; fig. 1); the remaining individual belonged to haplogroup T, which likely represents recent European admixture (Macaulay et al. 1999).
Dual Genetic Origins of Polynesians
Based on the NRY and mtDNA data, we identified a dual genetic heritage of Polynesians, containing both Melanesian and Asian genetic components. However, these 2 components differed between the paternally inherited Y chromosome and the maternally inherited mtDNA (table 2). Overall in Polynesia, the proportion of Melanesian haplogroups was 11-fold higher for Y chromosomes (65.8%) than for mtDNAs (6%), and of Asian haplogroups was more than 3-fold higher for mtDNAs (93.8%) than for Y chromosomes (28.3%). The proportions of Asian NRY and mtDNA haplogroups in Polynesia were not correlated (Spearman R = 0.43, P = 0.34, excluding Tokelau due to small sample size), and the correlation for Melanesian haplogroups was somewhat higher but not statistically significant (R = 0.60, P = 0.21, excluding Niue and Tokelau for small sample size). In addition, no correlation between all NRY haplogroups and mtDNA sequence for Polynesian populations was observed (Mantel test based on FST: R = 0.243, P = 0.25, excluding Niue and Tokelau). The discrepancy between the amount of Asian versus Melanesian NRY and mtDNA haplogroups of Polynesians could reflect uxorilocal (matrilocal) residence in ancestral Polynesian society (Hage 1998; Hage and Marck 2003), as this would have resulted in more admixture of Asian migrants with Melanesian males than females before their colonization of the Pacific. This explanation finds some support in the proportions of Melanesian and Asian haplogroups in the coastal and island Melanesians included in this study (table 2); those Island Melanesians received a larger contribution of Asian mtDNAs (29.4–72.5%) than of Asian Y chromosomes (5.3–37.7%) from the ancestral Polynesians, as expected given that the respective societies (Tolai, Trobriand Islanders, Bereina-Mekeo) are of virilocal (patrilocal) residence.
Moreover, inferred European genetic components were 15-fold higher for the Y chromosome (4.5%) than for mtDNA (0.3%), in keeping with the expectation that European men would have contributed more genes to Polynesians than European women. The fact that we find lower levels of inferred European ancestry in Polynesia than reported in other studies (Hurles et al. 1998; Capelli et al. 2001) may reflect the care taken to exclude individuals whose genealogical history indicated European ancestry. In our study, we only used Polynesian individuals with Polynesian family history in both parental lines (Trent et al. 1986). We also found no evidence for a genetic contribution from the New World as proposed first by Heyerdahl (1950) and identified previously by Y chromosome analysis but interpreted as a recent genetic contribution (Hurles et al. 2003): the most frequent NRY haplogroups in Native Americans are subgroups of haplogroup P-M74/M45(xR-M173) (Lell et al. 2002), which was not observed in our Polynesian samples.
Genetic Heterogeneity among Polynesian Populations
We analyzed the frequency distributions of the Asian and Melanesian mtDNA and NRY haplogroups in Polynesia (table 2), to ascertain if there is significant genetic heterogeneity among Polynesian groups. With respect to mtDNA, haplogroups PM and B4a vary significantly in frequency (χ2 exact test: P = 0.0017 and P = 0.03, respectively, based on 100,000 Monte Carlo simulations). Sample sizes for the remaining mtDNA haplogroups are too small to test for significant differences in frequency (Table S4, Supplementary Material online), although the frequency of Melanesian mtDNA haplogroups is higher in Fiji (20.5%) than elsewhere in Polynesia (0–7.7%) (table 2 and fig. 1). With respect to NRY haplogroups, O-M122, K-M9, M-M104, and C-M208 showed highly significant frequency differences among Polynesian groups (P < 0.0001, M-M104: P = 0.00019); the other haplogroups either occurred sporadically or only in single populations. Thus, all haplogroups for which sample sizes are sufficient exhibit significant frequency differences among Polynesian groups. This most likely reflects founder events during the colonization of the various islands and/or subsequent genetic drift due to small population sizes. However, many of these haplogroups also show gradients in frequency across the Pacific (fig. 4), with the frequency of one haplogroup (C-M208) significantly and positively correlated with longitude (Spearman's R = 1, P < 0.01), and the correlations for 2 other haplogroups (K-M9 and PM) approaching statistical significance (R = −0.77, P = 0.07 and R = 0.68, P = 0.09, respectively). Such frequency gradients are not expected if founder events occurred at random across the Pacific but instead suggest that there was an increasing tendency for founder events as the more eastern islands were colonized. This interpretation receives further support from the previous observation of an inverse correlation between mtDNA and Y-DNA diversity and the time of colonization of Pacific islands (Hurles et al. 2003). It is also consistent with a study that found a significant association between migration distance from Southeast Asia and loss of heterozygosity for autosomal microsatellite loci, which included a small number of Polynesian groups (Lum et al. 2002). Thus, Pacific voyaging was regular rather than haphazard.
We also noticed striking differences in genetic diversity between groups from different Polynesian islands and for different measures of diversity (Table S5, Supplementary Material online). Diversity of NRY haplogroups and mtDNA HV1 sequences declines from west-to-east (fig. 4), with negative correlations that are approaching statistical significance (Spearman, NRY: R = −0.77, P = 0.07; mtDNA: R = −0.71, P = 0.07). Thus, our data provide evidence for a west-to-east settlement of Polynesia with additional evidence from the frequency and diversity distribution of the Polynesian DYS385 triplication and the Polynesian haplogroup K-M353 (see above).
Are the different Melanesian and Asian NRY and mtDNA haplogroups in Polynesia today the result of a single wave of migration or multiple migrations? To address this question, we performed network analyses as described previously (Kayser, Brauer et al. 2000), and demographic analyses, for the most frequent Polynesian NRY (C-M208, K-M9, M-M4, O-M122) and mtDNA (PM and B4a) haplogroups using associated Y-STR and mtDNA sequence haplotypes, respectively (figs. 2 and 3). All networks exhibit a consistent pattern with one Polynesian haplotype at high frequency that is shared between all (or almost all) Polynesian groups, and most other Polynesian haplotypes connected via 1 or 2 mutational steps only (figs. 2 and 3; Table S3, Supplementary Material online). This star-like pattern, identified in 4 independent NRY and 2 independent mtDNA haplogroups, indicates a strong founder effect with subsequent population expansion in Polynesia and implies that the number of founding Y-STR and mtDNA haplotypes per haplogroup was low in Polynesia. Evidence for the Polynesian founder effect was also reported previously based on other genetic marker systems (Trent, Mickleson et al. 1988; Flint et al. 1989).
If Polynesian ancestors did migrate to coastal/island Melanesia from Asia, mixed with coastal/island Melanesians (thereby obtaining Melanesian Y chromosomes and mtDNA types and leaving behind “Asian” Y chromosomes and mtDNA types), and then left Melanesia and colonized Polynesia, then the degree of haplotype sharing should be the same for haplogroups of Asian versus Melanesian origin because there was a single “separation” of an ancestral group of Polynesians from ancestral Melanesians. The fact that there is extensive sharing of Asian haplotypes, but not Melanesian haplotypes, between Polynesians and Melanesians today, therefore, could indicate that Melanesian haplotypes were present earlier in Polynesia (perhaps in Fiji), leading to greater divergence between Polynesians and Melanesians for haplogroups of Melanesian origin than for haplogroups of Asian origin. However, there are large gaps in the sampling of coastal/island Melanesians, which would need to be filled in before one could be certain that there is truly a difference in patterns of haplotype sharing between Polynesians and Melanesians for haplogroups of Asian versus Melanesian origin.
Fijian Genetic History
Fiji represents the most western islands of Polynesia, and Fijians share some features of physical and cultural traits with Melanesians (for overview see Frost 1979), whereas the Fijian dialects are closely related to Polynesian languages (Ross et al. 2003). The NRY and mtDNA data also indicate a closer relationship between Fijians and Melanesians than between other Polynesians and Melanesians. This is evidenced by the following: 1) the highest overall frequency of mtDNA haplogroups of Melanesian origin in Polynesia (20.5%) is observed in Fiji—it is also the only Polynesian group where all 4 Melanesian mtDNA haplogroups observed in Polynesia are found (table 2 and fig. 1); 2) Fiji displays the highest diversity of Melanesian NRY haplogroups in Polynesia and shows the second highest frequency of Melanesian haplogroups (78.5%) in Polynesia with all 5 major haplogroups being present (table 2 and fig. 1); 3) in the K-M9 network, most Fijian haplotypes are more closely associated with Melanesian than with Polynesian haplotypes (fig. 2C); 4) Fiji displays the highest frequency of M-M4 (24.3%), which elsewhere only exists in Melanesia (2%), where it most likely originated but today mostly occurs as subgroup M-P34 (28–74%). Thus, M-M4 in Fiji represents an old Melanesian lineage that left Melanesia prior to the M-P34 mutation rising in appreciable frequency. On the other hand, there is also a strong Polynesian association of Fijians: 1) in the C-M208 network, all but one of the Fijian haplotypes are shared with Polynesians (fig. 2A); 2) in the O-M122 network, 2 Fijian haplotypes (5 of 7 men) are shared with other Polynesians (fig. 2B); 3) the DYS385 triplication, for which a Polynesian origin is assumed, was observed in Fiji but not in Melanesia (table 1); 4) some M9 haplotypes are shared with other Polynesians (fig. 2C); 5) the Polynesian haplogroup K-M353 was only observed in Fiji and Futuna (but not in Melanesia) and probably arose in Fiji. Moreover, Fijians appear between the Polynesian cluster and the Coastal/Island Melanesian cluster in the FST-based 2-dimensional MDS plots from mtDNA haplotypes as well as from NRY haplogroups (fig. 5), although the latter MDS plot should be interpreted more carefully as indicated by the relatively high stress value. In addition, Fiji shows the highest overall genetic diversity from all Polynesian groups for both Y chromosome and mtDNA markers.
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Two-dimensional plots from multidimensional scaling analyses based on FST distances from (A) NRY haplogroup frequencies and (B) mtDNA sequence haplotypes. NRY haplogroups R-M173 and F-M89 were omitted due to their assumed European origin; Niue (NRY) and Tokelau (NRY, mtDNA) were omitted due to small sample size. For population abbreviations, see Table S5(Supplementary Material online). Geographic regions are highlighted.
These results indicate the central role of Fiji in further Polynesian migrations; the fact that Fiji has the highest genetic diversity, and that all Polynesian groups have a subset of the diversity in Fiji, indicates that humans probably first migrated to Fiji and that subsequent settlement of Polynesia probably came from Fiji. This is in agreement with archaeological evidence showing that the oldest findings of Lapita pottery in Polynesia are from Fiji (3,200 years ago). Having originated from the Bismark Archipelago in Island Melanesia, Lapita was first introduced to Polynesia in Fiji, and there was a rapid expansion of the Lapita cultural complex from Fiji eastward into other parts of Polynesia (Futuna, Tonga, Samoa) as suggested indirectly by finding younger Lapita dates elsewhere in Polynesia (2,900–2,100 years ago), but also directly, for example, by the presence of Fijian potsherds in Tonga (Kirch 2000).
An alternative explanation is that following initial colonization, Fiji continued to receive migrants and genes from Melanesia and that humans continued to disperse from Fiji to Polynesia. Although there is archaeological evidence to support this view (for summary see Kirch 2000), the genetic results do not suggest substantial ongoing contact between Fiji and Melanesia, as separate expansions of Y haplogroups C-M208 and K-M9 (both of Melanesian origin) in Fiji/Polynesia versus Melanesia are evident in the networks (fig. 2A and B). Ongoing contact between Melanesia and Fiji should result in more sharing of haplotypes between Melanesia and Fiji, which is not observed. Moreover, Y haplogroup M-M4 (of Melanesian origin) has its highest frequency in Fiji and exists in Melanesia mostly as its derived subgroup M-P34; ongoing contact should have brought more M-P34 chromosomes to Fiji. However, the low frequency elsewhere in Polynesia of other Melanesian Y and mtDNA haplogroups existing in Fiji precludes definitive conclusions, and additional sampling between mainland New Guinea and Fiji (e.g., from the Solomon Islands, Vanuatu, and New Caledonia) is needed to further investigate the amount of ongoing genetic contact between Melanesia and Fiji.
Our study provides evidence for a dual genetic origin of Pacific Islanders in Asia and Melanesia. This is in agreement with the Slow Boat hypothesis of Polynesian origins (Kayser, Brauer et al. 2000) according to which Polynesian ancestors originated in Asia, moved eastward, and mixed extensively with local Melanesians before colonizing the Pacific Islands. Although dating methods revealed somewhat similar entries of NRY/mtDNA haplogroups into Polynesia, haplotype sharing suggests that haplogroups of Melanesian origin may have appeared earlier in Polynesia than those of Asian origin, although more extensive sampling in Melanesia is needed to confirm this observation. The striking difference observed here between Asian and Melanesian contributions to the paternal and maternal gene pool of Polynesians suggests an admixture bias toward more Melanesian men, perhaps as result of uxorilocal (matrilocal) residence and matrilineal descent in ancestral Polynesian society (Hage and Marck 2003). The identified east-west gradient in the frequency distribution of some NRY/mtDNA haplogroups suggests an increasing tendency for founder events as the more eastern islands were colonized and also implies that Pacific voyaging was regular rather than haphazard. The gradual west-to-east decrease of overall NRY/mtDNA diversity in addition to the frequency distribution of the Polynesian DYS385 triplication provide genetic evidence for a west-to-east settlement of Polynesia. Fiji has played a pivotal role in the history of Polynesia either by having had received an earlier migration wave from Melanesia or by subsequent intensive contacts with Melanesia. In order to differentiate between these scenarios, additional sampling between mainland New Guinea and Fiji (e.g., from the Solomon Islands, Vanuatu, and New Caledonia) is needed. Based on the data presented here, Polynesians can be regarded as an admixed population (especially Fijians), although it should be pointed out that autosomal data are needed in addition to the Y/mtDNA data presented here for a more comprehensive estimate of Polynesian genetic admixture. Nevertheless, we predict that Polynesians should be of interest for admixture mapping of disease genes. For example, Polynesians have an extraordinarily high frequency of Type 2 diabetes (Zimmet et al. 1990), which may reflect past selection on genes involved in nutrition metabolism for a “thrifty genotype” (Neel 1962). Polynesians thus may prove of interest not only because of their fascinating history and extraordinary accomplishments in colonizing the Pacific but also from what we may learn about complex diseases that affect other populations.
Supplementary Tables S1–S5 and Figures S1 and S2 are available at Molecular Biology and Evolution online (http://www.mbe.oxfordjournals.org/).
We are deeply grateful to all volunteers for contributing cheek swab or blood samples and additionally to the following colleagues for providing DNA samples: N. Saha, A. G. Soemantri, A. S. M. Sofro, K. Bhatia, J. Kuhl, N. Kretchmer, D. Bugawan, E. Hagelberg, S. Ulijaszek, K. Katayama, J. Martinson, B. Budowle, and C. Tyler-Smith. We thank D. Mueller, A. Fiedler, and A. Gross for DNA extractions, as well as D. Kappei for technical assistance in DNA typing. C. Schwarz and B. Hoeffner are acknowledged for DNA sequence analysis. R.B.R. is grateful to the Thomas J. Watson Foundation for financial support. The Max Planck Society is acknowledged for financial support of this study.
Funding to pay the Open Access publication charges for this article was provided by the Max Planck Society.
Arndt von Haeseler, Associate Editor
© 2006 The Authors
This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/2.0/uk/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.
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First published online: August 21, 2006
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