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. 2014 Mar 6;94(3):426-36.
doi: 10.1016/j.ajhg.2014.02.003.

Early Austronesians: into and out of Taiwan

Albert Min-Shan Ko  1 , Chung-Yu Chen  2 , Qiaomei Fu  1 , Frederick Delfin  3 , Mingkun Li  1 , Hung-Lin Chiu  4 , Mark Stoneking  5 , Ying-Chin Ko  6
Affiliations

Early Austronesians: into and out of Taiwan

Albert Min-Shan Ko et al. Am J Hum Genet. .

Abstract

A Taiwan origin for the expansion of the Austronesian languages and their speakers is well supported by linguistic and archaeological evidence. However, human genetic evidence is more controversial. Until now, there had been no ancient skeletal evidence of a potential Austronesian-speaking ancestor prior to the Taiwan Neolithic ~6,000 years ago, and genetic studies have largely ignored the role of genetic diversity within Taiwan as well as the origins of Formosans. We address these issues via analysis of a complete mitochondrial DNA genome sequence of an ~8,000-year-old skeleton from Liang Island (located between China and Taiwan) and 550 mtDNA genome sequences from 8 aboriginal (highland) Formosan and 4 other Taiwanese groups. We show that the Liangdao Man mtDNA sequence is closest to Formosans, provides a link to southern China, and has the most ancestral haplogroup E sequence found among extant Austronesian speakers. Bayesian phylogenetic analysis allows us to reconstruct a history of early Austronesians arriving in Taiwan in the north ~6,000 years ago, spreading rapidly to the south, and leaving Taiwan ~4,000 years ago to spread throughout Island Southeast Asia, Madagascar, and Oceania.

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Figures

Figure 1
Figure 1
The Liangdao Man Skeleton Side and top profile of the ancient specimen found in a supine flexed position without coffin or burial items.
Figure 2
Figure 2
Liangdao Man’s mtDNA Haplogroup There are 29/29 complete matches from rCRS to haplogroup E (based on Phylotree v.15), 2/4 defining positions toward E1 (13254C, 14577C), and 2 additional positions that are not haplogroup defining (315.1C, 16519C). Thus, the Liangdao Man (black star) is haplogroup E (and ancestral to E1).
Figure 3
Figure 3
Haplogroup E among Austronesian Speakers Related to the Liangdao Man Shown are 67 haplotypes obtained from 104 whole mtDNA genome sequences belonging to haplogroup E, collected from this study and published data from the Philippines, Malaysia, Indonesia, and Melanesia, and from two M9a sequences from Han in southern China. The Liangdao Man sequence (black star) is an intermediate between E and E1. The Formosan sequences are most similar to the Liangdao Man sequence, where the closest sequences (red stars) only differ by four nucleotide changes, and these are found in two Ami (with differences at np 4248-6340-6620-10834) individuals under E1a and one Atayal (with differences at np 4248-6620-10834-14766) individual under E1a1. The polymorphic positions are indicated on branches; recurrent mutations are underlined.
Figure 4
Figure 4
Into and Out of Taiwan (A) Map location of highland Formosan tribes and DAPC plot; individuals are dots, groups are circles, color corresponds to eight Formosan groups and two Han groups in gray. (B) Stepping-stone haplotype sharing across 12 groups. Han/lowland Taiwanese groups are in gray and the frequency of haplotypes shared with/with other groups are colored according to the map. (C) Formosan nucleotide diversity declines from north (N) to south (S). (D) Formosan tribal relationships based on complete mtDNA sequences; dotted lines are non-Formosan groups.
Figure 5
Figure 5
Suggested Migration Route for Early Austronesians Into and Out of Taiwan and the Worldwide Distribution of Haplogroup E1 (A) Geographic regions in China of foxtail millet domestication (shaded) delimited by Nanzhuangtou, Cishan, and Yuezhuang, and of rice domestication (shaded) in the Yangtze River Valley. Shown are (1) early Austronesians in the Fuzhou region, (2) entry into north Taiwan, and (3) rapid north-south dispersal along the west coast and crop cultivation at Nanguanli. (4) One Austronesian language subgroup from Taiwan is ancestral to the Proto-Malayo-Polynesian language subgroup in the Philippines. (B) Frequency of haplogroup E1 across Taiwan, Island Southeast Asia, Madagascar, and Near Oceania.
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