Genetic variation in South Indian castes_ evidence from Y-chromosome, mitochondrial, and autosomal polymorphisms _ SpringerLink

11/19/2019 Genetic variation in South Indian castes: evidence from Y-chromosome, mitochondrial, and autosomal polymorphisms | SpringerLink
Genetic variation in South Indian castes:

evidence from Y-chromosome,
mitochondrial, and autosomal
polymorphisms
BMC Genetics
December 2008, 9:86 | Cite as
WS Watkins (1)
R Thara (2)
BJ Mowry (3) (4)
Y Zhang (1)
DJ Witherspoon (1)
W Tolpinrud (1)
MJ Bamshad (5)
S Tirupati (2)
R Padmavati (2)
H Smith (3) (4)
D Nancarrow (4)
C Filippich (3) (4)
LB Jorde (1) Email author (lbj@genetics.utah.edu)
1. Department of Human Genetics, University of Utah, , Salt Lake City, USA

2. Schizophrenia Research Foundation, , Chennai, India
3. Queensland Centre for Mental Health Research, , Wacol, Australia
4. Queensland Institute of Medical Research, , Herston, Australia
5. Department of Pediatrics, University of Washington School of Medicine, , Seattle,
USA
Open Access
Research article
First Online: 12 December 2008
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Abstract
Background
https://link.springer.com/article/10.1186/1471-2156-9-86 1/49
Major population movements, social structure, and caste endogamy have influenced
the genetic structure of Indian populations. An understanding of these influences is
increasingly important as gene mapping and case-control studies are initiated in
South Indian populations.
Results
We report new data on 155 individuals from four Tamil caste populations of South
India and perform comparative analyses with caste populations from the neighboring
state of Andhra Pradesh. Genetic differentiation among Tamil castes is low (RST =
0.96% for 45 autosomal short tandem repeat (STR) markers), reflecting a largely
common origin. Nonetheless, caste- and continent-specific patterns are evident. For
32 lineage-defining Y-chromosome SNPs, Tamil castes show higher affinity to
Europeans than to eastern Asians, and genetic distance estimates to the Europeans
are ordered by caste rank. For 32 lineage-defining mitochondrial SNPs and
hypervariable sequence (HVS) 1, Tamil castes have higher affinity to eastern Asians
than to Europeans. For 45 autosomal STRs, upper and middle rank castes show
higher affinity to Europeans than do lower rank castes from either Tamil Nadu or
Andhra Pradesh. Local between-caste variation (Tamil Nadu RST = 0.96%, Andhra
Pradesh RST = 0.77%) exceeds the estimate of variation between these geographically
separated groups (RST = 0.12%). Low, but statistically significant, correlations
between caste rank distance and genetic distance are demonstrated for Tamil castes
using Y-chromosome, mtDNA, and autosomal data.
Conclusion
Genetic data from Y-chromosome, mtDNA, and autosomal STRs are in accord with
historical accounts of northwest to southeast population movements in India. The
influence of ancient and historical population movements and caste social structure
can be detected and replicated in South Indian caste populations from two different
geographic regions.
Keywords
Short Tandem Repeat Caste System Lower Caste Caste Group
Tribal Population
These keywords were added by machine and not by the authors. This process is
experimental and the keywords may be updated as the learning algorithm improves.
Electronic supplementary material

The online version of this article (doi: 10.1186/1471-2156-9-86
(https://doi.org/10.1186/1471-2156-9-86)) contains supplementary material, which
is available to authorized users.
Background
The origins and genetic affinities of India's populations have been debated
extensively [1, 2, 3, 4, 5, 6]. Archaeological studies document human occupation of
the subcontinent from the lower Paleolithic through the Neolithic, including a
flourishing ancient civilization in the Indus Valley [7]. The historical record
documents an influx of Vedic Indo-European-speaking immigrants into northwest
India starting at least 3500 years ago [8, 9, 10, 11]. These immigrants spread
southward and eastward into an existing agrarian society dominated by Dravidian
speakers [12]. With time, a more highly-structured patriarchal caste system
developed [7, 9, 10]. India is now broadly characterized by Indo-European (e.g.
Hindi, Urdu, and Punjabi) speaking populations found in the central and northern
regions and by Dravidian (e.g. Tamil, Telugu, and Kannada) speaking populations in
the southern and southeastern regions. The extent to which ancient and
contemporary migrations, and the more recent inception of a hierarchical caste
system, have influenced the genetic composition of modern Indian populations
remains controversial.
A number of studies have addressed the genetic contribution of other Eurasian

populations to Indian caste and tribal populations [1, 2, 3, 6, 13, 14, 15, 16, 17, 18].
They have arrived at somewhat different conclusions regarding the origins of castes,
their relationships to each other, and their relationship to populations outside India.
These discordances can be attributed, in part, to differences in sampling strategies
and the varied effects of gene flow between the typically endogamous castes and
tribes [14, 19, 20, 21].
Several trends regarding the origin and affinities of Indian populations have
emerged. The predominantly south and east Asian mtDNA haplogroup M is found in
more than half of individuals from a wide sampling of castes [5, 6, 13, 22] and is
nearly fixed in some Austro-Asiatic tribal populations [6]. This haplogroup is
uncommon in western European populations [23, 24]. In contrast, some paternally-
inherited Y-chromosome lineages are more closely related to lineages originating in
central Asians and Europeans [1, 13, 25, 26]. Genetic distances estimated from
autosomal polymorphisms have typically demonstrated that caste populations tend to
occupy a position intermediate between European and East Asian populations [8, 27,
28, 29].
The genetic affinities among the more than 2000 extant caste populations of India,
however, are complex. Genetic distances between caste populations from the state of
Andhra Pradesh, India, are correlated with differences in caste rank, suggesting that
endogamy and differential inter-caste gene flow influences genetic structure [30].
Several studies have found a similar pattern, [31, 32, 33] but others have not [6, 34].
Higher rank castes may show closer affinity to European populations than do other
caste populations [13]. Recent Y-chromosome data suggest a higher affinity between
tribal populations and castes of lower rank [35].
These results support historical accounts of nomadic pastoralists from central and
northwestern Eurasia integrating with existing local populations, and either
introducing a system of social stratification or becoming members of the existing
upper castes [8, 9, 35]. Yet, the occurrence of Y-chromosome haplogroups L, H, R2,
and R1a in both caste and isolated tribal populations suggests much of the existing
Indian population structure is very old [5]. Additionally, the high diversity of Y
haplogroups R1a1 and R2 in both South Indian and Indus valley populations has led
to the suggestion that there is little, if any, genetic influence from other Eurasians on
the castes of South India [3].
A broad study of 24 castes from various locations throughout India concluded that
genetic data were not congruent with "sociocultural" affinities due to high rates of
gene flow [6]. Yet, this study and others [1, 36] have suggested a clinal (north to
south) contribution of central Asian Y-chromosomal lineages to caste populations.
Due to well-established clines in gene frequencies across India, especially in the
north-south direction, [2, 34, 36] comparisons of castes from different geographic
locations can conflate clinal variation with variation that may exist between local
caste groups. Therefore, it is important to obtain large, carefully chosen samples from
the same geographic locale to determine whether previous results [13] indicating
caste-related genetic structure can be replicated in other regions of India [37].
Additionally, because single linkage groups such as the non-recombining region of
the Y-chromosome or the mtDNA genome may be strongly influenced by genetic drift
or selection, the use of a large number of independent autosomal polymorphisms can
greatly improve the reliability of estimates of population relationships.
In this study we analyze four castes of different rank sampled from Tamil Nadu, the
southern-most state of the Indian subcontinent. The genetic relationship among the
Tamil castes, their relationship to castes from the neighboring state of Andhra
Pradesh, and their affinity to other Eurasian populations are examined using Y-
chromosome, mtDNA, and autosomal polymorphisms. We show that the genetic
affinities between Indian castes from Tamil Nadu and other Eurasians are broadly
congruent with patterns observed previously for castes from Andhra Pradesh. These
results strengthen the conclusions drawn from our previous analyses regarding caste
relationships in South India and suggest reproducible patterns regarding the genetic
influence of ancient and historical events on the Indian caste system.
Results
Y-chromosome haplogroups
We evaluated the genetic relationship between Tamil castes, eastern Asians, and
Europeans using 32 lineage-specific Y-chromosome SNPs. The sampling locations for
the Tamil castes and a comparative set of castes from the neighboring state of Andhra
Pradesh are shown in Figure 1. Y-haplogroups F*, H1, J2/J2a, L1, R1a1, and R2 reach
appreciable frequencies (> 5%) in most castes. Common Y-haplogroups are typically
shared among castes (Tables 1 and 2). Haplogroup R (predominantly R1a1 (27%) and
R2 (11%)) is the most common major lineage in the Tamil castes, followed by H (21%,
predominantly H1), L (13%, predominantly L1), J (11%, predominantly J2), and F*
(10%).
Table 1
Y-chromosome haplogroup frequencies for South Indian castes and major population
groups
Tamil Nadu Andhra Pradesh Major Geographic Groups
NTS E. S.
Haplogroup Upper Middle Lower Upper Middle Lower Europeans
Upper Asians Indians
C 0.073 0.027 . 0.029 0.030 0.025 0.056 0.018 0.036 0.034
F* 0.049 0.054 0.140 0.206 0.030 0.050 0.204 . . 0.103
G 0.024 . . . 0.030 . 0.019 . . 0.009
H 0.098 . . . . . 0.019 . . 0.016
H1 0.024 0.189 0.163 0.353 0.152 0.250 0.185 . . 0.193
I . . . . . . . 0.316 . .
J2 0.049 . 0.023 . 0.091 0.100 0.056 0.018 . 0.053
J2a 0.073 0.054 0.140 0.088 . 0.050 . . . 0.056
K* . 0.054 . . . . . . 0.071 0.006
L 0.024 . 0.023 0.029 . 0.025 . . . 0.016
L1 0.073 0.108 0.209 . 0.061 0.163 0.130 . . 0.118
O . . . . . . . . 0.464 .
O3 . . . . . . 0.019 . 0.429 0.003
Q . . . 0.088 . 0.025 0.019 0.018 . 0.019
R1 . . . . . . . 0.035 . .
R1a1 0.342 0.432 0.186 0.206 0.515 0.188 0.167 0.053 . 0.267
NTS E. S.
R1b3 . . . . 0.030 . . 0.544 . 0.003
R2 0.171 0.081 0.116 . 0.061 0.125 0.130 . . 0.106
Table 2
Y-chromosome haplogroup counts for South Indian castes and major population
groups
NTS E. S.
C 3 1 0 1 1 2 3 1 1 11
F* 2 2 6 7 1 4 11 0 0 33
G 1 0 0 0 1 0 1 0 0 3
H 4 0 0 0 0 0 1 0 0 5
H1 1 7 7 12 5 20 10 0 0 62
I 0 0 0 0 0 0 0 18 0 0
J2 2 0 1 0 3 8 3 1 0 17
J2a 3 2 6 3 0 4 0 0 0 18
K* 0 2 0 0 0 0 0 0 2 2
L 1 0 1 1 0 2 0 0 0 5
L1 3 4 9 0 2 13 7 0 0 38
O 0 0 0 0 0 0 0 0 13 0
O3 0 0 0 0 0 0 1 0 12 1
Q 0 0 0 3 0 2 1 1 0 6
R1 0 0 0 0 0 0 0 2 0 0
R1a1 14 16 8 7 17 15 9 3 0 86
NTS E. S.
R1b3 0 0 0 0 1 0 0 31 0 1
R2 7 3 5 0 2 10 7 0 0 34
Total 41 37 43 34 33 80 54 57 28 322
(See Materials and Methods for marker information.)
Figure 1
Map of South India. A map of the four major states of South India
shows the sample locations for the caste populations (figure adapted
from Google maps).
Some between-caste trends are suggested by the data. The F* lineage is found at
higher frequencies in lower castes than in upper or middle castes. The R1a1 lineage
occurs at a higher frequency in upper vs. lower castes and differed significantly in
frequency in Andhra upper vs. Andhra lower (p < 0.05). These trends appear in
castes from both Tamil Nadu and Andhra Pradesh. Lineage H also reaches
substantial frequency in the Tamil lower caste but is less common in upper and
middle castes. Lineage J2, previously shown to be distributed in a northwest to
southeast gradient, [3] was present in all castes but not correlated with caste rank.
mtDNA haplogroups
Tamil castes are characterized by high frequencies of mitochondrial M and N super-

family lineages, and all South Indian lineages could be assigned to either M or N
clades (Tables 3 and 4). Both major haplogroup super-families are deep-rooting in
South Indian populations, with diversity estimates for N (0.01589, n = 63) exceeding
that for M (0.01044, n = 92), based on HVS1 data. In contrast to the South Indian
mtDNA haplogroup pool, the eastern Asian and European groups have
predominantly either M or N lineages, respectively. High diversity and deep-
coalescence dates (> 40 K ybp) for both major mtDNA superfamilies are consistent
with an ancient and continuous presence of populations in South India that greatly
predates the documented history of the caste system.
Table 3
mtDNA haplogroup frequencies for South Indian castes and major population groups
NTS E. S.
M-lineages
M* 0.463 0.622 0.442 0.882 0.667 0.600 0.685 . 0.464 0.615
C 0.024 . . . . . . . 0.036 0.003
D . . . . . . . . 0.179 .
G . . . . . 0.013 . . . 0.003
Z . . . . . . . . 0.036 .
M subtotal 0.487 0.622 0.442 0.882 0.667 0.613 0.685 0 0.715 0.621
N-lineages
N* . . 0.023 . . . 0.037 . 0.071 0.009
W . . . . . . . 0.035 . .
R 0.024 0.081 0.186 0.029 0.030 0.138 0.074 0.035 0.214 0.090
R5 0.073 . . . 0.030 0.013 0.111 . . 0.034
J . . . . . 0.013 . 0.105 . 0.003
T 0.024 . . . . 0.025 0.019 0.088 . 0.012
HV 0.073 0.108 0.047 0.029 0.091 0.063 0.019 0.579 . 0.059
U 0.122 . . . . . . . . 0.016
NTS E. S.
U5 . 0.027 . . . . . 0.018 . 0.003
U5a1 . 0.027 . . . . . 0.070 . 0.003
U5a/b . . . . . 0.013 . . . 0.003
U2 0.024 . . . . . . . . 0.003
U2-K 0.098 0.027 0.209 0.059 0.061 0.100 0.019 . . 0.084
U4 0.024 . . . . . . . . 0.003
U7 0.049 0.108 0.093 . 0.091 0.025 0.019 . . 0.050
U8b . . . . . . . 0.070 . .
U9 . . . . 0.030 . 0.019 . . 0.006
N subtotal 0.511 0.378 0.558 0.117 0.333 0.390 0.317 1.000 0.285 0.378
(Lineages are grouped by M or N super-family; *could not be further resolved)
Table 4
mtDNA haplogroup counts for South Indian castes and major population groups
NTS E. S.
M-lineages
M* 19 23 19 30 22 48 37 0 13 198
C 1 0 0 0 0 0 0 0 1 1
D 0 0 0 0 0 0 0 0 5 0
G 0 0 0 0 0 1 0 0 0 1
Z 0 0 0 0 0 0 0 0 1 0
N-lineages
N* 0 0 1 0 0 0 2 0 2 3
W 0 0 0 0 0 0 0 2 0 0
R 1 3 8 1 1 11 4 2 6 29
R5 3 0 0 0 1 1 6 0 0 11
J 0 0 0 0 0 1 0 6 0 1
T 1 0 0 0 0 2 1 5 0 4
HV 3 4 2 1 3 5 1 33 0 19
U 5 0 0 0 0 0 0 0 0 5
U5 0 1 0 0 0 0 0 1 0 1
NTS E. S.
U5a1 0 1 0 0 0 0 0 4 0 1
U5a/b 0 0 0 0 0 1 0 0 0 1
U2 1 0 0 0 0 0 0 0 0 1
U2-K 4 1 9 2 2 8 1 0 0 27
U4 1 0 0 0 0 0 0 0 0 1
U7 2 4 4 0 3 2 1 0 0 16
U8b 0 0 0 0 0 0 0 4 0 0
U9 0 0 0 0 1 0 1 0 0 2
Total 41 37 43 34 33 80 54 57 28 322
(Lineages grouped by M or N superfamily; *could not be further resolved; see

Materials and Methods for marker information.)
To further examine potential western and central Eurasian contributions to South

Indian castes, mitochondrial U lineages, defined by coding variant 12308G, were
analyzed in greater detail (Table 5). U haplogroup subtypes were assigned using key
HVS1 variants as previously described [4, 38]. South Asian lineages U2a and U2c are
common in Tamil and Andhra castes. U7 is the most prevalent U lineage in Tamil and
Andhra castes. U7 is also common in Iran, Pakistan, and northern India, [39]
suggesting an affinity between Dravidian populations from South India and
populations to the north and west. A comparison of HVS1 for U7 haplogroups (10)
with Indian/Pakistani HVS1 sequences available in the mtDB database (4) revealed
similar but non-identical motifs, suggesting ancient rather than very recent gene flow
between northwestern and southern India. A notable between-caste difference is
observed for the mtDNA haplogroup U data in that the Tamil lower caste sample has
a lower frequency of U haplogroups (all subclades) than Tamil upper castes (0.317 vs.
0.059, p < 0.05) or middle castes. This trend is also present in the Andhra sample,
but it is not significant.
Table 5
mtDNA haplogroup U counts for South Indian castes and Europeans
Major geographic
Tamil Castes Andhra Castes
groups
NTS S.
Upper Middle Lower Upper Middle Lower Europeans
Upper Indians
U1a 5 1 1 0 0 0 0 0 7
U2a 1 0 2 0 1 3 0 0 7
U2b 0 0 0 1 0 0 0 0 1
U2c 2 0 4 1 0 4 1 0 12
U2d 0 0 0 0 0 0 0 0 0
U2e 2 0 0 0 0 0 0 0 2
U2i 0 0 2 0 0 0 0 0 2
U4 1 0 0 0 0 0 0 0 1
U5 0 2 0 0 0 1 0 0 3
U5a 0 0 0 0 0 0 0 5 0
U7 2 4 4 0 3 2 1 0 16
K 0 0 0 0 2 1 1 4 4
Total 13 7 13 2 6 11 3 9 55
Genetic distances
We calculated genetic distances between Tamil castes, Europeans, and East Asians
and compared these results to those from upper, middle, and lower caste groups from
the neighboring state of Andhra Pradesh. The genetic distance estimates reveal
several distinct patterns (Table 6).
Table 6
Genetic distance estimates between South Indian castes, Europeans, and eastern
Asians
Y-Chromosome a Autosomal STRs b mtDNA a
E. E. E.
Europeans Europeans Europeans
Asians Asians Asians
Tamil Upper 0.1730 0.6379 0.0026 0.0067 0.2059 0.0772
NTS Upper 0.1947 0.7094 0.0046 0.0082 0.2640 0.0356
Tamil Middle 0.2737 0.5075 0.0037 0.0077 0.1861 0.0810
Tamil Lower 0.3652 0.7060 0.0052 0.0069 0.5951 0.0383
Andhra
0.1992 0.9352 0.0023 0.0082 0.3124 0.0284
Upper
Andhra
0.2527 0.4431 0.0041 0.0064 0.2401 0.0239
Middle
Andhra
0.2693 0.4885 0.0053 0.0070 0.3264 0.0154
Lower
(aSlatkin's linearized FST distance; bDSW distance)
For Y-chromosome polymorphisms, all castes have smaller distances to Europeans

than to eastern Asians. For mtDNA polymorphisms, all castes have smaller distance
estimates to eastern Asians than to Europeans. For Y-chromosome data, the genetic
distance estimates to the Europeans is ordered by caste rank. These trends appear in
castes from both geographic regions.
A neighbor-joining network depicts the between-population relationships based on

Y-chromosome data (Figure 2). The NTS Upper caste is more closely related to the
Andhra Upper caste than to the other Tamil castes, a finding consistent with a
common language (Telugu) shared by the NTS Upper and Andhra upper castes. All
castes are closer to Europeans than to eastern Asians, and basal haplogroup R is
common, especially in the upper castes and Europeans. The inset, however, shows
that haplogroups derived from R are not commonly shared between this sample of
Europeans and southern Indians. Affinity between the groups is driven largely by
basal characters (R, F* and H) that have contrasting frequency patterns.
Figure 2
Genetic distances for Y-chromosome data. A neighbor-joining

network depicts the genetic distance estimates between South Indian
castes, Europeans, and East Asians for 32 Y-chromosome SNPs. The pie
diagrams indicate the proportion of each major Y-chromosome lineage
found in each population. The inset shows the proportions of Y-
chromosome R sub-lineages. Inset circle size is proportional to the total
number of R lineages.
A neighbor-joining network based on distance estimates from 45 STRs shows a

greater affinity of all castes to Europeans than to eastern Asians (Figure 3). With the
exception of the NTS Upper (Telugu and Kannada speaking) Brahmins, castes of
similar rank from different geographic locations tend to branch at similar locations
within the network. Within each geographic region, the distances to other Eurasians
(both Europeans and East Asians) increases with decreasing caste rank.
Figure 3
Genetic distances for autosomal data. A neighbor-joining network

depicts the genetic distance estimates (DSW) between South Indian
castes, Europeans, and East Asians for 45 autosomal STRs.
The network based on mitochondrial distance estimates shows little between-caste

rank organization, yet reveals the greater affinity of all castes to eastern Asians for
maternal lineages (Figure 4). Basal U haplogroups are less frequent in lower rank
castes from both southern India locations. The inset shows that only a few high-
resolution U haplogroups (U5, K) are shared between Europeans and South Indians.
Figure 4
Genetic distances for mitochrondrial data. A neighbor-joining

network depicts the genetic distance estimates between South Indian
castes, Europeans, and East Asians for 32 mtDNA SNPs and 411 bp of
HVS1 sequence. Pie diagrams indicate the proportion of major mtDNA
lineages found in each population. The inset shows the proportion of
mtDNA U sub-lineages. Inset circle size is proportional to the total
number of U lineages.
Genetic structure
The proportion of genetic variation distributed within and between South Indian
castes was assessed by an analysis of molecular variance (AMOVA) (Table 7). The
Tamil South Indian castes are only modestly differentiated from one another: 0.96%
of STR variance occurs between Tamil castes. A similar value of 0.77% for between-
population (caste) difference is observed in the Andhra castes. A smaller fraction,
0.12%, is attributable to geographic differences between Tamil and Andhra locations
and was not significantly different from zero. Removal of the NTS Upper caste from
the comparison yielded a non-significant but higher value of 0.28%. These findings,
based on multiple unlinked loci, suggest that social structure has had a larger impact
on caste population structure in these South Indian samples than geographic
separation.
Table 7
AMOVA for Y-chromosome, 45 autosomal STRs, and mtDNA
Tamil Castes Andhra Castes South Indians
(4 castes) (3 castes) (2 states, 7 castes)
Y-chromosome
Among states - - -1.88a
Among castes 4.45 5.52 5.06
Within castes 95.55 94.48 96.82
Autosomal STRs
Among states - - 0.12a, b
Among castes 0.96 0.77 0.85
Within castes 99.04 99.23 99.02
mtDNA
Among states - - -0.47a
Among castes 4.50 0.17a 2.41
Within castes 95.50 99.83 98.06
(Values are shown as percentages. All values differ significantly from a null
distribution (p < 0.05) unless indicated by [a]. bThis value increases to 0.28 without
the NTS Upper caste but remains non-significant. Tamil castes: Upper, NTS Upper,
Middle, Lower; Andhra castes: Upper, Middle, Lower)
Y-chromosome and mtDNA estimates of molecular variance between castes samples

from either Tamil Nadu or Andhra Pradesh also exceed the estimate for between-
group variation for the two geographic regions. Between-caste variation for mtDNA
in Tamil populations is greater than that for Andhra populations. This may be partly
due to regionally high female mobility in Andhra castes as previously reported [20,
30]. As expected, for all genetic systems, the vast majority of all variation occurs
within populations.
The degree of population subdivision among Indian castes was estimated using a
model-based clustering method implemented in STRUCTURE (ver. 2.1). The best
estimate of the number of clusters (K) was consistently one for the Tamil Indians.
The best estimate of the K clusters was also one for Tamil and Andhra castes
together. This result indicates that individuals from castes spanning the Indian social
hierarchy from two independent geographic regions are not sufficiently differentiated
to allow clustering into groups based on genetic data from 45 STR polymorphisms
alone. This finding is consistent with the low RST values for these populations but
may also reflect the limited power of 45 STRs to distinguish such closely related
populations. Estimates for heterozygosity and repeat variance in these populations
also indicate no substantial between-caste differences or excess homozygosity in
these caste groups (Table 8).
Table 8
STR heterozygosity and variance for South Indian caste populations
Hexp Hobs Alleles per locusa Variancea
0.728
Tamil Upper 0.700 7.1 3.4
(+/- 0.020)
0.716
NTS Upper 0.683 6.6 3.2
(+/- 0.019)
0.718
Tamil Middle 0.681 6.8 3.3
(+/- 0.020)
0.716
Tamil Lower 0.699 6.4 3.3
(+/- 0.021)
0.732
Andhra Upper 0.708 6.7 3.6
(+/- 0.018)
0.721
Andhra Middle 0.693 7.6 3.4
(+/- 0.020)
0.719
Andhra Lower 0.704 7.5 3.4
(+/- 0.020)
0.692
E. Asians 0.697 6.5 3.4
(+/- 0.025)
0.716
Europeans 0.695 7.3 3.2
(+/- 0.017)
(aAverage variance in repeat number over 45 STR loci)
We evaluated the correlation between caste rank and genetic distance using a Mantel
test (Table 9). For each test, a correlation between pairwise genetic and pairwise caste
rank distances matrices using the Tamil caste individuals was calculated. For Tamil-
speaking populations, all genetic systems produced low, significant positive
correlations. Y-chromosome haplogroup data yielded the highest positive correlation
with caste rank (ρ = 0.26, p < 0.01). Inclusion of the non-Tamil speaking Brahmins
decreases the correlation for all systems.
Table 9
Spearman's correlation between genetic distance and caste rank.
Tamil – speaking castes All Tamil castes
Y-chromosome 0.26* 0.15*
45 STRs 0.06* 0.03
mtDNA 0.10* 0.06*
(* p < 0.01)
Discussion
Using a geographically well-defined sample of caste populations from Tamil Nadu,

India, this study arrives at many conclusions similar to those from our previous
studies of caste populations from Andhra Pradesh, India [13, 20, 30]. In both cases,
there is extensive sharing of Y and mtDNA haplogroups among castes, and the overall
level of inter-caste differentiation is low. This finding is consistent with many other
studies of genetic structure and gene flow patterns among caste populations [6, 32,
33, 40].
Paternally-inherited Y-chromosome SNPs show that caste populations have greater

affinity to a sample of Europeans than to a sample of eastern Asians. Unlike the Y-
chromosome data, maternally-inherited mtDNA polymorphisms demonstrate a
contrasting pattern – castes, regardless of rank, have higher affinity to eastern Asians
than to Europeans. These patterns were present in samples from both geographical
locations suggesting that South Indian paternal lineages have been more
substantially influenced by western or central Eurasians compared to South Indian
maternal lineages. Unlike our previous study of Andhra castes, [13] direct haplogroup
sharing between Tamil castes and our sample of Europeans is more limited,
suggesting a potentially greater time depth for the development of these patterns.
More extensive sampling will be required to resolve this difference.
Using Y-chromosome data, Tamil castes of different rank have differential affinities
to our sample of Europeans, with upper castes demonstrating greater affinity than
lower castes. Genetic distances are weakly correlated with caste rank distances and
correlations from Y-chromosome data are stronger than correlations based on
mtDNA or autosomal data. This pattern argues for a differential contribution of male
lineages to castes of different rank and limited male mobility between castes in South
India.
An interesting difference between the data sets from Andhra Pradesh and Tamil
Nadu is also observed. For the former sample, inter-caste distance based on mtDNA
polymorphisms (HVS1 sequence) demonstrated a strong relationship to caste rank,
while distances based on Y-chromosome data did not. This was interpreted as
evidence of historical upward female mobility in the caste system [30]. (We note,
however, that the primary reason for a lack of correlation between Y-chromosome
distances and caste rank was close affinity between the upper-caste Brahmin and
lower-caste Relli samples [20].) In contrast, the Tamil Nadu samples show a higher
correlation between Y-chromosome distances and caste rank than between mtDNA
distances and caste rank. This difference likely reflects differential apportioning of
individuals as the caste system originated or subsequent differences in male-female
mobility patterns.
Recently, several studies have underscored the complexity of Y-chromosome

variation in Indian populations. Sahoo et al. (2006) presented evidence that the R1a
haplogroup has attained high frequencies and high diversity in northern India,
central Asia, and eastern Europe. They also reported high frequencies of Y-
chromosome haplogroup H in caste and tribal populations and provided compelling
evidence for an origin of haplogroup H in South India. Upon further analysis, their
data show that, as in our study, the frequency of haplogroup R lineages is higher in
Brahmins (upper rank) than in lower-rank castes (0.53 vs. 0.41), while the frequency
of H lineages is lower in Brahmins than in lower castes (0.15 vs. 0.34).
In a study of broadly distributed Indo-European and Dravidian castes, Sengupta et al.

(2006) suggested that the majority of Indian Y-chromosome haplogroups are at least
10,000 to 15,000 years old as gauged by Y-chromosome microsatellite diversity, thus
predating the origin of the caste system. The antiquity and complex geographic
distribution of the R1a1 and R2 haplogroups led these authors to conclude that the
majority of the subcontinent Y-chromosomes arrived in or before the early Holocene
(10,000 years ago) rather than in a later Indo-European expansion. Likewise, and
concordant with other studies of tribal Indian populations, [5] we observe Y-
chromosome R1a1 lineages in South Indian tribal Irula (unpublished data), a
population substantially differentiated from South Indian castes [18].
An examination of the R and H haplogroup frequencies of Indo-European-speaking

castes reported by Sengupta et al. (2006) shows that, as in our study, R haplogroup
frequencies in upper castes exceeded those of middle and lower castes (0.62, 0.35,
and 0.38, respectively), while H haplogroup frequencies were lowest in upper castes
(0.14), intermediate in middle castes (0.38), and most frequent in lower castes
(0.44). For Dravidian castes, R (0.62) was more frequent than H (0.14) in upper
castes while R and H had similar (within 6%) frequencies in middle and lower castes.
A recent analysis of caste and tribal populations from eastern India (Orissa)
demonstrated Indo-European influences on paternal caste lineages [41]. Brahmins
showed high Y-chromosome affinity to eastern Europeans (M17, haplogroup R1a1).
In contrast, maternal mtDNA polymorphisms revealed primarily Indian-specific

lineages. Taken together, our studies and at least three other studies of Y-
chromosome lineages in Indian castes demonstrate that upper castes show genetic
affinity to populations residing north and northwest of the Indian subcontinent. This
affinity appears, in part, to result from varying frequencies of Y-chromosome R
lineages and older South Asian lineages such as F* and H.
Indian mtDNA lineages demonstrate high diversity, suggesting that a majority of

Indian maternal lineages are also relatively old and likely predate historically
documented expansion events [38, 42]. Older, deep-rooting mitochondrial lineages
belonging to the N macrolineage are prevalent in western Eurasia and are distributed
in a West – East cline, with high frequencies in Anatolia and Iran and moderate
frequencies in Pakistan and northwestern India [43]. In this study we observe higher
frequencies of basal U lineages in upper castes than in lower castes. Higher resolution
haplogroup results, however, show little evidence of between caste differences. This
may indicate differences in founding populations. More likely, though, it may suggest
ancient migration and integration of various U haplogroups into different pre-caste
populations with subsequent, non-uniform lineage sorting and differentiation over
time. In contrast, and consistent with early human expansion across South Asia, the
predominantly Asian M clade mitochondrial haplogroups account for more than half
of all Indian mitochondrial lineages and reach their highest frequencies in lower caste
and tribal groups [6, 13].
While Y-chromosome and mtDNA polymorphisms yield valuable information, it

must be borne in mind that they each represent a single linkage group. Estimates
based on these systems are thus subject to a high level of stochastic variability [44,
45]. In addition, the Y-chromosome and mtDNA may both have been affected by
natural selection, [46, 47] which can further complicate the interpretation of
population history. Coalescence dates based on these systems must also be viewed
with appropriate caution, in part because of their large confidence intervals. More
importantly, a coalescence date is not necessarily a reliable indicator of the founding
date of a population [45] because these dates are affected by the size of the founder
population and by subsequent gene flow patterns. To gain a more complete and
reliable portrait of population history, multiple, independent autosomal
polymorphisms should also be examined.
Our analysis of 45 unlinked autosomal STRs reveals that in Tamil Nadu, genetic
distances between castes are positively correlated with caste rank. A similar pattern
was detected in upper, middle, and lower rank castes of Andhra Pradesh using these
STRs [20] and Alu and L1 insertion polymorphisms [13]. An analysis of the Kallar,
Vanniyar, and Pallar castes, which also reside in Tamil Nadu, showed that upper –
lower caste distance estimates (0.0553) exceeded those for upper – middle castes
(0.0329) and middle – lower castes (0.0515) [40]. Majumder et al. [37, 48] presented
Y-chromosome, mtDNA, and autosomal data from several caste populations in Uttar
Pradesh. Subsequent analysis indicated that caste rank was correlated with genetic
distance for all three types of systems [20]. Similar correlations have been observed
in a number of other studies of Indian populations [31, 33, 49]. A relatively greater
affinity between upper-caste populations and Europeans has been observed for
autosomal polymorphisms in our Andhra Pradesh and Tamil Nadu samples and in a
number of other analyses of autosomal data [6, 50, 51].
Although significant correlations between caste-rank and genetic distances are

apparent, model-based clustering algorithms did not detect structure within the
Tamil or Andhra populations. We suggest that this finding results from the low
amount of differentiation between all caste groups but also from a lack of sufficient
power in 45 unlinked STRs to detect high-resolution population structure. With ~250
K SNPs typed in a subset of the Andhra upper and Andhra lower castes, individuals
can be clustered into these population groups using genotype information alone [52].
Likewise, using > 950 K SNPs, the Tamil upper and Tamil lower castes demonstrate
group-specific clustering by principal component analysis (unpublished data).
Considering the complex history of Indian populations, it is not surprising that some
studies demonstrate an association between caste rank and genetic distance, whereas
others do not. A recent study of 15 geographically dispersed Indian populations
residing in the United States using 1200 markers found little evidence for caste or
geographic structure [53]. However, sampling strategy (relocated vs. in situ) or other
factors, such as a very wide geographic dispersion of the study populations, may
confound correlations if they exist. Admixture and gene flow can also vary
substantially between caste populations in the various regions of India. Linguistic
differences may influence the genetic structure of local caste populations [34]. The
linguistically different NTS Upper caste Brahmins showed several differences in
comparison to the other Tamil castes in this analysis. Yet, because Indian populations
show only a small amount of genetic differentiation, [17, 53] a large number of
autosomal loci will be necessary for adequate power to detect consistent patterns of
variation if they are present [54, 55]. Ancestry-informative autosomal
polymorphisms, high-density genotyping, and extensive population sampling will
provide better resolution of the relationships between Indian and other Eurasian
populations.
The results presented here underscore the complexity of the Indian caste system.
Although other interpretations may be possible, our data are consistent with a model
in which nomadic populations from northwest and central Eurasia intercalated over
millennia into an already complex, genetically diverse set of subcontinental
populations. As these populations grew, mixed, and expanded, a system of social
stratification likely developed in situ, spreading to the Indo-Gangetic plain, and then
southward over the Deccan plateau. A strong patrilineal social structure,
accompanied by a developing practice of caste endogamy, may have contributed to an
asymmetric apportioning of Y-chromosome, autosomal, and to a lesser extent,
mtDNA lineages. Remnants of these patterns can still be detected in some of the
inhabitants of peninsular South India.
Conclusion
Genetic variation between South Indian castes from Tamil Nadu is low (RST =
0.0096). Tamil caste Y-chromosomes and STR alleles are more similar to Europeans
than to eastern Asians, and genetic distance estimates to Europeans are ordered by
caste rank. In contrast, Tamil caste mtDNA shows greater similarity to eastern Asians
than to Europeans. Low, but statistically significant, correlations between genetic
distance and caste rank can be demonstrated for the Tamil-speaking populations.
These patterns likely reflect asymmetric influences of ancient and historical processes
on the caste system as it developed. These findings provide a general replication of
our analysis of ranked castes from the neighboring state of Andhra Pradesh, India
[13]. For the caste populations analyzed here, between-caste genetic differentiation
exceeds that due to geographic (between-state) differentiation, a finding that may be
of considerable interest when initiating linkage mapping [56] and case-control
association studies in South Indian populations.
Methods
Study Subjects
Study subjects were recruited from four caste groups in Tamil Nadu, India. Tamil-
speaking Brahmins (41), Mudaliars (43), and Dalits (Harijans) (34) were sampled in
Chennai or from rural locations near Chennai. Caste rank was assigned using the
traditional varna of Brahmin (Brahmin, upper ranking), Mudaliar (Sudra, middle
ranking), and Dalit (scheduled caste – outside the traditional caste system, lower
ranking). A second sampling of Brahmins (37) was obtained in Kanchipuram, located
~70 km southwest of Chennai. The Kanchipuram Brahmin group is linguistically
diverse, containing Kannada- and Telugu-speaking Brahmins that relocated from the
neighboring states of Andhra Pradesh and Karnataka. This group of upper castes
individuals is referred to subsequently as the non-Tamil speaking (NTS) Upper caste.
This study was approved by the Schizophrenia Research Foundation, Chennai, India
and by the Wolston Park Hospital Ethics Committee, Brisbane. Approvals were also
obtained from the Indian Council of Medical Research and the Indian Ministry of
Commerce. Written, informed consent was obtained from all participants.
A comparative European sample of northern European and French ancestry (57), and
eastern Asians of Chinese, Japanese, and S.E. Asian ancestry (28) have been
previously described [28, 57, 58]. Because all samples were required to have data for
all genetic systems thus excluding females, sample sizes are smaller than previously
reported. The comparative sample of populations from Andhra Pradesh, India
includes upper-caste Brahmins (33), middle-caste Kapus and Yadavas (80), and
lower-caste Malas, Madigas, and Rellis (54) [13].
Data collection
DNA was extracted from venous blood using standard procedures. Hypervariable
sequence 1 (HVS1), corresponding to base pairs 16000 – 16410, was amplified by
PCR and sequenced using BigDye 3.1 dye-terminator fluorescent sequencing
chemistry and an Applied Biosystems (ABI) 3100 automated sequencer.
Lineage and sub-lineage identifying single nucleotide polymorphisms (SNPs) for the
mitochondria (32 markers) and Y-chromosome (32 markers) were selected from the
literature [47, 59, 60, 61, 62, 63, 64]. Lineage-defining mitochondrial coding region
markers used in the study are L2-C10810T, M-C10400T, C-A13263G, D-C4883T,
preE-G4491A, E-G7598A, G-A4833G, Z-T9090C, N-C10873T, N1d-C6713T, Y-
A7933G, W-G8994A, R-T12705C, R5-T8594C, J-A12612G, T-T10463C, H/V-
T14766C, U-A12308G, U6-A3348G, U6a-G7805A, U5-T3197C, U5a1-A14793G,
U5a/b-A7768G, U2-K-A1811G, U2-A3720G, U2-A9545G, U3-G9266A, U4-T4646C,
U7-C5360T, U7-C8137T, K/U8-G9055A, and U9-G3531A. Y-chromosome lineages
and markers used are C-M216, F*-M89, G-M201, H-M52, H1-M82, H1a-M36, H1b-
M97, H1c-M138, I-M170, J2-M172, J2a-M410, K*-M9, K1-SRY9138, K2-M70, L-
M20, L1-M76, M-M5, N-LLY22g, O-M175, O3-M122, P*-M74, Q-P36, Q3-M3, R*-
M207, R1-M173, R1a-SRY10831.2, R1a1-M17, R1a1a-M56, R1a1b-M157, R1a1c-M87,
R1b3-M269, and R2-M124.
Mitochondrial and Y-chromosome SNPs were genotyped by fluorescent primer

extension using SNaPshot chemistry (ABI). Primers were annealed to amplification
products adjacent to the polymorphic site and extended by one nucleotide using the
manufacturer's recommendations. Extension products were pooled and resolved on a
36-cm capillary array. Four to eight SNPs were assayed per multiplex. Forty-five
STRs, predominantly tetranucleotide repeats, were amplified using 5'-NED, -PET, -
VIC, or -6-FAM labeled primers using standard PCR conditions and resolved in 5
fluorescent multiplex runs on an ABI 3100. STR loci are UT1091, UT1201, UT1205,
UT1220, UT1227, UT1228, UT1232, UT1239, UT1243, UT1257, UT1313, UT1352,
UT8067, UT868, UT871, UT901, UT919, and vWFII. These STRs and mtDNA
polymorphisms, were typed in comparative populations as described previously [13,
18, 20, 57, 58]. Y-chromosome, STR, and mtDNA genotype data is provided in the
Additional_file 1.
To allow a direct comparison of Y-chromosome haplogroups from Tamil Nadu castes

to those from Andhra Pradesh castes, we typed individuals from Andhra Pradesh for
26 of the 32 lineage-defining SNPs. A Y-haplogroup was assigned to each sample by
the presence of one or more derived-state alleles, and the remaining alleles were
inferred. This SNP panel allowed further refinement of the haplogroups previously
reported for the Andhra Pradesh samples [13, 30].
Data analysis
Haplogroups for the Y-chromosome (32 SNPs) and mtDNA (32 SNPs and 411bp
HVS1 sequence) were assigned using SNP data. Mitochondrial haplogroups were
assigned to a haplogroup based on the most probable consensus of polymorphic
changes or resolved using previously published mtDNA HVS1 motifs as a guide [62].
Thirty-one exceptions to the canonical mtDNA phylogeny occurred on 27 mtDNA
haplogroups, and these haplogroups with recurrent mutations were assigned to the
most likely haplogroup based on HVS1 sequence data [4, 6]. The variant 7598A,
defining mtDNA lineage M-E, was found in 2 Tamil and 1 Andhra individuals who
share identical HVS1 motifs but lack the preE 4491A variant. Between-caste
haplogroup differences were evaluated for significance using Fisher's exact test.
Diversity estimates (FST, RST, and AMOVA) for Y-chromosome, mtDNA, and
autosomal STRs were calculated using the ARLEQUIN 3.0 software package [65].
AMOVA statistics were evaluated for significance by comparison to an empirical
distribution generated by random permutation of genotypes or haplogroups. A
general age estimate for mtDNA coalescent dates was calculated by the method of Nei
[66] using a substitution rate of 2 × 10-7 substitutions/site/year [67].
Model-based analyses of population structure were performed using the

STRUCTURE program [68]. An estimate of the optimal number of clusters (K) for
the four Tamil castes was obtained from the posterior probabilities of K, P(X|K),
averaged over 10 runs for each value of K. A uniform prior probability distribution
was assumed on K = {1...n}, and burn-in and iterations were set to 10,000 each for
estimating the best K. Estimates of proportionate membership to three clusters were
averaged values from 10 independent STRUCTURE runs. Population admixture and
correlated allele frequencies were used in all analyses.
The correlation between genetic distance and caste rank was assessed by Mantel
matrix tests using Spearman's rank correlation. For all possible pairs of caste
individuals, inter-individual genetic distance estimates were calculated using
DNADIST (Y and mtDNA) [69] or the Dsw program (STRs) [70]. Next, each
individual was assigned a ranking (1, 2, or 3) for upper, middle, and lower caste
status. The difference in caste rank was calculated for all possible pairs of caste
individuals, yielding a full pair-wise matrix (155 × 155, or 118 × 118 for Tamil-
speakers only) of ordinal values (0, 1, 2). Spearman's rank correlation between the
genetic distance (Y-chromosome, mtDNA, or autosomal STRs) matrix and the caste
rank difference matrix was calculated. A significance level for the correlation was
determined by comparing the actual correlation to a distribution of correlations
generated by 10,000 random columnar permutations.
Notes
Acknowledgements
The authors thank the study individuals for their participation, J.R. Ayankaran for
help in recruiting samples, and J. Xing for helpful discussions. Chris Tyler-Smith
generously supplied primer sequences for LLY22g. This work was supported by NSF
grants SBR-9514733 and SBR-9512178.
Authors' contributions
WSW carried out the molecular studies, performed data analysis, and drafted the
manuscript. RT performed analysis and sample collection in India. BJM and DN
designed and partially funded the study. YZ performed genotyping and laboratory
experiments. DJW provided statistical consultation. WT performed genotyping and
other laboratory experiments. MJB helped acquire and analyze samples from Andhra
Pradesh. ST and RP performed sample collection in Tamil Nadu. HS and CF
performed sample extraction and laboratory analysis of samples from Tamil Nadu.
LBJ designed, coordinated, and funded the study. All authors read and approved the
final manuscript.
Supplementary material
12863_2008_652_MOESM1_ESM.xls (743 kb)
Additional file 1: Y-chromosome, STR, and mtDNA genotype data. Genotype data for
South Indians, Europeans, and eastern Asians. (XLS 743 KB)
12863_2008_652_MOESM2_ESM.pdf (1.9 mb)
Authors’ original file for figure 1
12863_2008_652_MOESM3_ESM.pdf (140 kb)
12863_2008_652_MOESM4_ESM.pdf (69 kb)
12863_2008_652_MOESM5_ESM.pdf (118 kb)
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11/19/2019 Genetic variation in South Indian castes: evidence from Y-chromosome, mitochondrial, and autosomal polymorphisms | SpringerLink

Genetic variation in South Indian castes:

December 2008, 9:86 | Cite as

1. Department of Human Genetics, University of Utah, , Salt Lake City, USA

Electronic supplementary material

A number of studies have addressed the genetic contribution of other Eurasian

Tamil Nadu Andhra Pradesh Major Geographic Groups

C 0.073 0.027 . 0.029 0.030 0.025 0.056 0.018 0.036 0.034

F* 0.049 0.054 0.140 0.206 0.030 0.050 0.204 . . 0.103

G 0.024 . . . 0.030 . 0.019 . . 0.009

H 0.098 . . . . . 0.019 . . 0.016

H1 0.024 0.189 0.163 0.353 0.152 0.250 0.185 . . 0.193

J2 0.049 . 0.023 . 0.091 0.100 0.056 0.018 . 0.053

J2a 0.073 0.054 0.140 0.088 . 0.050 . . . 0.056

K* . 0.054 . . . . . . 0.071 0.006

L 0.024 . 0.023 0.029 . 0.025 . . . 0.016

L1 0.073 0.108 0.209 . 0.061 0.163 0.130 . . 0.118

O3 . . . . . . 0.019 . 0.429 0.003

Q . . . 0.088 . 0.025 0.019 0.018 . 0.019

Tamil Nadu Andhra Pradesh Major Geographic Groups

R1b3 . . . . 0.030 . . 0.544 . 0.003

R2 0.171 0.081 0.116 . 0.061 0.125 0.130 . . 0.106

Tamil Nadu Andhra Pradesh Major Geographic Groups

Tamil Nadu Andhra Pradesh Major Geographic Groups

(See Materials and Methods for marker information.)

Tamil castes are characterized by high frequencies of mitochondrial M and N super-

Tamil Nadu Andhra Pradesh Major Geographic Groups

M* 0.463 0.622 0.442 0.882 0.667 0.600 0.685 . 0.464 0.615

C 0.024 . . . . . . . 0.036 0.003

N* . . 0.023 . . . 0.037 . 0.071 0.009

R5 0.073 . . . 0.030 0.013 0.111 . . 0.034

J . . . . . 0.013 . 0.105 . 0.003

T 0.024 . . . . 0.025 0.019 0.088 . 0.012

HV 0.073 0.108 0.047 0.029 0.091 0.063 0.019 0.579 . 0.059

Tamil Nadu Andhra Pradesh Major Geographic Groups

U5 . 0.027 . . . . . 0.018 . 0.003

U5a1 . 0.027 . . . . . 0.070 . 0.003

U5a/b . . . . . 0.013 . . . 0.003

U2-K 0.098 0.027 0.209 0.059 0.061 0.100 0.019 . . 0.084

U7 0.049 0.108 0.093 . 0.091 0.025 0.019 . . 0.050

U9 . . . . 0.030 . 0.019 . . 0.006

(Lineages are grouped by M or N super-family; *could not be further resolved)

Tamil Nadu Andhra Pradesh Major Geographic Groups

Tamil Nadu Andhra Pradesh Major Geographic Groups

(Lineages grouped by M or N superfamily; *could not be further resolved; see

To further examine potential western and central Eurasian contributions to South

mtDNA haplogroup U counts for South Indian castes and Europeans

Y-Chromosome a Autosomal STRs b mtDNA a

Tamil Upper 0.1730 0.6379 0.0026 0.0067 0.2059 0.0772

NTS Upper 0.1947 0.7094 0.0046 0.0082 0.2640 0.0356

Tamil Middle 0.2737 0.5075 0.0037 0.0077 0.1861 0.0810

Tamil Lower 0.3652 0.7060 0.0052 0.0069 0.5951 0.0383

(aSlatkin's linearized FST distance; bDSW distance)

For Y-chromosome polymorphisms, all castes have smaller distances to Europeans

A neighbor-joining network depicts the between-population relationships based on

Genetic distances for Y-chromosome data. A neighbor-joining

A neighbor-joining network based on distance estimates from 45 STRs shows a

Genetic distances for autosomal data. A neighbor-joining network

The network based on mitochondrial distance estimates shows little between-caste