AnthropologicalScience 109(4),257-273, 2001
Population

Genetic

Studies
GroupsII. Serumon Nine AboriginalEthnicof Taiwan
Protein

Systems

Isao Yuasa1, Kazuo Umetsu2, Kazutoshi Ago3, Cheih-Shan Sun4,
I-Hung Pans, Takafumi Ishida6, Naruya Saitou7, and Satoshi Horai8
1Departmentof LegalMedicine,TottoriUniversitySchoolof Medicine,Yonago
2 Departmentof ForensicMedicine,YamagataUniversitySchoolof Medicine,Yamagata
3Departmentof LegalMedicine, Facultyof Medicine,KagoshimaUniversity,Kagoshima
4TaidongHospital, Taidong,Taiwan
5Departmentof Bacteriology,Collegeof Medicine,NationalTaiwanUniversity,Taipei,Taiwan
6Departmentof BiologicalSciences, GraduateSchoolof Science,Universityof Tokyo,Tokyo
7Laboratoryof EvolutionaryGenetics,NationalInstituteof Genetics,Mishima
8Departmentof BiosystemsScience, The GraduateUniversityfor AdvancedStudies,Hayama(ReceivedMay 16, 2001; acceptedAugust31, 2001)
Abstract As part of our comprehensive genetic study of the aboriginal Taiwan
ese(Gaoshan), we have reported the distribution of allele frequencies for 13 redcell enzyme and 4 serum protein polymorphisms in all nine aboriginal groups. In
this study, 10 additional serum protein polymorphisms for AHSG, C3, C6, C7,
GC, HP, ITIH1, PI, TF and ZAG were typed. Common and variant alleles prevail
ingin East Asian populations were found together with several new variants. The
distribution of allele frequencies differed markedly among the nine aboriginal
groups and from that in East Asian populations. The large diversity among theaboriginal groups was confirmed by analysis of the genetic affinity on the basis of
Nei's standard genetic distance and Cavalli-Sforza's chord distance, which were
calculated using the present and previous data from 27 red cell enzyme and serum
protein loci. The analysis based on 15 red cell enzyme and serum protein locirevealed that they, in particular the Puyuma and Ami, had strong affinity with
Filipino among several East Asian populations.
Keywords:
morphisms

aboriginal Taiwanese, Gaoshan, population study, serum protein poly

Introduction
The Gaoshan (Mountain Highlander) are one of the 56 recognized ethnic groups
in China and are an aboriginal population in Taiwan. They consist of nine linguisticallyCorresponding
author:IsaoYuasa
Departmentof LegalMedicine,
TottoriUniversity
Schoolof Medicine
86Nishimachi,
Yonago,Tottori683-8503,Japan
E-mail:yuasai@grape.med.tottori-u.ac.jp

258

I. Yuasaet al.

and culturally different groups: the Atayal, Saisiat, Bunun, Tsou, Ami, Puyuma,
Rukai and Paiwan who live on the main island of Taiwan, and Yami who live on
Lanyu Island, located off the southeast coast of the main Island (Du and Yip, 1993;
Jin et al., 1999). There have been few comprehensive population genetic studies
done on all the nine aboriginal groups in Taiwan at the same time. Our project wasundertaken to collect blood samples from all nine groups, to analyze genetic markers
including red cell antigens and enzymes, serum proteins, mtDNA and Y chromosome,and to understand the relationships among these groups and the genetic
affinity with other populations in East Asia (see Jin et al., 1999). As a part of thisproject, we preliminarily reported the data on a red cell enzyme, adenosine deaminase(Jin et al., 1995) and on four serum protein polymorphisms including the R
subcomponent of the first component of the complement (C1R), complement factor
I (IF) and orosomucoid (ORM1 and ORM2) (Umetsu et al., 1994, 1995). Recently
we have begun a new series of comprehensive study and have reported the data on 12
red cell enzyme polymorphisms (Jin et al., 1999). Also, another group investigatedthe HLA system in all nine aboriginal Taiwanese groups (Lin et al., 2000).
Concerning serum protein polymorphisms in the aboriginal Taiwanese, only sev
eralpolymorphisms
including haptoglobin (HP), group-specific component (GC,vitamin D-binding protein), and gamma and kappa chains of immunoglobulin (GMand KM) have been studied in limited numbers of ethnic groups among the aborigi
nalTaiwanese (Fong, 1974; Nakajima and Ohkura, 1971; Matsumoto, 1988). The
Atayal group living in Toroko were studied in detail for six serum protein polymor
phismsincluding the third component of the complement (C3), complement factor B(BF), GC, HP, transcobalamin II (TC2) and Transferrin (TF) as well as 13 otherblood groups and red cell enzyme polymorphisms (Chen et al., 1985). Recently, newpolymorphisms and microheterogeneity of some serum proteins have been demon
stratedby the application of isoelectric focusing. These genetic markers are usefulfor population studies. Subsequently to the first report of this new series (Jin et al.,1999), this paper presents the typing results of ten additional loci for serum proteinsin all nine aboriginal groups of Taiwan.

Materials and Methods
A total of 659 blood samples (Umetsu et al., 1994, 1995; Jin et al., 1995, 1999)
were examined for ten genetic markers including alpha-2-HS-glycoprotein (AHSG),
C3, the sixth and seventh components of the complement (C6, C7), GC, HP, interalpha-trypsin-inhibitor H 1 chain (ITIH1), alpha-1-antitrypsin (PI), TF and Zn-al
pha-2-glycoprotein(ZAG).
C3 and HP were typed using agarose and polyacrylamide gel electrophoresis,
respectively. The others were determined by isoelectric focusing on thin-layer polyacrylamide

SerumProteinPolymorphismsin AboriginalTaiwanesePopulations

259

gels. Allele frequencies were calculated by the standard allele counting
method. Nei's standard genetic distance (Ds) (Nei, 1987) and Cavalli-Sforza's chord
distance (Dc) (Cavalli-Sforza and Edwards,1967) were calculated by PHYLIP 3.5c
(Felsenstein, 1993). Phylogenetic trees were constructed using the unweighted pairgroupmethod with arithmetic mean (UPGMA) and neighbor joining method (NJ)
(Sokal and Sneath, 1967; Saitou and Nei, 1987) with the help of PHYLIP 3.5c
(Felsenstein,1993) and Tree View (Page, 1996).

Results
Distribution of allele frequencies
The distribution of allele frequencies for the ten genetic markers examined isshown in Table 1.
AHSG*1 was predominant over AHSG*2 in the nine groups like all the Asian
populations investigated. The allele frequency for AHSG*2 ranged from 34% in the
Saisiat to 2% in the Atayal. A combined frequency for AHSG*2 in the nine groups(13%) was low in comparison with other East Asian populations. In the Honshu
island and Okinawa of Japan AHSG*2 was observed at the frequency of 27% and23%, respectively (Yuasa and Umetsu, 1988). The aboriginal Taiwanese had no
AHSG*5, which was observed at polymorphic frequencies in Okinawa, a neighbor
ingpopulation located close to the north coast of Taiwan (Yuasa and Umetsu, 1988).
The data in the Ami were not in agreement with Hardy-Weinberg's law. The reason
for discrepancy is not known.
The C3 system is generally monomorphic due to the occupation of C3*S in East
Asian populations studied (Roychoudhury and Nei, 1988; Nishimukai et al., 1985).
The Atayal in Toroko was reported to have only C3*S (Chen et al., 1985). Similarly,
six of the nine aboriginal groups including the Saisiat, Ami, Puyuma, Rukai, Paiwan
and Yami were monomorphic. The three other groups, the Tsuo, Bunun and Atayal,
had C3*F07tsouor C3*F06atayal,which subtly differed in mobility from the C3*F07
and C3*F06 found in Europeans, respectively (Nishimukai, personal commun.). It isnoteworthy that C3*F07tsouwas observed at an allele frequency of as high as 10% in
the Tsou. Also, one Bunun individual possessed this variant.
Concerning the C6 system, C6*A, C6*B and C6*B2 were common alleles in East
Asian populations including Japanese and Han Chinese, where C6*A was observed
at a high frequency of more than 40%, but was somewhat lower in frequency thanC6*B (Roychoudhury and Nei, 1988). All aboriginal groups except the Bunun had
the three common alleles. The frequencies varied among the nine groups, where
C6*A frequency changed from 58% in the Atayal to 18% in the Tsou. A variant
allele, C6*Ataiwanese,with a slightly lower isoelectric point than C6*A, was observedin the Rukai and Paiwan at polymorphic frequencies of 3.7% and 4.2%, respectively.

260

Table

I. Yuasa et al.

1.

Allele frequencies

of ten loci for serum

proteins

in nine Aboriginal

Taiwanese

populations

In the C7 system, the Atayal, Ami, Rukai and Yami had one predominant allele,
C7*1 and three minor alleles, C7*2, C7*3 (=C7*5) and C7*4 as seen in other East
Asian populations including Japanese and Han Chinese (Roychoudhury and Nei,
1988). However, the Saisiat, Tsou, Bunun and Paiwan were devoid of C7*3 and the
Puyuma were lacking in C7*4.

SerumProteinPolymorphismsin AboriginalTaiwanesePopulations

261

The GC system is determined by three common alleles, GC*1F, GC*1S and GC*2
and many rare variant alleles. The distribution of allele frequencies has been studied
in detail all over the world, as reviewed by Kamboh and Ferrell (1986). The Atayal in
Toroko had a GC*2 frequency of 17% (Chen et al., 1985). Similarly, the aboriginal
Taiwanese were shown to have relatively low GC*2 frequencies (23% in the Rukai to
6% in the Tsou). The frequencies of GC*1F were higher than those in the Japanese,
Korean and Han Chinese (Kamboh and Ferrell, 1986). No variants alleles such as
GC*1A2, GC*1A3, GC*1A8, GC*1A9 and GC*1C2, characteristic of East Asianand native American populations, were observed in the aboriginal Taiwanese (Constans
et al., 1985). A new variant allele, GC*1A8saisiat,located closely but very slightly
anodally to GC*1A8, was observed in the Saisiat at a polymorphic frequency of
4.7%. Two other rare variants, GC*1Cami and GC*2Cbunun,were found in the Ami and
Bunun groups, respectively.
The HP system was controlled by two common alleles, HP*1 and HP*2. The
frequency of HP*1 in the nine aboriginal groups varied from 67% in the Yami to 23%
in the Saisiat. The frequency in the Atayal group in this study (30%) was similar tothat in the Atayal from Toroko (26%) (Chen et al., 1985), which was comparable tothe data in most East Asian populations (Roychoudhury and Nei, 1988).
At the ITIH1 locus, three common alleles, ITIH1*1, ITIH1*2 and ITIH1*3 havebeen observed together with a few variant alleles in several East Asian populations
examined so far. ITIH1*1 and ITIH1*2 were observed at frequency of about 50%.
However, ITIH1*1 was predominant over the other alleles in Korean, Han Chinese,and Thai. In contrast, Japanese had higher ITIH1*2 frequencies (Vogt et al., 1991;
Yuasa et al., 1991; Hou et al., 1993; Harada et al., 1994). In the aboriginal Taiwan
ese,only ITIH1*1 and ITIH1*2 were observed without variant alleles. The ITIH1*1
was predominant and its frequencies varied from 89% in the Atayal to 55% in the
Tsou.
The PI system is determined by three major alleles, PI*M1, PI*M2 and PI*M3 in
East Asians, as summarized by Kamboh (1985). The aboriginal Taiwanese had fairly
high PI*M1 frequencies, which were comparable to those in Japanese, Korean, Han
Chinese and Thai (Kamboh, 1985). PI*M3 was exceptionally predominant overPI*M2 in the Rukai. The Yami were lacking in PI*M3. No variants were detected inany aboriginal groups, while some variant alleles have been observed in East Asian
populations: PI*Etokyowas widespread in Japanese and Northern Han Chinese (Yuasa
et al., 1984; Ying et al.,1985).
In the TF system, TF*C1 and TF*C2 were found, and TF*C1 was predominant
over TF*C2 in East Asian populations, as reviewed by Kamboh and Ferrell (1987).
Similarly, TF*C1 was a major allele in the aboriginal Taiwanese. TF*Dchina,characteristicof East Asian and native American populations, was also observed in the
Puyuma and Paiwari. It was reported that the Atayal in Toroko had no TF*Dchina

262

Table

I. Yuasa et al.

2.

Genetic

distances

Figuresabove

averageheterozygosity.

Allele frequency

and below

Ds and Dc among

the diagonalnine Aboriginal

Taiwaneseare Ds and Dc, respectively,

data for the 13 red cell enzyme

and 14 serum protein

populations

and thoseon the diagonal

areloci were used.

(Chen et al., 1985).
The ZAG is known to be almost monomorphic in Japanese, Korean, Han Chinese,
Thai and Filipino except for the Papuan. Several rare variants, however, were ob
servedin some East Asian populations (Nakayashiki et al., 1992). No aboriginal
groups showed any variation at the ZAG locus.
Genetic distance analysis among nine aboriginal Taiwanese
Average heterozygosities (gene diversities) and genetic distances (Ds and Dc)
were calculated using a total of 27 systems including 14 serum protein systems
investigated in the present and previous studies (Umetsu et al., 1994, 1995) and 13
red cell enzyme systems reported previously (Jin et al., 1995, 1999). The average
heterozygosities are shown on the diagonal in Table 2. The values ranged from 0.215
in the Atayal to 0.259 in the Paiwan. All the nine aboriginal groups except for the
Paiwan showed rather small within-population variations in comparison with German(0.254) and Japanese (0.274).
The genetic distances Ds and Dc between each pair of the nine aboriginal Taiwan
esegroups calculated using the allele frequencies for the 27 loci are given above andbelow the diagonal in Table 2, respectively. In the Ds distance, the largest distancewas obtained between the Saisiat and Tsou or Yami, and relatively high values wereobserved between the Atayal and Tsou or Yami. The smallest distance was obtained
between the Puyuma and Ami, and relatively low values were observed between the
Puyuma and Paiwan or Rukai. In the Dc distance, the largest distance was obtained
between the Paiwan and Yami, and relatively high values were observed between the
Tsou and Atayal or Siasiat. The smallest distance was obtained between the Puyuma

Serum Protein Polymorphisms
and Ami,and relatively

betweenPaiwanand a close

lowvaluesand Ami. A distantrelationshipbetweenin Aboriginal Taiwanese Populations

were

observed

relationship
Puyumabetweenbetweenand Amiwere

PuyumaTsou

263
and Rukai

and Saisiat

confirmed

andor Atayal,

by twogenetic

distances.

Figure1.

UPGMA trees for the nine aboriginal groups of Taiwan based on Ds (A) and Dc (B)distance matrix from Table 2.

264

Figure 2.

I. Yuasa et al.

NJ trees for the nine aboriginal
distance matrix from Table 2.

groups of Taiwan based on Ds (A) and Dc (B)The UPGMA trees constructed by using the Ds and Dc distances are shown in Fig.
1. The clustering pattern (Fig. 1A) based on Ds distances was somewhat different
from that (Fig. 1B) based on the Dc distances. On the Ds-UPGMA tree, the nineaboriginal groups were divided into two major clusters. The Ami and Puyuma werethe most closely related to each other, followed by the Paiwan and Rukai. These fourgroups were closely distributed in the southeastern part of Taiwan. The Bunun,
Saisiat and Atayal, living in the central and northern parts of Taiwan, were also
included in this cluster. The Yami and Tsou formed another cluster, though the Yami
lives on Lanyu Island and the Tsou settles in central region of Taiwan. However, thebranch between them was long. The Dc-UPGMA tree confirmed the close relation
shipamong the Puyuma, Ami, Paiwan and Rukai. The Bunun, Saisiat and Atayal
were clustered with these four groups. The Tsou and Yami did not form another loose
cluster and the Yami was the most remote from the other groups. These two groups
were notably different in clustering patterns between the two UPGMA trees.
The NJ trees constructed by using the Ds and Dc distances are shown in Fig. 2.
The clustering pattern (Fig. 2A) based on Ds distances was fairly similar to that (Fig.
2B) based on Dc distances. The Puyuma, Ami, Paiwan and Rukai appeared to be inclose proximity to each other on the two NJ trees. These results were similar to thoseobserved on the UPGMA trees. The Yami, Tsou and Bunun were loosely clustered
and were distant from the Saisiat. The branches of the Atayal, Saisiat, Tsou and Yami
were always long. The branching points of the Atayal were remarkably different
between the two NJ trees.

SerumProtein

Polymorphisms

in Aboriginal

Taiwanese

Populations

265

Comparison of nine aboriginal Taiwanese populations with eight other populations
As shown in Table 3, average heterozygosities and genetic distances (Ds and Dc)
were calculated among 17 human populations including the nine aboriginal Taiwan
esegroups and eight other human populations using allele frequency data for 15 redcell enzyme and serum protein loci. The average heterozygosities ranged from 0.268
in Atayal to 0.339 in Paiwan in the nine aboriginal Taiwanese groups, while theyranged from 0.324 in German to 0.352 in southern Chinese among the other eight
populations. Small within-population variations in the aboriginal groups wereconfirmed in comparison with those in the other populations, though a limited
number of loci were used.
The Ds and Dc distances were also similar to the results obtained using 27 loci.
The Puyuma showed a strong affinity with Ami, followed by the Paiwan and Rukai.
The largest distances were observed between the Yami and Saisiat in the Ds distanceand between the Yami and Paiwan in the Dc distance. In both distances the Yami,
Atayal, Saisiat and Tsou apparently showed the fairly large values for some East
Asian populations, while the other groups showed the small values for some Asian
populations, especially for Filipino. The values were smaller than those between
some aboriginal Taiwanese groups. German among the eight populations had thelargest values for each aboriginal Taiwanese group.
The UPGMA trees constructed by using the Ds and Dc distances are shown in Fig.
3. The clustering pattern (Fig. 3A) based on Ds distances was fairly different fromthat (Fig. 3B) based on Dc distances. On the Ds-UPGMA tree, the 17 populations
were divided into two major clusters. One consisted of the nine aboriginal groups,
but Filipino was included in this cluster with a strong affinity with the Puyuma andAmi. The other consisted of all East Asian populations except for the Filipino.
German was also included in this cluster.
On the Dc-UPGMA tree, German formed the most external branch. The Yami was
located at the position of the out-group to all the remaining 15 populations, whichwere divided into two major clusters. One consisted of the eight aboriginal Taiwan
esegroups, in which the Puyuma and Paiwan showed the strongest affinity and theSaisiat was apart from the others.
The NJ trees constructed by using the Ds and Dc distances are shown in Fig. 4.
The nine aboriginal Taiwanese groups formed a large cluster in the both trees. The
clustering patterns were somewhat different, but the differences were similar to
those observed for 27 loci. The nine aboriginal Taiwanese groups showed a closeaffinity with Southeast Asian populations including Filipinos, Thai and SouthernChinese, but were distant from Northeast Asian populations. Some aboriginal Tai
wanesegroups were apparently close to Filipino than other aboriginal Taiwanesegroups.

2
Table

3.

Genetic

Figuresabove

Allelefrequency

distancesand below

Ds and Dc among

the diagonal

Taiwanese,are Ds and Dc, respectively,

data for 7 red cell enzymeTF) loci were used.

Aboriginal

AK ACP ADA

ESDGermanand East Asian populations

and thoseon the diagonal

GPT PGD

and PGMare average

heterozygosity.

and 8 serum protein

AHSG

GC

HP IF ORM1

ORM2,

PI andSerumFigure 3.

Protein

Polymorphisms

in Aboriginal

Taiwanese

Populations

UPGMA trees for the nine aboriginal groups of Taiwan and eight other populations
based on Ds (A) and Dc (B) distance matrix from Table 3.

267

268

Figure 4.

I. Yuasa et al.

NJ trees for the nine aboriginal groups of Taiwan and eight other populations basedon Ds (A) and Dc (B) distance matrix from Table 3.

Discussion
In the early investigations of red cell enzyme and serum protein systems, weobserved a significant degree of genetic diversity among the nine groups of theaboriginal Taiwanese (Jin et al., 1995, 1999; Umetsu et al., 1994, 1995). In this study
almost all aboriginal Taiwanese groups showed rather small average heterozygosity
in comparison with German and Japanese. However, the large diversity among themwas confirmed by the present study, in which allele frequencies varied from group togroup at as many as nine loci. In particular, the AHSG, C6 and HP systems wereconspicuous for their diversity. The HLA study also showed that the nine aboriginal
Taiwanese groups were highly homogeneous within each group, but diversified
among them (Lin et al., 2000). In the early studies 11 variant alleles including
ADA*Taiwan1, ADA*Taiwan2, ESD*7, GPT1*6, PGM1*6, C1R*9, IF*B2,
ORM1*B9, ORM1*Q0, ORM2*H2 and ORM2*H19 were observed (Jin et al., 1995,
1999; Umetsu et al., 1994, 1995). In this study seven variant alleles including
C3*F07tsou,C3*F06atayal,C6*Ataiwanese,
GC1A8saisiat,GC*1Cami,GC*2Cbunun,
and TF*Dchina,
were observed at four of the ten loci. Some of them were new variants specific for theaboriginal Taiwanese and were polymorphic with a frequency of more than 1%.
ORM1*Q0 and ORM2*H19 were relatively widespread in the aboriginal Taiwanese,but the other variant alleles were restricted to only a few groups. It is of interest that
the Yami group living on Lanyu Island had no such variant alleles. These findings

SerumProteinPolymorphismsin AboriginalTaiwanesePopulations

269

suggest the low level of gene flow among the nine aboriginal groups. The diversity
must also have come from the effect of founder principle and the bottleneck effect.
The tree analysis confirmed the large genetic differentiation among the nineaboriginal Taiwanese groups. In particular, the Atayal, Saisiat, Tsou and Yami al
wayshad a long branch and were far apart form each other, though the branchingpatterns were different from tree to tree. In contrast, the Puyuma, Ami, Paiwan andRukai were always clustered and related to each other. The Bunun were often
included in this group. The clustering patterns were fitted with their geographicaldistribution because they are neighboring with each other at southeastern parts of
Taiwan. Though the Saisiat had a long branch, they were also clustered with a
Puyuma-Ami-Paiwan-Rukai
cluster on the NJ trees. The results described here,
however, were different from those of the HLA study (Lin et al., 2000). It showedthat the Rukai and Paiwan were the most closely related to each other, followed by
the Tsou and Bunun. These four groups were further clustered with the Atayal andSaisiat. The Puyuma, Ami and Yami became more remote from them.
The languages of the aboriginal Taiwanese are classified into four groups of theAustronesian language: the Atayal and Yami speak Atayalic and Malayo-Polynesian,respectively. Tsou and Rukai use Tsouic. The remaining five groups speak Paiwanic(Ruhlen 1987; see Jin et al., 1999). Thus, genetic relationships based on red cell
enzyme and serum protein polymorphisms were fairly identical to the linguisticclassification. The long branch of the Atayal, Tsou and Yami on the trees are consis
tentwith the linguistic classification, because each of the three groups has used theirown peculiar language diversified by a long isolation. Paiwanic is spoken by the
Puyuma, Ami, Paiwan, Bunun and Saisiat, who also formed a large cluster on the NJ
trees. However, the Rukai, who use Tsouic, were clustered with a Puyuma-AmiPaiwan group, but not with Tsou. The Rukai may have genetically been influenced
by the Bunun, Puyuma and Paiwan, because they are surrounded with the Bunun,
Puyuma and Paiwan, but separated from the Tsou. As described above, the Rukai
and Paiwan shared C6*Ataiwanese
at relatively high frequencies.
According to the 'express-train to Polynesia' model (Bellwood 1991; Diamond
1988, 1997), the homeland of the aboriginal Taiwanese was south China. A study of
mtDNA and nuclear DNA variation in the Atayal, Ami, Puyuma, and Bunun sug
gestedthat they had temporally deep roots, probably, in central or south China(Melton et al., 1998). The different waves of migration must have brought ancestral
populations of aboriginal people to the main island of Taiwan. It is noteworthy that
the Atayal and Yami were clustered together with long branches on the Dc-NJ tree.
Chen et al. (1985) cited a reference of Hsu and Pan (1945) and described that the
Atayal group living in Toroko were believed to be the first of all aboriginal Taiwan
esewho moved to Taiwan. If it is correct, the Atayal and Yami may be direct
descendants of the first people, though they live most distantly from each other270

I. Yuasaet al.

among the nine aboriginal Taiwanese groups.
Chen et al. (1985) studied the Atayal group living in Toroko using 19 red cell
antigen, serum protein, and red cell enzyme polymorphisms and concluded that the
Atayal group had a strong genetic affinity with populations from the Philippines andThailand. Melton et al. (1998) described that the aboriginal Taiwanese most resembledpopulations from the Philippines, but this was the result of migration from
Taiwan south to the Philippines. In addition, they deduced that the Ami among thefour aboriginal groups investigated was most probable emigrants. Linguistic andarchaeological evidence agrees that Taiwan is the most likely Austronesian homeland(Bellwood 1991; Diamond 1988, 1997; Gray and Jordan, 2000). In this study
the aboriginal Taiwanese showed genetic affinity with Filipino and to less extent
with Thai and southern Chinese. In accordance with the results of Melton et al.
(1998), the Ami and Puyuma were most closely related to Filipino. Nevertheless,these results were inconsistent with the linguistic classification. The language ofFilipino as well as the Yami belongs to the Malayo-Polynesian, but not to the
Paiwanic (Ruhlen, 1987). mtDNA studies have suggested that Taiwanese spreadthroughout Polynesia by way of Philippines and East Indonesia (Melton et al., 1995;
Sykes et al., 1995; Richards et al., 1998). However, the HLA study of all nineaboriginal Taiwanese showed that they were close to Papua New Guinea Highlandersand Javanese in phylogenetic analysis (Lin et al., 2000). A study of the Y
chromosome markers suggested that Southeast Asia had provided a genetic source
for two independent migrations, one toward Taiwan and the other toward Polynesia
through island Southeast (Su et al., 2000).
Thus, there are two major anthropogenetic interests in the aboriginal Taiwanese:
their genetic origin and colonization to Polynesia. The aboriginal Taiwanese are a
key population to clarify prehistoric Austronesian dispersals. To solve such prob
lemswe are now investigating all nine aboriginal Taiwanese groups for the bloodgroup antigens, mtDNA and Y chromosome.

Acknowledgments
We wish to thank Dr. Hiroaki Nishimukai, Department of Legal Medicine, Ehime
University School of Medicine, for reference typing of the C3 variants found in thisstudy. We are also grateful to Mr. Minoru Endoh and Ms. Miwa Asahi for their
excellent technical clerical assistance. This study was supported by Grants-in-Aid
for the International Research Program ("Human Genetic Studies on Native Minor
ityGroups in Taiwan") to S. H. and by Grants-in-Aid for Scientific Research on the
Priority Areas ("Prehistoric Mongoloid Dispersals" and "The Origin of the Japaneseand Japanese Culture") to I.Y., K.U., N.S. and S.H, from the Ministry of Education,
Science, and Culture of Japan.

SerumProtein

Polymorphisms

inAboriginal

Taiwanese

Populations

271

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