THIS PAPER CAN NOT BE CITED EXCEPT IN THE CONTEXT OF IWC MEETINGS
SC/56/PFI4

An update of the mitochondrial DNA and microsatellite analyses
in western North Pacific Bryde’s whale
Luis A. Pastene, Mutsuo Goto, and Naohisa Kanda
The Institute of Cetacean Research, 4-5 Toyomi-cho, Chuo-ku, Tokyo 104-0055, Japan

ABSTRACTOn the basis of the JARPNII data presented at the 54SC meeting in Shimonoseki, the Committee agreed that
there was no reason to change the stock structure of Bryde's whales inhabiting western North Pacific in theexisting RMP trials. Further analyses, however, were also recommended in order to pursue a full and detailed
discussion at future meetings. In this paper, we describe the results of mitochondrial DNA control region
sequencing and microsatellite analyses of Bryde's whales samples obtained from different localities of western
North Pacific: JARPNII samples from 2000 to 2003 and the archived samples of past commercial whaling off
Japan (Ogasawara whaling ground and Pelagic samples). A total of 36 haplotypes were detected from sequencevariations at 299bp of mtDNA control region analyzed for a total of 401 animals. All of the 17 microsatellite locianalyzed for a total of 385 animals were polymorphic over the samples. Chi-square statistics were used to test
genetic heterogeneity between sexes within each of the samples, among the samples collected from a same area
in different years (i.e. four JARPNII samples and two Ogasawara area samples) and among all the samples fromdifferent localities in the western North Pacific. Both genetic markers showed the same results as none of thetests were statistically significant. In contrast samples from the western North Pacific showed marked mtDNA
and microsatellite differences with a sample of Bryde’s whale from the eastern Indian Ocean. Our results
strongly indicate that the different samples from the western North Pacific came from a single population. These
results provide no reason to change the stock structure scenario being considered in the RMP trials.

INTRODUCTION
The comprehensive assessment of North Pacific Bryde’s whale was started during the 47th meeting of the IWC
Scientific Committee (IWC, 1996). On the basis of the information derived from genetics and non-genetics
analyses, the Committee defined five stocks: the East China Sea stock (including inshore waters of Kochi), thewestern North Pacific stock, the eastern Tropical stock, the Gulf of California stock and the Solomon
Island/Southeast Asia dwarf stock.
In 1998, the Committee agreed to place two sub-areas (sub-area 1 and 2 divided at 180°) in the western North
Pacific Stock, which created two alternative stock hypotheses for IST to be tested. In 1999, however, somemembers of the Committee concerned that the size of the sub-area 1 was very wide and limited information wasavailable for some parts to make firm conclusion of the structure of the western North Pacific stock. Specifically,
they pointed out that the number of sample used for genetic analyses was scarce in some parts of the sub-area 1
as shown in Appendix 14 of Annex D in IWC (1999).
After the first two years of JARPNII surveys, the results of further genetic analyses were presented (Pastene etal., 2002). The analyses utilized both mitochondrial DNA sequencing and microsatellite analyses for new
samples from a region of the sub-area 1 where the previous genetic analyses that used only historical samples did
not covered. In the analyses, mtDNA control region sequencing analysis involved three localities in the western
North Pacific: JARPN II (2000 and 2001), Ogasawara whaling ground and pelagic operation in the western
central North Pacific. Overall there were no significant mtDNA haplotype frequencies differences among these
localities. Similar results were observed from microsatellite analysis. The authors concluded, therefore, that there
was no evidence to support additional stock structure in the western North Pacific.
After discussion of the JARPNII DNA information at the 54SC meeting, the Committee agreed that the results
reported provided no reason to change the stock structure in the existing trials, but that as much work as possible
should be undertaken before the next meeting to allow a full and detailed discussion at that time. In the1discussion related to the new Implementation Schedule defined at the 54SC meeting it was noted that activities
were still within the Pre-Implementation Review.
In this document, we conducted mtDNA sequencing and microsatellite analyses to examine genetic stock
structure of western North Pacific Bryde's whales. New samples obtained during 2002 and 2003 JARPNII
surveys were added to the analyses, and the number of microsatellite loci used was three times more than that of
Pastene et al. (2002). We believe that these additions should increase the resolution of analyzing stock structureof Bryde's whales.
MATERIALS AND METHODS
Samples
Samples of the ordinary form Bryde’s whales were obtained from different localities in the western North Pacific(Table 1). JARPN II samples included the surveys conducted from 2000 to 2003. Archived samples included
past coastal commercial whaling off Ogasawara and off Taiji and pelagic commercial whaling (Fig. 1). The
samples used for the microsatellite analysis were identical to those used for the mtDNA analysis except some in
the Ogasawara and pelagic samples, which were analyzed with either of the two methods. For comparative
purpose, we used a sample of Bryde’s whale from the eastern Indian Ocean (EIO, n=23) taken in November
1978 during a scientific whaling operation.
MtDNA
Molecular genetic methods used to obtain mtDNA control region sequences were reported in Pastene et al.
(1997a). Following recommendations from the Committee, differences in the frequency of haplotypes among thesamples were tested by the randomized chi-square test (Roff and Bentzen, 1989) with a total of 10,000
permutations in each test. Heterogeneity tests were conducted to test the differences between sexes within thesamples, test the differences among the four JARPNII samples as well as between the two samples from
Ogasawara area (Taiji and Ogasawara), and test among the samples from three different localities: JARPNII,
Pelagic and Ogasawara.
Microsatellites
Genotypes were scored at 17 microsatellite loci: EV1, EV14, EV21, EV94, EV104 (Valsecchi & Amos 1996),
GT011 (Bérubé et al. 1998), GT023, GT310, GT575 (Bérubé et al. 2000), GATA028, GATA053, GATA098,
GATA417, GGAA520, TAA031 (Palsbøll et al. 1997), DlrFCB14, and DlrFCB17 (Buchanan et al. 1996).
Primer sequences and PCR profiles basically follow those of the original authors. All the statistical tests wereconducted using the computer program GENEPOP (Raymond and Rousset, 1995). Statistical tests wereconducted to test the differences between sexes within the samples, test for the deviation from Hardy-Weinberg
genotypic proportion in the samples, test the differences among the four JARPNII samples as well as between
two samples from Ogasawara area (Taiji and Ogasawara), and test among the samples from three different
localities: JARPNII, Pelagic and Ogasawara. Decision of statistically significance on hypothesis testing was
made using the chi-square value obtained from summing the negative logarithm of P-values over the total loci(Fisher, 1950). When needed, sequential Bonferoni correction (Rice 1989) was applied.
Genetic relationshipCavalli-Sforza's cord measures (Cavalli-Sforza and Edwards, 1967) between all pairs of samples were calculated
from mtDNA haplotype frequencies and microsatellite allele frequencies, respectively, using GENDIST of the
PHYLIP (Felsenstein, 1993) program after 1000 replicates of bootstrap analysis. These measures were used tocreate unrooted neighbor-joining trees with NEIGHBOR of the PHYLIP program to analyze the relationships
between Bryde's whale samples. The obtained trees were visualized using the TreeView PPC program (Page,
1996). For this analysis, the samples from eastern Indian Ocean (N=23) were used as outgroups.
RESULTS
MtDNA
Sequence variations at 299bp of the mtDNA control region resulted in 36 unique haplotypes. Haplotype
diversity ranged from 0.670 in the 00NP to 0.888 in the 01NP with an average of 0.816. Without the 00NP, it
was similar among the samples that ranged from 0.800 to 0.816. Percent nucleotide diversity ranged from 0.495
in the Taiji and 1.424 in the 01NP with an average 0.968. Without the Taiji, it was from 0.910 to 1.424. The first
three most frequent haplotypes were common among the samples (see Table 2 in SC/56/PFI5).

2

No significant difference was detected between sexes in each of the samples (Table 2). No significant
difference was found among the four JARPNII samples (Table 4) as well as between the Taiji and Ogasawara
samples. The former four were combined into one as JARPNII and the latter two as Ogasawara for further
analyses. Finally, no significant difference was detected among the samples from the three different localities in
the western North Pacific (Table 5). In contrast, statistically significant difference was detected between thesamples from western North Pacific and that from eastern Indian Ocean (Table 6).
Microsatellites
All loci were polymorphic at each of the samples analyzed (data not shown). The number of alleles per locus
ranged from two at DlrFCB14 to 19 at GATA28 with an average of 8.6. Heterozygosity was quite similar
among the samples ranged from 0.656 in the Taiji sample to 0.681 in the Pelagic sample with an average of
0.669.
No significant difference was detected between sexes in each of the samples (Table 2). A few loci with p-valueless than 5% could be due to chance effect as the test for all loci combined showed no significance at all of thesamples. No significant departure from the Hardy-Weinberg proportions was detected at the 17 loci in each of
the samples (Table 3). Again, a few loci with p-value less than 5% could be due to chance effect as the test for
all loci combined showed no significance at all of the samples. No significant differences were found among thefour JARPNII samples after the correction for the multiple tests (Table 4). Similarly, no significant difference
was found between the Taiji and Ogasawara samples (Table 4). The former four were combined into one as
JARPNII and the latter two as Ogasawara for further analyses. Finally, no significant difference was detectedamong the samples from the three different localities after the correction for the multiple tests (Table 5). In
contrast, statistically significant difference was detected between the samples from western North Pacific andthat from eastern Indian Ocean (Table 6).
Genetic relationships
Genetic relationships of the samples were clearly depicted in unrooted neighbor-joining trees obtained from thefrequencies of mtDNA haplotypes and microsatellite alleles, respectively (Fig. 2). The samples from the western
North Pacific were clustered together very closely while the outgroup EIO sample was quite distant.
DISCUSSION
Pastene et al. (1997b) examined the intra- and inter-oceanic patterns of genetic differences in the Bryde’s whales,
and revealed striking genetic differences among the samples from the different oceanic regions but no significant
differences within the WNP. For that study only two archived samples of past commercial whaling, however,were available for the western North Pacific Bryde's whales: Ogasawara and Pelagic.
Pastene et al. (2002) further analyzed stock structure of Bryde's whales with additional samples from thewestern North Pacific and additional genetic markers. They utilized two hyper-variable genetic markers, namely,
mtDNA control region sequencing and microsatellites to increase resolution and added two JARPNII samples(2000 and 2001 surveys) to the analyses that allowed them to cover wider area in the western North Pacific.
JARPN II surveys obtained the samples from higher latitudes within sub-area 1 approximately between 35° and
45°N compared to the archived samples between 20°N and 30°N, yet different longitudinal localities within subarea 1 (Fig. 1). In Pastene et al. (2002), both mtDNA and microsatellite results showed similar pattern of stock
structure to Pastene et al. (1997b). That is, striking genetic differences were observed at the inter-oceanic level,
but not at intra-oceanic level.
The present study included two additional samples from JARPNII obtained in 2002 and 2003, increasing
sample size for the analysis in the intra-oceanic level. In addition, the number of microsatellite loci analyzed in
the present study (17 loci) was much greater than that in the previous study (5 loci). These improvements shouldhave increased the power of detecting population differentiation.
Genetic similarity of the samples from the western North Pacific was observed from all of the analyses
performed in this study using both of the genetic markers, indicating that these samples came from a geneticallysame group of Bryde's whales. If multiple breeding populations exist, we think we should have detected
temporal genetic differences among samples from multiple-year samplings in feeding grounds. In contrast, eventhough the sample size of the EIO sample was small, substantial genetic difference was detected against the
samples from the western North Pacific. Therefore, our analysis strongly supports the single population
inhabiting western North Pacific.

3

The most effective way to address questions on stock identity is to consider results from several techniques,
genetics and non-genetics (Donovan, 1991; Pastene et al., 2000; Perrin, 2001; Rugh et al., 2003). That is the way
our group has used to address stock identity questions in the North Pacific minke whale, Antarctic minke whaleand bowhead whale. The genetic results of the present study do not support additional stock structure in thewestern North Pacific Bryde’s whale and these results are supported by the analysis of other non-genetic
approaches conducted before or during the CA.
Nemoto (1959) examined mark-recapture data from the Pacific side of Japan and demonstrated movement of
whales between Ogasawara and Sanriku (northern part of Honshu Island). Ohsumi (1978) studied the movementof Bryde’s whale using mark-recapture data. He found that some whales marked near the Equator wererecaptured in both the pelagic grounds of the western North Pacific and in coastal areas of Japan. He used this tosuggest that Bryde’s whales found in pelagic waters belong to the same stock as those found near the coast ofJapan. In a more recent analysis of mark-recapture data, Kishiro (1996) suggested that Bryde’s whales
summering in the whaling grounds (coastal and offshore), winter over a wide latitudinal range, between 1°S and
25°N. The author could not find evidence of more than one stock in the western North Pacific whaling grounds.
Kato and Yoshioka (1993) examined the areal and temporal variations of biological parameters in the western
North Pacific Bryde’s whale with the objective of identifying populations. Mean body length, mean body length
at maturity, pregnancy rate and seasonality of breeding were found to vary on a temporal and spatial basis.
However these authors considered that the variations in these parameters were due to differences in whaling
regulations between coastal and pelagic whaling and under-representation of pregnant females in coastal samples.
They also concluded that there was no evidence supporting the occurrence of different stocks in the western
North Pacific.
We conclude therefore that available genetic and non-genetic evidences support the single stock scenario forwestern North Pacific Bryde’s whale.
ACKNOWLEDGMENTS
We thank researchers and crewmembers of the JARPN II surveys from 2000 to 2003 for sample collections. We
also thank H. Kato, National Research Institute of Far Seas Fisheries, for access to historical samples of the
Bryde’s whale from different oceanic regions and H. Hatanaka, Institute of Cetacean Research, for usefulcomments.
REFERENCES
Bérubé, M., Aguilar, A., Dendanto, D., Larsen, F., Notarbartolo-di-Sciara, G., Sears, R., Sigurjónsson, J., UrbanRamirez, J. and Palsbøll, P.J. 1998. Population genetic structure of North Atlantic, Mediterranean Sea andSea of Cortez fin whales, Balaenoptera physalus (Linnaeus, 1758); analysis of mitochondrial and nuclear
loci. Mol. Ecol. 7:585-599.
Bérubé, M., Jørgensen, H., Mcewing, R. and Palsbøll, P.J. 2000. Polymorphic di-nucleotide microsatellite lociisolated from the humpback whale, Megaptera novaeanglliae. Mol. Ecol. 9:2181-2183.
Buchanan, F.C., Friesen, M.K., Littlejohn, R.P. and Clayton, J.A. 1996. Microsatellites from beluga whale
Delphinapterus leucas. Mol. Ecol. 5:571-575.
Cavalli-Sforza, L.L. and Edwards, A.W. 1967. Phylogenetic analysis: models and estimation procedures. Amer.
J. Human Genet. 19:233-257.
Donovan, G. 1991. A review of IWC stock boundaries. Rep. int. Whal. Commn (special issue 13): 39-68.
Felsenstein, J. 1993. PHYLIP (Phylogeny Inference Package) 3.5c. University of Washington, WA, Seattle.
Fisher, R.A. 1950. Statistical Methods for Research Workers. 11th ed. Oliver and Boy, London.
International Whaling Commission. 1996. Report of the Scientific Committee, Annex G. Report of the subcommittee on North Pacific Bryde’s whales. Rep. int. Whal. Commn. 46:147-59.
International Whaling Commission. 1999. Report of the Scientific Committee, Annex D. Report of the subcommittee on the Revised Management Procedure. J Cetacean Res. Manage. 1 (Suppl.):61-116.

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Kato, H. and Yoshioka, M. 1993. Biological parameters and their areal and temporal variations of Bryde’s
whales in the North Pacific, with special reference to stock identification. Paper SC/45/O8 presented to the
IWC Scientific Committee, April 1993 (unpublished). 18pp.
Kishiro, T. 1996. Movements of marked Bryde’s whales in the western North Pacific. Rep. int. Whal. Commn
46:421-8.
Nemoto, T. 1959. Food of baleen whales with reference to whale movements. Sci. Rep. Whales Res. Inst., Tokyo
14:149-290.
Ohsumi, S. 1978. Bryde’s whales in the North Pacific in 1976. Rep. int. Whal. Commn 28:277-80.
Page, R.D.M. 1996. TREEVIEW: An application to display phylogenetic trees on personal computers. Comput.
Appli. Biosci. 12:357-358.
Palsbøll, P.J., Bérubé, M., Larsen, A.H. and Jørgensen, H. 1997. Primers for the amplication of tri-and tetramer
microsatellite loci in baleen whales. Mol. Ecol. 6:893-895.
Pastene, L.A., Goto, M., Palsboll, P.J. and Kato, H. 1997a. Population structure of the Bryde’s whale(Balaenoptera edeni) in the North Pacific based on analysis of mitochondrial DNA control region
sequences. Document SC/49/NP5 presented to the IWC Scientific Committee, September 1997
(unpublished). 12pp.
Pastene, L.A., Goto, M., Itoh, S., Wada, S. and Kato, H. 1997b. Intra- and inter-oceanic patterns ofmitochondrial DNA variation in the Bryde’s whale, Balaenoptera edeni. Rep. int. Whal Commn. 47:569574.
Pastene, L. A., Goto, M. and Fujise, Y. 2000. Review of the studies on stock identity in the minke whale
Balaenoptera acutorostrata from the North Pacific. Paper SC/F2K/J1 presented to IWC ScientificCommittee Workshop to review JARPN, February 2000(unpublished).
Pastene, L.A., Goto, M. and Kanda, N. 2002. Mitochondrial DNA and microsatellite analyses in the western
North Pacific Bryde’s whale, including samples from JARPNII 2000 and 2001 surveys. Appendix 10,
pp146-155. In: Fujise, Y., Kawahara, S., Pastene, L.A. and Hatanaka, H. (eds.) Report of 2000 and 2001
feasibility study of the Japanese Whale Research Program under special permit in the western North
Pacific-Phase II (JARPNII). Paper SC/54/O17 presented to the IWC Scientific Committee, 2002
(unpublished).
Perrin, W. 2001. Draft review of the utility of non-genetic data for differentiating stocks of whales. Paper
SC/53/SD2 presented to the IWC Scientific Committee, July 2001 (unpublished).
Raymond, M. and Rousset, F. 1995. GENEPOP (version 1.2): population genetics software for exact tests andecumenicism . J. Heredity 86:248-249.
Rice, W.R. 1989. Analyzing tables of statistical tests. Evolution 43:223-225.
Roff, D.A. and Bentzen, P. 1989. The statistical analysis of mtDNA polymorphisms: chi-square and the problem
of small samples. Mol. Biol. Evol. 6:539-45.
Rugh, D., DeMaster, D., Rooney, A., Breiwick, J., Shelden, K. and Moore, S. 2003. A review of bowhead whale(Balaena mysticetus) stock identity. J. Cetacean Res. Manage 5(3):267-279.
Valsecchi, E. and Amos, W. 1996. Microsatellite markers for the study of cetacean populations. Mol. Ecol.
5:151-156.

5

Fig. 1: Sampling localities in the western North Pacific Bryde’s whales used in this study.

Fig. 2 Unrooted neighbor-joining trees based on Cavalli-Sforza's cord measure (Cavalli-Sforza and
Edwards 1967) from frequencies of a) mtDNA haplotypes and b) microsatellite alleles that portray thegenetic relationships among the Bryde's whales samples collected from the western North Pacific(JARPNII, Pelagic, Ogasawara) and the eastern Indian Ocean (EIO). Bootstrap values (%) from 1,000
replicates are given for b). No bootstrap value was smaller than 100% from 1000 replicates in a).

6

Table 1: Bryde's whales samples used in this study

Year Period

MtDNA
Female Male
Total

Sample nameSource

00NP
01NP
02NP
03NP
Pelagic
Off Taiji
Ogasawara

JARPNII
2000 Aug.-Sep. 21
JARPNII
2001 May-July 30
JARPNII
2002 July-Sep. 25
JARPNII
2003 May-Aug. 31
Past commercial 1979 Apr.-May 43
Past commercial 1983 Apr.-June 11
Past commercial 1984 Apr.-June 31

21
13
25
19
70
9
52

42
43
50
50
113
20
83

Microsatellite
Female Male
Total

21
30
25
31
37
12
30

Table 2. Heterogeneity tests between sexes within each of the samples.
Locus

00NP

01NP

02NP

03NP

Pelagic Off Taiji Ogasawara

mtDNA
GATA98
EV104
GT011
GA53
GATA417
DlrFCB14
DlrFCB17
GT23
EV14
GT310
EV1
EV94
GGAA520EV21GT575
GATA28
TAA31
Total loci

0.758
0.044
0.635
0.249
0.386
0.255
0.272
0.567
0.872
0.648
0.235
0.379
0.438
0.552
0.635
0.785
0.803
0.458
0.638

0.351
0.991
0.721
0.747
0.009
0.600
1.000
0.084
0.350
0.117
0.395
0.652
0.747
0.931
0.363
0.374
0.741
0.741
0.587

0.828
0.515
0.479
0.698
0.168
0.912
0.549
0.100
0.165
0.670
0.133
0.156
0.178
0.839
0.848
0.524
0.327
1.000
0.522

0.457
0.803
0.102
0.710
0.609
0.962
1.000
0.589
0.865
0.264
0.203
0.315
0.233
0.646
0.905
0.537
0.728
0.478
0.909

0.288
0.566
0.754
0.030
0.367
0.035
0.557
0.223
0.531
0.208
0.955
0.663
0.925
0.123
0.030
0.647
0.975
0.311
0.177

7

0.900
0.586
0.070
0.755
0.529
0.777
0.356
0.440
0.584
0.769
0.582
0.653
0.640
0.902
0.440
0.563
0.263
0.176
0.835

0.234
0.322
0.737
0.322
0.257
0.482
0.737
0.072
0.119
0.662
0.308
0.106
0.737
0.994
0.864
0.074
0.986
0.274
0.382

21
13
25
19
61
9
51

42
43
50
50
98
21
81

Table 3. Test for the deviation from Hardy-Weinberg genotypic proportion at microsatellite
loci in each of the samples.
Locus

00NP

01NP

02NP

03NP

Pelagic Off Taiji Ogasawara

GATA98
EV104
GT011
GA53
GATA417
DlrFCB14
DlrFCB17
GT23
EV14
GT310
EV1
EV94
GGAA520EV21GT575
GATA28
TAA31
Total

0.781
0.268
1.000
0.499
0.703
0.532
0.972
0.860
0.046
0.378
0.194
0.784
0.488
0.967
0.351
0.303
1.000
0.878

0.786
0.846
0.878
0.989
0.994
0.750
0.999
0.596
0.393
0.131
0.501
0.424
0.279
0.622
0.690
0.792
1.000
0.995

0.551
0.540
0.653
0.714
0.753
0.768
0.013
0.753
0.991
0.085
0.709
0.840
0.556
0.840
0.794
0.201
0.592
0.813

0.309
0.983
0.285
0.100
0.008
0.774
0.976
0.307
0.124
0.946
0.019
0.230
0.639
0.662
0.759
0.817
0.608
0.192

0.386
0.662
0.839
0.370
0.445
0.533
0.948
0.406
0.372
0.443
0.454
0.007
0.668
0.061
0.709
0.325
0.223
0.318

Table 4. Heterogenity tests among the four JARPNII samples
and between the Taiji and Ogasawara samples.
Test results
Locus

JARPNII

Taiji x Ogasawara

mtDNA
GATA98
EV104
GT011
GA53
GATA417
DlrFCB14
DlrFCB17
GT23
EV14
GT310
EV1
EV94
GGAA520EV21GT575
GATA28
TAA31
Total micro.

0.245
0.620
0.100
0.515
0.173
0.359
0.010
0.290
0.515
0.033
0.493
0.454
0.967
0.173
0.361
0.749
0.707
0.254
0.101

0.192
0.433
0.552
0.604
0.913
0.131
1.000
0.628
0.977
0.853
0.622
0.983
0.226
0.356
0.660
0.790
0.741
0.092
0.942

8

0.333
0.595
0.404
0.893
0.289
0.377
0.264
0.001
0.151
0.178
0.114
0.410
0.180
0.423
0.841
0.760
0.214
0.052

0.912
0.333
0.822
0.665
0.895
0.263
0.440
0.075
0.061
0.610
0.259
0.057
0.598
0.236
0.362
0.674
0.645
0.377

Table 5. Heterogeneity tests among the samples
from the different localities: JARPNII, Pelagic,
and Ogasawara.
Locus

Test results

mtDNA
GATA98
EV104
GT011
GA53
GATA417
DlrFCB14
DlrFCB17
GT23
EV14
GT310
EV1
EV94
GGAA520EV21GT575
GATA28
TAA31
Total micro.

0.201
0.034
0.969
0.165
0.539
0.620
0.465
0.932
0.073
0.778
0.760
0.278
0.016
0.133
0.624
0.224
0.512
0.538
0.165

Table 6. Heterogeneity tests between the samples
from the western North Pacific and that fromthe eastern Indian Ocean.
Locus

Test results

mtDNA
EV104
GT011
GA53
GATA417
DlrFCB14
DlrFCB17
GT23
EV14
GT310
EV1
EV94
GGAA520EV21GT575
GATA28
TAA31
Total micro.

0.000
0.000
0.000
0.576
0.557
0.115
0.010
0.000
0.005
0.000
0.052
0.000
0.000
0.000
0.011
0.000
0.000
Highly signif.

9