Annals of Human Biology

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Forensic features and phylogenetic analyses of the

population of Nayagarh (Odisha), India using 23 Y-
STRs

Muktikanta Panda, Ramkishan Kumawat, Shivani Dixit, Awdhesh Narayan

Sharma, Hari Shankar, Gyaneshwer Chaubey & Pankaj Shrivastava

To cite this article: Muktikanta Panda, Ramkishan Kumawat, Shivani Dixit, Awdhesh Narayan
Sharma, Hari Shankar, Gyaneshwer Chaubey & Pankaj Shrivastava (2022): Forensic features and
phylogenetic analyses of the population of Nayagarh (Odisha), India using 23 Y-STRs, Annals of
Human Biology, DOI: 10.1080/03014460.2022.2039762

To link to this article: https://doi.org/10.1080/03014460.2022.2039762

Published online: 02 May 2022.

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ANNALS OF HUMAN BIOLOGY
https://doi.org/10.1080/03014460.2022.2039762

RESEARCH PAPER

Forensic features and phylogenetic analyses of the population of Nayagarh

(Odisha), India using 23 Y-STRs
Muktikanta Pandaa,b
, Ramkishan Kumawatc , Shivani Dixita, Awdhesh Narayan Sharmab, Hari Shankara,
Gyaneshwer Chaubeyd and Pankaj Shrivastavaa,b

a
Department of Home (Police), DNA Fingerprinting Unit, State Forensic Science Laboratory, Government of MP, Sagar, India; bDepartment
of Anthropology, Dr. Harisingh Gour Vishwavidyalaya (A Central University), Sagar, Madhya Pradesh, India; cDNA Division, State Forensic
Science Laboratory, Jaipur, Rajasthan, India; dDepartment of Zoology, Cytogenetics Laboratory, Banaras Hindu University, Varanasi, India

ABSTRACT ARTICLE HISTORY

Aim: The present study was designed to explore the STR diversity and genomic history of the inhabi- Received 12 October 2021
tants of Nayagarh district of Odisha, India. We also tested the proficiency of the most recent, new gen- Revised 21 December 2021
eration PowerPlexR Y23 multiplex system for forensic characterisation and to decipher the Accepted 20 January 2022
phylogenetic affinities.
KEYWORDS
Subjects and methods: The genetic diversity and polymorphism among 236 healthy unrelated male Y-STRs; forensic DNA;
volunteers from Nayagarh district of Odisha, India was investigated. This investigation was carried out Odisha;
via 23 Y-chromosomal STRs using capillary electrophoresis. Nayagarh; haplogroup
Result: A total 223 unique haplotypes were reported. Discrimination capacity (DC), gene diversity (GD)
and power of discrimination (PD) were observed as 0.945, 0.999999999998333, and 0.99999999999794,
respectively. Polymorphic information content (PIC) and matching probability (PM) were reported as
0.999999999925535 and 2.06  1012, respectively. Simultaneously, the haplogroup analysis character-
ised with C2, E1b1a, E1b1b, G2a, H1, I2a, J2a, J2b, L, O, O1, O2, Q, R1a, R2, and T haplogroups, disclos-
ing the possible geographical relatedness of the studied population to different areas of the world.
Conclusion: Phylogenetic analysis with previously reported Indian and Asian populations showed the
genetic closeness of the studied population to different Indian populations and the Bangladeshi popu-
lation of Dhaka, whereas the Bhotra population of Odisha and Han population of China showed much
less genetic affinity.

Introduction with diverse stages of economic as well as cultural develop-

India is a landmass with well-known geographical and lin-
guistic diversities. The continent comprises more than 4000
anthropologically identified ethnic groups (Papiha 1996;
Majumder 1998; Bhasin and Walter 2001; Basu et al. 2003;
Indian Genome Variation Consortium 2008). Odisha state is
the part of Indian continent situated in the Eastern part, at
the coast line of the Bay of Bengal. This political territory
was a part of Gondwana supercontinent around 140 million
years ago (MYA) (Satapathy 2018). The discovery of
Acheulian tools dated to the Lower Palaeolithic era indicated
towards the beginning of human history in Odisha during
the same period (Ghosh 1990). Existing literature indicates
towards different ethnic stocks inhabitating the area over
various time periods, which later on became amalgamated
(Behura and Mohanty 2006). The population of Odisha is
comprised of individuals with enormous diversity with
respect to their social system, cultural configuration, linguis-
tic affiliations and biological make-up. This diversity could be
visible in the different pockets of geography. The population
consists of a large number of tribes and caste groups found
ment, encompassing wide ethnic, cultural and linguistic vari-
eties (Behura and Mohanty 2006). This diversity encourages
the anthropologist and forensic biologist to explore the gen-
etic structure of unexplored endogamous groups at small
diverse geographic pockets with reference to the suitable
genetic markers.
Geographically, the Nayagarh district comes under Odisha
state which lies between 19 540 to 20 320 North Latitude
and 84 290 to 85 270 East Longitude. The district is situated
at the bank of river Mahanadi and is with a number of
streams. Hilly ranges are seen in the West whereas the North
East consists of fertile valleys. The district was formed in
1993 after the splitting of the Puri district into three parts.
Geographically, the territory is surrounded by Khordha,
Kandhamal, Cuttack and Ganjam in East, West, North and
South directions, respectively. The political history of this
region dates back to the 13th century, when Baghela dyn-
asty from the far Rewa area of Madhya Pradesh state came
and set up a new kingdom. Historically the district is known
for its four Garjat states named Nayagarh, Ranpur, Daspalla

CONTACT Pankaj Shrivastava pankaj.shrivastava@rediffmail.com, ecsdsjbp@gmail.com Department of Home (Police), DNA Fingerprinting Unit, State
Forensic Science Laboratory, Government of MP, Sagar, India

Authors with equal credential.
ß 2022 Informa UK Limited, trading as Taylor & Francis Group
2 M. PANDA ET AL.
and Khandapara (Nayagarh District Government of Odisha j
District Administration Portal j India). The territory is spread
over 3890 sq km with a total population of 962789
(Nayagarh District Population Census 2011–2021, Orissa liter-
acy sex ratio and density). Back in the history of human
settlement, the territory was considered as the homeland of
two indigenous communities named Savara and Kandha. It
was later occupied by other migratory castes and state-based
societies on a large scale who settled there and believed
themselves to be native. The population of Nayagarh consti-
tutes a distinct culture, in the form of different dialects, rit-
uals and geographical adaptation from the other parts of the
state of Odisha. More than forty types of the endogamous
ethnic groups reside in this small pocket of geography with
complex vertical and horizontal hierarchy. This encourages a
specified genetic investigation of the area which could pro-
vide information from evolutionary, human settlement and
forensic points of view.
Y-STRs analysis has been extensively used for efficient out-
comes in population genetics studies, anthropological stud-
ies and in forensic identification. The non-recombining
region (NRY) of the Y-chromosome follows the paternal
mode of inheritance. The patterns of current and past gene
flow among several populations can be deciphered through
the analysis of Y-chromosomal variants (Oppenheimer 2012).
Other investigations related to forensic applications, paternity
analysis and human migrations also use the Y-chromosome
polymorphism as a suitable factor (Jobling and Tyler-Smith
2000; Quintana-Murci et al. 2001). The Y-chromosome short
tandem repeats (Y-STRs) are commonly used genetic markers
which belong to the NRY region. These are popularly known
for their applications in demographic history and population
structure analysis, deciphering the lineage relationship in
males, inter-population genetic differentiation and also for
Figure 1. Geographical distribution of the studied population.
their use in forensic molecular biology for the task of per-
sonal identification (Prinz et al. 1997; Roewer et al. 2005;
Lowery et al. 2013; Adnan et al. 2016; Li et al. 2016;
Heraclides et al. 2017; Iacovacci et al. 2017). Interestingly, the
Y-STR can likewise analyse the closely and distantly related
population, defining their potential use in forensic applica-
tion (Ballantyne and Kayser 2012).
In this study, our aim is to discourse the two issues (1) to
test the proficiency of 5-dye based PowerPlexR Y23 multiplex
system (Promega, USA) and evaluate the forensic characteris-
tics among the general male population of Nayagarh district
of Odisha, India, and (2) to explore the phylogenetic affinities
and population structure of the studied population. To the
best of our knowledge, no previous studies have been con-
ducted among any population in any region of Odisha state
using 23 extended Y-marker sets. The findings of this study
could help in deciphering the wide and extensive range of
haplotype data among the general male population of
Nayagarh district.
Subjects and methods
Sample collection
Venous blood samples were collected from 236 healthy unre-
lated male volunteers from different locations of Nayagarh
district of Odisha, India (Figure 1), excluding the minors. All
the persons included in this study were interviewed before
sampling about their relatedness. Before recruiting individu-
als it was ensured that there was no consanguineous or
intercommunity marriage within the latest three generations
for each participant. Written informed consent was taken
from all the volunteers before sample collection.

DNA extraction, PCR amplification and Y-
STR genotyping
Here, we skipped the DNA extraction process and directly
amplified the whole blood samples through PCR as
described in our previously published study (Srivastava et al.
2019; Kumawat et al. 2020). The five-dye based commercial
multiplex kit, Promega PowerPlexR Y23 System (Promega
Corporation, Fitchburg, WI, USA) was used to co-amplify the
23 YSRs (DYS19, DYS389I, DYS389II, DYS385a/b, DYS390,
DYS391, DYS392, DYS393, DYS437, DYS438, DYS439, DYS448,
DYS456, DYS458, DYS635, YGATAH4, DYS481, DYS533,
DYS576, DYS643, DYS549 and DYS570) with 96 well Verity
PCR (Thermo Fisher Scientific, Wilmington, DE, USA). PCR
amplification was carried out as per the manufacturer’s
protocol, excluding the reaction volume which was custom-
ised to 10 ll. The amplified DNA fragments were separated
on a 3500XL Genetic Analyser (Thermo Fisher Scientific,
Waltham, MA, USA) by capillary electrophoresis (CE) and sub-
sequent analysis, as well as Y-STR profile generation, was car-
ried out using GeneMapperR HID Software version 1.6
(Thermo Fisher Scientific, Waltham, MA, USA).
Quality control procedures
Here, different guidelines and recommendations, i.e. recom-
mendations of the DNA Commission of the International
Society of Forensic Genetics (Gusmao et al. 2006; Roewer
et al. 2020) and updated guidelines regarding the publica-
tion of population genetics data (Carracedo et al. 2013) were
followed to carry out the Y-STR genotyping procedures. Our
laboratory had passed the quality assurance test of the
GITAD, Spain (http://gitad.ugr.es/principal.htm) as well as
YHRD, Germany (www.yhrd.org). For quality control purposes,
different laboratory internal control standards, as well as kit
controls (DNA 2800 M as þ ve control), were assigned. In
order to detect the profile quality, different parameters like
the total ratio of peak height as well as inter-locus and inter-
colour balance were analysed for the Y-STR profiles (not
shown here). The generated data of Y-STR haplotypes were
approved from YHRD database (https://yhrd.org/) with the
accession number YA004721, which had been published in
Release 64 of the database.
Data management and statistical analysis
The direct counting method was used to calculate the allelic
frequencies for each 23 Y-STR loci. The haplotype frequen-
cies were calculated using Arlequin software v3.5 (Excoffier
and Lischer 2010). Simultaneously, the genetic diversity (GD)
and haplotype diversity (HD) were estimated by the follow-
ing formula:
 X 
GD or HD ¼ n 1  Pi2 =ðn  1Þ
where, “Pi” specifies the frequency of the ith allele or haplo-
type and “n” specifies the number of samples considered
into the study/analysis. The discrimination capacity (DC) was
determined by following formula:
ANNALS OF HUMAN BIOLOGY 3
DC ¼ h=n
where, “h” specifies the number of different observed haplo-
types and “n” specifies the total sample size.
Multidimensional scaling (MDS) (Kruskal 1964) and ana-
lysis of molecular variance (AMOVA) (Excoffier et al. 1992) to
access the genetic affinity among the study and compared
population were performed using the online tools in Y-
chromosome STR Haplotype Reference Database (YHRD)
(http://www.yhrd.org). Principal component analysis (PCA)
plot and neighbor-joining (NJ) tree construction were
achieved using PAST software (Hammer et al. 2001).
Microsoft Excel, 2016, software was used for the construction
of Heatmap. For AMOVA, PCA, NJ tree and Heatmap, the
studied population data were compared with 15 different
Indian and Asian populations reported previously, viz.
Odisha, India [Bhotra]; Assam, India [Indian] (Singh et al.
2018); Jammu and Kashmir, India [Indian]; Jharkhand, India
[Indian] (Imam et al. 2018); Karnataka, India [Indian]; Madhya
Pradesh, India [Indian] (Shrivastava et al. 2017); Maharashtra,
India [Indian], Rajasthan, India [Indian] (Kumar et al. 2020;
Kumawat et al. 2020); Tamil Nadu, India [Indian]; Uttar
Pradesh, India [Indian]; Uttarakhand, India [Indian]; West
Bengal, India [Indian]; Dhaka, Bangladesh [Bangladeshi]
(Alam et al. 2010); Beijing, China [Han] (Kwak et al. 2005;
Nothnagel et al. 2017; Lang et al. 2019); China [Han] (Zeng
et al. 2014).
Y-haplogroup prediction
Whit Athey’s Haplogroup Predictor has been shown to pre-
dict the haplogroup more efficiently than any other available
software, and it has been found suitable for prediction in
mixed, as well as small, population samples (Petrejc
ıkov
a
et al. 2014; Dogan et al. 2016; Emmerova 2017). Therefore,
haplogroups were assigned from Y-STR haplotype data by
using Whit Athey’s Haplogroup predictor software (Athey
2005, 2006). Specific haplogroups for each haplotype were
predicted by considering the least score at 20 and least
probability at 85% during the assignment.
Results and discussion
Y-STR allele frequencies and variant alleles
For each Y-STR locus, the allele frequencies and gene diver-
sity (GD) values have been mentioned in Tables 1 and 2,
respectively. For the male population of Nayagarh, Odisha, a
total of 44 alleles were observed at 23 Y-STR loci. The range
of allele frequency was found to be between 0.004 and
0.797. Locus-wise, the number of alleles seen were in the
range of 4 (for locus DYS389I, DYS437 and DYS439) to 19
(for locus DYS385a/b). 55 allelic combinations were observed
at the multi-copy loci DYS385a/b. A total of 33 allelic micro-
variants were observed at 11 markers (Table 3); DYS576 (16.3
in single copy), DYS448 (17.2 in single copy), DYS389II (31.1
in single copy and 28.2 in two copies), DYS437 (13.2 in single
copy), DYS570 (18.3 in single copy), DYS635 (23.1 in two cop-
ies), DYS392 (11.2 in four copies,10.2 in two copies and 14.1
 4

Table 1. Observed allele frequency for the population of Nayagarh, Odisha, India (n ¼ 236).
Allele DYS19 DYS389I DYS389II DYS390 DYS391 DYS392 DYS393 DYS437 DYS438 DYS439 DYS448 DYS456 DYS458 DYS481 DYS533 DYS549 DYS570 DYS576 DYS635 DYS643 YGATAH4 DYS385a/b
7 0.004 0.004 0.017 Allele Frequency
8 0.004 0.021 0.089 8,15.2 0.004
9 0.042 0.004 0.254 0.008 0.004 0.169 9,9 0.004
9.2 0.008 9,14 0.004
10 0.720 0.085 0.280 0.326 0.140 0.013 0.424 0.030 10,10 0.004
10.1 0.004 10,14 0.004
10.2 0.008 10,17 0.004
M. PANDA ET AL.

11 0.212 0.678 0.047 0.428 0.390 0.242 0.097 0.153 0.331 10.1,16 0.004
11.2 0.017 11,11 0.030
11.3 11,13 0.008
12 0.093 0.017 0.008 0.203 0.008 0.225 0.538 0.589 0.110 0.394 11,14 0.182
13 0.030 0.555 0.004 0.123 0.547 0.004 0.059 0.042 0.008 0.072 0.250 0.008 0.025 0.233 11,15 0.059
13.2 0.004 11,15.2 0.004
14 0.131 0.347 0.047 0.182 0.797 0.055 0.042 0.042 0.030 0.034 0.013 11,16 0.008
14.1 0.004 11,18 0.004
15 0.555 0.004 0.004 0.008 0.021 0.119 0.581 0.157 0.004 0.085 0.038 11,20 0.004
15.1 0.025 11,22 0.004
15.2 11.3,16.2 0.004
16 0.229 0.081 0.258 0.347 0.157 0.110 12,12 0.017
16.2 12,13 0.004
16.3 0.004 12,14 0.017
17 0.055 0.013 0.051 0.246 0.148 0.203 12,15 0.008
17.2 0.004 12,16 0.004
18 0.004 0.208 0.013 0.144 0.220 0.267 0.004 12,17 0.013
18.3 0.004 12,18 0.004
19 0.004 0.339 0.017 0.237 0.267 0.004 12,19 0.004
19.2 0.004 13,13 0.017
20 0.381 0.008 0.089 0.059 0.131 13,14 0.004
21 0.038 0.055 0.025 0.021 0.017 0.275 13,15 0.013
22 0.233 0.153 0.085 13,16 0.025
23 0.169 0.487 0.356 13,17 0.034
23.1 0.008 13,18 0.025
24 0.047 0.203 0.140 0.085 13,19 0.008
25 0.148 0.309 0.085 0.030 13,20 0.017
26 0.034 0.047 0.055 0.013 14,14 0.030
27 0.004 0.025 14,15 0.034
28 0.059 0.025 0.008 14,16 0.013
28.2 0.008 14,17 0.025
29 0.216 14,18 0.008
30 0.229 14,19 0.013
31 0.178 0.004 14,20 0.013
31.1 0.004 14,21 0.008
32 0.068 15,15 0.047
33 0.004 15,16 0.055
15,17 0.047
15,18 0.030
15,19 0.042
15,20 0.021
15.2,15.2 0.004
16,16 0.008
16,17 0.025
16,19 0.004
16,20 0.004
17,17 0.004
17,19 0.004
20,20 0.004
 ANNALS OF HUMAN BIOLOGY 5

Table 2. Forensic interest parameters observed in the population of performed using Y-STR Haplotype Reference Database
Nayagarh, Odisha, India (n ¼ 236).
(YHRD) tool (http://www.yhrd.org) (Table 4). The genetic dis-
Locus GD PIC PM PD
tance between the studied and the compared populations
DYS19 0.6210 0.5707 0.3816 0.6184
DYS385a/b 0.9433 0.8541 0.0567 0.9433
has been represented in the neighbor-joining (NJ) Tree
DYS389I 0.5648 0.4806 0.4376 0.5624 (Figure 2), principal co-ordinate analysis (PCoA) (Figure 3)
DYS389II 0.8393 0.8152 0.1643 0.8357 and heat map (Figure 4). The study population was found to
DYS390 0.7796 0.7409 0.2237 0.7763
DYS391 0.4360 0.3851 0.5659 0.4341 be genetically closely affiliated to Jharkhand, India [Indian];
DYS392 0.5175 0.4919 0.4847 0.5153 Madhya Pradesh, India [Indian] and Dhaka, Bangladesh
DYS393 0.6267 0.5748 0.3760 0.6240 [Bangladeshi] populations by these analyses. On the other
DYS437 0.3463 0.3185 0.6552 0.3448
DYS438 0.6763 0.6106 0.3266 0.6734 hand, two inland populations of the Indian continent viz.
DYS439 0.6906 0.6270 0.3124 0.6876 Odisha, India [Bhotra] and Uttar Pradesh, India [Indian] as
DYS448 0.6963 0.6355 0.3067 0.6933
well as two other Asian populations viz. Beijing, China [Han]
DYS456 0.5911 0.5374 0.4114 0.5886
DYS458 0.7739 0.7373 0.2294 0.7706 and China [Han] showed distant relationships with the male
DYS481 0.7106 0.6794 0.2924 0.7076 population of Nayagarh, Odisha.
DYS533 0.6299 0.5760 0.3727 0.6273
DYS549 0.5816 0.5263 0.4209 0.5791
DYS570 0.8356 0.8106 0.1679 0.8321
DYS576 0.8009 0.7686 0.2025 0.7975 Haplogroup studies
DYS635 0.7679 0.7314 0.2354 0.7646
DYS643 0.7506 0.7174 0.2526 0.7474 Out of the 236 sampled male individuals, 211 individuals
YGATAH4 0.6830 0.6168 0.3199 0.6801 were assigned to Y-haplogroup through Whit-Athey’s pre-
PD: power of discrimination; GD: gene diversity; PIC: polymorphism informa- dictor. In this study, 16 haplogroups were reported viz. C2,
tion content; PM: matching probability.
E1b1a, E1b1b, G2a, H1, I2a, J2a, J2b, L, O, O1, O2, Q, R1a, R2,
and T (Figure 5). The subclade wise distribution of major
in single copy), DYS643 (9.2 in two copies and 10.1 in single haplogroups has been represented in Figure 6. The four
copy), DYS458 (15.1 in six copies and 19.2 in single copy), major reported haplogroups were R1a (36%), H1 (20.9%), O1
DYS385a (15.2, 10.1, and 11.3 in single copy) and DYS385b (10.9%) and R2 (7.6%) and these constituted >75% of the
(15.2 in three copies and 16.2 in single copy). Here no null total haplogroup distribution.
allele was observed.

Haplogroup R
Haplotype frequencies and forensic parameters
Haplogroup R1a has been most frequently seen in Eurasian
Out of 236 samples, a total of 223 unique haplotypes were regions as well as in different major geographical regions of
reported, of which 211 were observed once, 11 haplotypes the Indian continent (Karafet et al. 2008; Trivedi et al. 2008;
were observed twice and 1 haplotype was observed three Singh et al. 2018). Similarly, the subclade R2 has been found
times (Table 1). The proportion of unique Y-STR haplotypes
to be concentrated in the East Indian region (Sahoo et al.
was calculated as 0.94. The discrimination capacity (DC) was
2006). A Y-STR based study among a Rajasthani population
noted for the studied population as 0.945. The overall gene
also reported R1a and R2 subclades as the two chief hap-
diversity (GD) and power of discrimination (PD) values were
logroups (Kumawat et al. 2020). R1a has been widely seen
found as 0.999999999998333 and 0.99999999999794, respect-
among the populations affiliated with Dravidian, Indo-
ively. The 21 loci (excepting two loci i.e. DYS437 and DYS391
European (Indo-Iranian), Turkic, Slavic and Finno-Ugric lin-
from all 23) showed GD values to be above 0.5, which indi-
guistic crowds (Underhill et al. 2010). The Nayagarh, Odisha
cated that high polymorphism exists in the Y-lineage among
population is affiliated with different languages like the Odia
the male population of Nayagarh, Odisha. Simultaneously, the
(Indo-European), Sora (Austro-Asiatic) and Kui (Dravidian) lan-
polymorphic information content (PIC) and matching prob-
guages. The subclade R1a was the most abundant hap-
ability (PM) were observed as 0.999999999925535 and
logroup seen among 76 individuals and the subclade R2 was
2.06  10–12, respectively. The locus-wise data for GD, PIC, PM
the fourth most abundant haplotype, i.e. seen among 16
and PD are provided in Table 3. DYS385a/b locus was found
to be the most informative locus due to the highest values of individuals in the present study, and together they contrib-
GD (i.e. 0.943263430048837), PIC (i.e. 0.854144679258519) uted 43.6%. Hence, the present findings strongly support the
and PD (i.e. 0.943263430048837). All the discussed forensic previous outcomes regarding the geographical and linguistic
relevance parameters possessed good values making the affiliation of two subclades viz. R1a and R2.
PowerPlexV Y-23 multiplex system potentially suitable for the
R

forensic application among the male individuals of the Haplogroup H

studied population.
Population cross comparison and phylogenetic analysis
An AMOVA test based on the Rst and p values distances
between the studied and compared populations was
Haplogroup H is primarily restricted to the Indian
Subcontinent and classified by two subclades, viz. H1 and H2
(Sengupta et al. 2006; Karafet et al. 2008; Debnath et al.
2011). The subclade H1 was the second most abundant hap-
logroup found among 44 individuals in the present study.
 6

Table 3. Haplotypes observed in the population of Nayagarh, Odisha, India (n ¼ 236).

Haplotype
Code Count DYS576 DYS389I DYS448 DYS389II DYS19 DYS391 DYS481 DYS549 DYS533 DYS438 DYS437 DYS570 DYS635 DYS390 DYS439 DYS392 DYS643 DYS393 DYS458 DYS385a DYS385b DYS456 YGATAH4
1 1 14 13 18 24 15 10 23 13 10 10 14 16 21 24 13 11.2 11 14 15 14 15 16 11
2 1 14 13 18 25 15 10 24 12 12 9 14 19 20 22 11 11 9 12 15 15 17 15 12
3 1 14 13 18 31 14 10 22 11 12 9 14 18 20 24 11 11 9 12 17 13 14 15 11
4 1 14 13 19 25 15 10 23 12 12 9 15 19 20 22 11 10 11 12 17 8 15.2 15 12
5 1 14 13 19 26 15 10 23 12 13 10 14 19 21 22 11 11 9 12 17 13 13 15 11
M. PANDA ET AL.

6 1 14 13 19 32 17 10 31 11 10 9 15 19 21 24 12 14 10 12 14 13 15 13 11
7 1 14 13 20 24 14 10 26 11 11 8 14 15 23 23 11 11 10 13 15 15 15 15 12
8 1 14 14 19 25 16 10 26 12 12 11 15 18 23 24 11 7 11 13 15 11 15.2 15 12
9 1 15 12 19 28 14 10 22 13 13 10 14 15 24 22 12 11 9 11 17 13 13 16 12
10 1 15 13 19 25 15 10 24 12 12 10 16 14 22 22 12 14 11 12 18 9 9 15 11
11 1 15 13 19 29 15 10 22 12 11 10 14 18 18 25 12 12 10 15 17 14 15 14 11
12 1 15 13 19 29 15 10 24 12 12 10 16 14 22 22 12 11 11 12 18 9 14 15 11
13 1 15 13 19 29 15 11 21 13 12 8 14 20 20 21 11 11 9 12 16 15 16 15 11
14 1 15 13 19 29 15 11 24 11 12 9 14 15 21 22 12 11 10 12 17 14 18 15 12
15 1 15 13 19 29 16 10 23 12 10 10 14 17 20 23 12 13 12 14 16 15 20 14 11
16 1 15 13 21 24 15 11 24 13 11 9 14 18 21 23 11 11 12 12 17 13 13 15 12
17 1 15 14 20 31 15 10 21 12 12 11 14 17 23 24 10 10.2 10 13 16 11 14 15 12
18 1 16 12 18 27 15 10 23 11 10 10 14 17 21 25 12 13 12 13 15.1 15 19 16 11
19 1 16 12 19 28 14 10 22 13 12 10 15 15 23 22 13 14 10 11 17 13 17 16 12
20 1 16 13 18 28 15 10 21 13 13 8 14 18 21 21 12 11 8 12 16 15 16 16 11
21 1 16 13 18 29 15 10 21 13 12 8 14 18.3 21 21 10 11 9 14 16 16 17 15 11
22 1 16 13 18 30 16 9 22 13 10 10 14 15 21 25 13 13 12 14 17 14 14 16 11
23 1 16 13 19 24 15 10 23 12 11 9 14 18 21 22 11 11 10 12 17 14 14 15 12
24 1 16 13 19 25 14 10 22 12 11 11 16 18 25 23 10 11.2 11 14 17 12 12 16 11
25 1 16 13 19 25 15 9 22 10 12 9 14 16 20 23 11 11 7 12 17 15 16 16 11
26 1 16 13 19 28 14 10 25 13 11 11 16 16 26 22 11 10 10 14 14 13 20 15 11
27 1 16 13 19 28 15 9 22 10 12 9 14 16 20 23 11 11 7 12 17 15 16 16 11
28 1 16 13 19 29 14 10 25 13 12 11 16 15 24 23 10 10 10 14 17 13 20 15 13
29 1 16 13 19 29 14 10 26 11 11 11 16 17 24 23 12 10 9 14 18 11 22 15 12
30 1 16 13 19 29 15 10 23 12 12 9 14 16 20 22 11 11 9 12 17 15 17 15 12
31 1 16 13 19 29 15 11 23 13 10 9 14 16 21 24 12 11 9 13 17 13 16 13 11
32 1 16 13 19 29 15 11 23 13 10 9 15 16 21 24 12 11 9 13 18 13 16 13 11
33 2 16 13 19 30 15 10 23 12 11 9 14 18 21 22 11 11 10 12 17 14 14 15 12
34 1 16 13 20 25 16 10 28 11 12 10 14 15 21 25 13 11 8 13 16 15 16 15 11
35 1 16.3 14 17 33 15 10 23 12 13 11 14 19 23 24 10 11 10 13 16 11 14 15 13
36 1 16 14 18 31 15 10 22 12 11 9 15 18 22 24 12 11 9 12 18 13 17 13 11
37 1 16 14 18 31 15 10 24 12 12 9 14 17 20 22 11 11 9 13 17 14 17 17 11
38 1 16 14 19 29 15 10 25 11 12 9 14 17 20 21 11 11 9 12 16 15 17 15 12
39 2 16 14 19 31 15 10 22 12 11 9 14 16 20 23 11 11 9 12 18 16 17 15 12
40 1 16 14 19 31 15 10 23 12 12 9 14 18 21 22 11 11 8 12 16 15 17 15 12
41 1 16 14 20 25 17 10 24 13 12 10 14 19 21 22 11 11 7 13 20 15 16 15 12
42 1 16 14 20 30 14 10 28 12 11 11 16 18 24 25 11 10 10 14 16 13 18 15 12
43 1 17 12 19 25 15 10 24 13 12 9 14 20 20 22 11 11 11 13 18 15 16 15 12
44 1 17 12 19 25 15 10 24 13 12 9 15 16 20 22 11 10 11 13 18 15.2 15.2 15 12
45 1 17 12 19 28 14 10 23 12 12 10 15 16 24 22 12 15 10 11 16 12 18 16 12
46 1 17 12 19 28 15 10 23 13 11 9 15 17 22 25 11 11 9 12 15 13 16 13 11
47 1 17 12 19 28 15 9 23 13 11 9 15 17 22 25 11 11 9 11 15 13 16 13 11
48 1 17 12 19 29 14 10 24 12 11 9 14 16 22 24 11 11 10 13 16 14 20 13 12
49 1 17 12 20 30 14 10 22 12 12 9 15 14 22 23 11 11 9 13 15.1 14 19 15 12
50 1 17 13 18 25 15 13 23 12 12 10 14 16 20 22 11 11 9 12 16 15 17 15 12
51 1 17 13 18 29 15 10 23 12 12 9 14 19 21 22 12 11 8 12 17 15 16 17 12
52 1 17 13 18 29 15 10 23 12 12 9 14 20 21 22 12 13 8 12 17 15 16 17 12
53 1 17 13 18 29 16 10 24 12 10 9 15 17 22 24 12 11 9 12 16 13 17 13 11
54 1 17 13 19 24 16 10 25 12 11 11 15 19 24 23 11 10 11 13 18 11 15 15 12
55 1 17 13 19 25 15 10 21 12 12 9 14 18 20 23 11 13 10 12 18 15 16 16 12
(continued)
Table 3. Continued.
Haplotype
Code Count DYS576 DYS389I DYS448 DYS389II DYS19 DYS391 DYS481 DYS549 DYS533 DYS438 DYS437 DYS570 DYS635 DYS390 DYS439 DYS392 DYS643 DYS393 DYS458 DYS385a DYS385b DYS456 YGATAH4
56 1 17 13 19 30 15 10 21 12 12 9 14 19 21 22 10 11 10 12 17 15 15 15 12
57 1 17 13 20 25 15 11 23 12 12 11 14 18 23 24 10 11.2 10 13 16 16 16 16 13
58 1 17 13 20 25 16 10 22 12 12 11 14 18 23 25 10 11.2 8 13 15 14 16 16 13
59 1 17 13 20 28 14 10 25 13 11 11 16 17 24 23 10 10 10 14 16 13 17 15 12
60 1 17 13 20 29 14 10 25 12 13 11 14 18 21 23 11 13 9 13 19.2 11 18 16 10
61 1 17 13 20 29 14 10 26 11 11 8 14 15 23 23 11 11 10 13 15 15 15 15 12
62 1 17 13 20 30 16 10 22 12 12 11 14 18 23 25 10 11 8 13 15 11 14 16 13
63 1 17 13 20 30 16 10 22 13 12 11 14 18 23 25 10 11 8 13 15 11 14 16 13
64 1 17 13 20 30 16 10 22 13 12 11 14 18 23 25 10 11 8 13 15 12 14 16 13
65 1 17 13 20 30 16 11 23 12 12 11 14 19 23 26 10 11 10 13 17 11 14 16 13
66 1 17 13 20 30 17 11 23 13 12 11 14 19 23.1 25 10 11 10 13 15.1 11 14 16 13
67 1 17 13 20 31 16 10 23 13 12 11 14 18 23 26 10 11 8 13 15 11 14 16 13
68 1 17 13 20 31 17 11 23 12 12 11 14 19 23 24 10 11 10 13 16 11 15 15 12
69 1 17 13 20 31 17 12 23 12 12 11 14 19 23 24 10 11 10 13 16 11 15 15 12
70 1 17 13 21 29 14 10 25 14 11 12 15 18 24 24 11 10 9 14 17 14 18 15 12
71 1 17 13 21 31 16 11 23 13 13 11 14 18 23 25 10 11 10 13 15 11 14 15 13
72 1 17 14 18 25 15 10 23 12 12 9 14 15 20 22 11 11 9.2 12 18 16 17 15 11
73 1 17 14 18 25 15 10 23 12 12 9 14 18 20 22 11 13 8 12 18 15 17 15 12
74 1 17 14 18 26 15 10 23 12 10 10 14 16 21 25 12 13 12 13 17 15 18 15 11
75 1 17 14 18 30 17 10 24 14 12 10 14 16 22 22 12 11 9 12 18 15 18 15 12
76 1 17 14 18 31 15 10 22 12 10 9 15 17 21 24 12 11 10 12 19 15 15 13 11
77 1 17 14 19 25 15 9 25 12 12 10 16 16 24 22 13 11 10 13 17 12 17 15 12
78 1 17 14 19 30 15 10 23 12 11 11 16 20 25 23 11 10 10 13 15 13 18 16 12
79 1 17 14 19 30 15 10 25 12 11 11 16 19 25 23 11 10 10 13 16 13 18 16 12
80 1 17 14 19 30 15 10 26 12 11 11 16 20 25 23 11 10 10 13 15 13 18 16 12
81 1 17 14 19 30 16 10 28 11 12 10 15 17 21 21 12 11 12 13 16 13 17 15 11
82 1 17 14 20 25 15 10 23 12 12 11 14 21 23 24 10 11 10 13 17 11 14 15 13
83 1 17 14 20 25 17 12 22 13 12 11 14 19 23 25 10 11 10 14 16 20 20 14 12
84 1 17 14 20 30 14 10 25 13 11 11 16 18 25 24 11 10 10 13 17 14 20 17 11
85 1 17 14 20 30 15 10 23 12 12 11 14 20 23 24 10 11 11 13 15 11 14 15 12
86 1 17 14 20 30 15 10 25 12 11 11 16 19 25 23 11 10 10 13 16 13 18 16 12
87 1 17 14 20 30 15 11 23 11 12 7 14 18 21 22 11 11 9 13 17 15 17 16 12
88 1 17 14 20 32 15 10 24 14 12 11 14 20 23 23 10 11 10 13 16 11 14 15 12
89 1 17 14 20 32 15 10 24 14 12 11 15 20 23 23 10 11 11 13 16 11 14 15 12
90 1 17 14 21 30 15 10 24 12 10 10 14 16 21 23 12 11 8 14 16 10 17 14 10
91 1 18 12 17 24 15 10 23 12 11 10 14 17 23 25 11 11 12 14 16 15 15 16 11
92 1 18 12 18 29 15 10 23 11 10 10 14 14 21 25 12 13 11 14 16 14 15 16 11
93 1 18 12 19 24 15 10 24 13 11 11 15 15 24 23 11 14 10.1 14 19 12 12 15 12
94 1 18 12 19 30 15 10 23 13 13 10 14 15 24 22 12 14 10 11 16 13 18 18 11
95 2 18 12 21 29 15 11 23 12 9 10 16 18 21 22 11 11 12 14 15 13 15 14 11
96 1 18 13 18 25 15 10 25 12 12 9 14 18 21 22 12 11 9 12 14 15 18 15 12
97 1 18 13 18 28 15 11 23 12 10 10 14 16 21 26 12 14 12 14 16 14 19 15 10
98 1 18 13 18 29 14 11 23 12 10 10 14 14 22 25 12 13 11 14 16 15 20 16 11
99 1 18 13 18 29 15 11 23 12 10 10 14 16 21 25 12 13 12 13 17 15 19 16 11
100 1 18 13 18 29 15 11 25 12 10 10 14 17 21 25 11 13 11 14 16 14 15 16 11
101 1 18 13 18 30 15 10 23 11 10 10 14 16 21 25 12 13 11 13 16 15 20 15 11
102 1 18 13 18 30 15 10 23 11 10 10 14 16 21 25 12 13 12 13 16 15 20 15 11
103 1 18 13 18 30 15 10 24 12 10 10 14 16 21 25 11 13 12 14 15 15 20 16 11
104 1 18 13 19 28 14 10 27 13 11 11 14 15 24 23 10 10 10 14 16 13 19 16 11
105 1 18 13 19 29 15 10 22 12 12 9 14 18 19 22 11 11 9 11 18 15 16 15 13
106 2 18 13 19 30 16 10 23 12 11 9 14 18 21 22 12 11 9 12 17 14 17 15 12
107 1 18 13 19 31.1 16 10 23 12 11 9 14 18 21 22 12 11 9 12 17 14 17 15 12
108 1 18 13 20 24 13 10 26 14 11 10 14 19 21 25 12 11 11 13 19 14 15 15 11
ANNALS OF HUMAN BIOLOGY

109 1 18 13 20 24 16 10 22 12 12 13 14 17 23 25 11 11 10 13 16 11 14 15 13
110 1 18 13 20 25 15 10 25 12 12 11 14 20 23 25 10 11 13 14 13 14 14 16 13
(continued)
7
 8

Table 3. Continued.
Haplotype
Code Count DYS576 DYS389I DYS448 DYS389II DYS19 DYS391 DYS481 DYS549 DYS533 DYS438 DYS437 DYS570 DYS635 DYS390 DYS439 DYS392 DYS643 DYS393 DYS458 DYS385a DYS385b DYS456 YGATAH4
111 1 18 13 20 25 15 11 24 12 12 11 13.2 18 23 25 10 11 12 13 16 10 14 15 13
112 1 18 13 20 25 16 10 23 11 12 11 14 18 23 26 10 11 10 13 16 11 11 15 13
113 1 18 13 20 26 14 11 22 12 13 9 15 15 21 23 11 9 10 13 15 11 15 15 12
114 1 18 13 20 26 14 11 22 13 13 9 14 16 22 23 11 11 10 12 16 17 17 15 11
115 1 18 13 20 29 14 11 22 11 11 9 16 16 23 24 11 11 10 12 16 14 21 16 12
116 1 18 13 20 30 15 10 27 12 10 9 14 20 21 24 11 11 11 13 20 14 15 14 12
M. PANDA ET AL.

117 1 18 13 20 30 16 11 23 12 12 11 14 20 23 25 11 11 10 13 16 11 14 15 13
118 1 18 13 20 31 14 11 22 12 13 9 15 15 21 23 11 11 10 13 15 15 18 15 12
119 1 18 13 20 31 16 10 23 11 12 11 14 18 23 26 10 11 11 13 16 11 16 15 13
120 1 18 13 20 31 16 10 23 13 11 11 14 20 23 25 11 11 10 13 16 11 14 15 13
121 1 18 13 20 31 16 10 23 13 12 11 14 19 23 25 10 11 10 13 18 11 14 17 12
122 1 18 13 20 31 16 10 24 13 12 11 14 20 23 25 10 11 8 13 14 11 14 16 13
123 1 18 13 20 31 16 11 23 12 12 11 14 19 23 25 10 11 11 13 16 11 14 15 13
124 1 18 14 17 30 15 11 23 14 10 10 14 17 21 25 11 13 12 14 16 16 17 15 11
125 1 18 14 18 28 15 10 23 12 10 10 14 16 22 25 12 10 10 14 17 14 14 14 11
126 1 18 14 18 30 15 10 23 13 10 10 14 17 22 25 13 13 12 15 16 15 19 16 11
127 1 18 14 18 30 15 11 23 12 10 11 14 16 22 25 13 13 12 15 15 16 19 18 11
128 1 18 14 18 30 15 11 23 12 11 10 14 15 21 25 12 12 12 15 16 16 20 15 11
129 1 18 14 19 24 14 10 26 12 11 11 16 19 25 21 12 10 10 14 17 14 21 16 12
130 1 18 14 19 29 15 10 23 12 11 9 14 16 20 22 11 11 9 11 18 14 17 16 12
131 1 18 14 19 30 14 10 23 12 12 9 14 17 20 22 11 11 9 11 18 16 17 15 12
132 1 18 14 19 30 15 10 23 13 12 9 14 17 20 22 11 11 8 12 18 15 17 17 12
133 2 18 14 19 30 15 10 23 13 12 9 14 17 20 22 11 11 8 13 18 15 17 17 12
134 1 18 14 19 30 15 10 28 14 11 10 14 19 21 23 11 11 10 13 18 12 14 16 12
135 1 18 14 19 30 15 9 25 12 12 10 16 16 22 21 12 11 10 13 17 12 16 17 12
136 1 18 14 19 30 16 10 26 12 12 11 14 18 23 24 11 13 12 13 15 11 13 15 12
137 1 18 14 19 31 17 11 26 12 11 10 15 15 22 21 11 11 11 13 15 12 19 15 12
138 1 18 14 20 25 16 10 22 12 12 11 14 17 23 25 10 11 10 13 16 15 15 15 13
139 1 18 14 20 25 16 10 22 12 12 11 14 19 23 25 10 11 10 13 16 11 14 17 13
140 1 18 14 20 25 16 10 23 12 12 11 14 18 23 25 10 11 10 13 16 11 15 15 11
141 1 18 14 20 29 15 9 23 12 12 10 14 17 21 23 12 11 8 12 17 14 15 15 13
142 1 18 14 20 30 16 10 28 11 12 10 15 17 21 22 12 11 13 13 16 13 17 15 11
143 1 18 14 20 31 15 10 22 12 12 11 14 19 24 26 10 11 10 13 14 14 15 15 12
144 1 18 14 20 31 15 10 24 12 12 11 14 20 23 25 10 11 10 13 16 11 14 16 13
145 1 18 14 20 31 16 10 23 12 12 10 14 18 23 25 10 11 10 13 16 11 15 15 11
146 1 18 14 20 31 16 11 24 12 12 11 14 19 24 24 10 11 10 13 15.1 11 14 15 13
147 1 18 14 20 32 15 10 23 13 12 11 14 20 23 25 10 11 10 13 17 11 14 15 11
148 2 18 14 21 31 15 10 22 12 12 11 14 19 23 24 10 11 10 13 17 11 14 15 11
149 1 18 15 21 32 15 11 22 12 12 11 14 19 23 24 10 11 10 13 17 11 14 15 11
150 1 19 12 19 28 15 10 24 12 12 9 14 19 20 22 11 11 9 13 18 15 18 15 12
151 1 19 12 19 29 13 10 23 12 12 10 15 15 22 22 12 14 10 11 14 13 17 16 12
152 1 19 12 19 29 13 10 23 12 12 10 15 15 23 22 12 14 10 11 14 13 17 16 12
153 1 19 12 19 29 15 15 28 13 11 9 14 18 22 24 11 11 11 13 17 12 13 15 12
154 1 19 12 20 28 14 10 23 11 11 11 15 17 22 24 12 14 13 12 18 13 19 15 12
155 1 19 13 17.2 31 15 11 24 12 12 11 14 19 23 25 10 11 10 13 16 11 15 15 13
156 1 19 13 18 25 15 10 24 12 12 9 14 17 20 22 11 11 9 12 15 10.1 16 15 12
157 1 19 13 18 29 14 10 24 13 11 10 14 14 20 24 11 11 12 13 17 15 16 15 11
158 2 19 13 18 29 14 10 24 14 11 10 14 13 21 24 11 11 12 13 16 15 15 16 10
159 1 19 13 18 29 15 10 23 12 10 11 14 16 21 25 12 13 9.2 14 16 15 19 17 11
160 1 19 13 18 29 15 10 23 9 12 9 14 17 22 22 11 11 8 12 17 15 16 15 12
161 1 19 13 18 29 15 10 24 12 12 9 14 17 20 22 11 11 9 12 15 15 17 15 12
162 1 19 13 18 29 15 10 24 13 11 10 14 14 20 24 12 11 12 13 17 15 15 15 11
163 1 19 13 18 29 16 10 23 13 10 10 14 16 21 24 13 14 11 14 15 17 19 15 11
164 3 19 13 18 30 15 10 23 13 10 10 14 16 21 24 13 13 11 14 15 15 19 16 11
165 1 19 13 18 30 15 11 23 13 10 10 14 16 21 24 13 13 11 14 15 15 19 16 11
166 2 19 13 19 29 15 10 27 13 11 10 14 18 21 23 11 11 11 13 18 11 14 16 12
167 2 19 13 19 30 15 10 23 12 12 11 14 17 23 25 10 11 10 13 16 11 11 15 13
(continued)
Table 3. Continued.
Haplotype
Code Count DYS576 DYS389I DYS448 DYS389II DYS19 DYS391 DYS481 DYS549 DYS533 DYS438 DYS437 DYS570 DYS635 DYS390 DYS439 DYS392 DYS643 DYS393 DYS458 DYS385a DYS385b DYS456 YGATAH4
168 1 19 13 19 31 16 10 26 12 11 10 14 19 21 24 12 11 10 13 17 12 12 14 12
169 1 19 13 19 31 17 10 23 12 12 11 14 19 23 24 11 11 10 13 14 11 11 16 12
170 1 19 13 20 25 15 10 22 12 12 9 14 16 23 25 10 11 10 13 16 16 16 15 11
171 1 19 13 20 25 15 10 23 12 12 11 14 21 23 25 10 11 10 13 16 11 11 15 13
172 1 19 13 20 25 16 10 23 12 12 11 14 19 23 25 10 10.2 11 13 16 12 14 15 13
173 1 19 13 20 25 16 10 23 12 12 11 14 19 24 23 11 11 11 13 18 11 15 16 13
174 1 19 13 20 25 16 11 23 12 12 11 14 19 23 25 10 18 8 13 15 10 10 15 14
175 1 19 13 20 28.2 17 10 23 12 12 11 14 19 24 24 10 11 10 13 14 11 11 15 12
176 1 19 13 20 28.2 17 10 23 12 12 11 14 19 24 24 10 11 10 13 14 11 14 15 12
177 1 19 13 20 29 15 9 23 12 12 10 14 17 26 23 13 11 8 11 17 15 15 15 13
178 1 19 13 20 30 16 11 23 13 12 12 14 18 23 24 10 11 10 13 15 11 14 15 12
179 1 19 13 20 31 15 10 23 12 12 11 14 21 23.1 22 11 14 10 13 15.1 14 16 15 13
180 1 19 13 20 31 15 10 23 12 12 11 14 21 23 25 10 11 10 13 16 11 14 15 13
181 1 19 13 20 31 15 10 25 12 12 11 14 18 23 24 10 11 10 13 17 11 15 15 12
182 1 19 13 20 31 15 10 25 13 11 9 14 19 23 24 11 11 11 13 18 13 16 14 13
183 1 19 13 20 31 16 11 23 12 12 11 14 19 23 25 10 11 10 13 15 15 15 16 13
184 1 19 13 20 31 16 11 23 12 13 11 14 19 23 26 10 11 9 13 16 11.3 16.2 15 13
185 2 19 13 21 30 16 11 23 12 12 11 14 19 23 25 10 11 10 14 16 11 14 16 13
186 1 19 14 18 30 15 11 23 12 10 10 14 17 23 25 12 13 13 15 18 15 19 14 11
187 1 19 14 18 31 15 10 23 11 11 10 14 16 20 25 12 13 12 14 17 14 19 16 11
188 1 19 14 19 24 13 10 27 13 11 11 14 17 23 22 12 19 10 13 17 13 13 16 10
189 1 19 14 19 29 14 11 23 11 12 11 14 19 28 22 12 10 10 13 18 13 20 15 11
190 1 19 14 19 29 14 11 25 11 12 11 14 19 28 22 12 10 10 13 18 13 20 15 11
191 1 19 14 19 30 13 10 25 12 11 11 15 18 23 22 11 15 11 13 19 12 15 15 10
192 1 19 14 19 30 15 9 24 13 12 9 14 19 20 22 11 11 7 12 17 15 18 16 11
193 1 19 14 19 31 15 10 23 12 12 11 14 19 23 25 10 11 10 13 16 11 14 15 13
194 1 19 14 19 31 17 10 26 10 12 10 15 15 23 21 11 11 11 13 15 11 20 16 12
195 1 19 14 20 26 15 10 24 12 13 11 14 18 23 25 10 8 13 14 16 11 14 15 11
196 1 19 14 20 26 16 10 23 14 13 11 14 19 23 25 10 10 10 13 16 14 17 15 12
197 1 19 14 20 26 16 11 23 12 13 11 14 19 23 26 10 11 10 13 16 11 14 15 13
198 1 19 14 20 26 16 11 23 12 13 11 14 19 23 26 10 11 11 13 16 11 14 15 13
199 1 19 14 20 31 15 10 25 13 11 9 14 19 23 24 11 11 11 13 18 13 16 14 13
200 1 19 14 20 32 15 10 23 12 12 11 14 18 23 25 10 11 10 13 16 11 16 14 12
201 1 19 14 20 32 15 10 23 13 12 11 14 18 23 25 10 11 10 13 16 11 15 15 11
202 1 19 14 20 32 15 10 23 13 12 11 14 20 23 25 10 11 10 13 17 11 11 15 11
203 1 19 14 20 32 16 10 22 12 12 11 14 17 24 25 10 11 10 13 16 11 15 15 13
204 2 19 14 20 32 16 12 23 12 13 11 14 20 23 26 10 11 9 13 16 11 14 15 13
205 1 19 14 21 31 15 10 23 12 12 11 14 19 23 25 11 11 10 13 13 11 14 15 13
206 1 20 12 18 31 15 10 23 12 11 9 15 18 21 23 12 11 9 12 15 15 18 13 11
207 1 20 13 18 29 16 11 23 12 10 10 14 16 21 25 11 13 12 14 15 15 19 15 11
208 1 20 13 19 25 13 10 26 13 12 10 14 19 21 23 11 11 9 14 16 12 17 15 11
209 1 20 13 19 25 15 10 24 12 11 11 14 20 24 23 11 11 8 14 17 12 12 15 12
210 1 20 13 19 29 13 10 26 13 12 10 14 19 21 23 11 13 12 14 16 12 17 15 11
211 1 20 13 19 29 15 11 24 15 11 9 14 18 20 22 11 14.1 9 12 17 14 14 18 12
212 1 20 13 20 29 15 11 27 13 11 10 14 19 21 23 12 11 10 14 18 12 14 15 12
213 1 20 13 20 29 16 10 23 12 12 11 14 18 23 25 10 11 10 13 17 11 14 16 14
214 1 20 13 20 31 16 11 22 12 12 11 14 17 26 25 10 11 10 13 18 11 15 15 13
215 1 20 13 20 32 15 10 23 12 12 11 14 19 23 24 11 11 10 13 16 12 15 15 13
216 1 20 14 20 32 15 10 23 12 12 10 14 19 23 25 11 11 11 13 16 11 14 15 14
217 1 20 14 20 32 15 11 23 13 12 11 14 21 23 25 10 11 10 13 16 11 15 15 13
218 1 20 14 20 32 16 10 22 12 12 11 14 18 23 25 10 11 10 13 16 11 15 15 13
219 1 20 14 21 31 15 10 23 13 12 11 14 20 23 23 10 11 10 13 16 11 13 15 13
220 1 21 13 18 29 17 9 22 12 10 10 14 15 21 25 13 13 13 14 17 14 20 17 11
ANNALS OF HUMAN BIOLOGY

221 1 21 13 20 25 15 10 23 12 12 11 14 18 23 24 10 11 10 13 17 11 14 15 12
222 1 21 13 20 30 16 11 23 12 12 11 14 19 23 24 10 11 10 13 17 14 16 16 12
223 1 21 13 20 32 16 11 22 11 12 11 14 20 24 25 10 11 10 13 15.1 11 14 15 13
9
 10 M. PANDA ET AL.

Haplogroup O
West Bengal,

[Indian]

0.6086

0.0616
0.0073

0.3041
0.0974
0.0786

0.1311
0.5965
0.2223
0.0001
0.3912
India

0.067

Predominantly, the Haplogroup O has been seen in the East

0
0
0
Asian Y- gene pool and splits into two major subclades O1
and O2, together these occupy 60% of the total East Asian
India [Indian] India [Indian]
Uttar Pradesh, Uttarakhand,

0.0717
0.4762

0.0006
0.0033

0.3151
0.5367

0.5165
0.6886
0.5619

0.5387
0.2903
0.6867
0.0001

0.0006
Y-gene pool (Shi et al. 2005, p. 122; Yan et al. 2011, p. 4;

Zhong et al. 2011; Wang and Li 2013). We observed three
0
Table 4. Analysis of molecular variance pairwise distances based on Rst values between the studied population (Nayagarh, Odisha, India) and selected populations used for comparison from YHRD.

subclades among the studied population, viz. O (among 2

individuals i.e. 0.9%), O1 (among 23 individuals i.e. 10.9%)
0.0001

0.0002

0.0011

0.0015

0.0848
0.1051
and O2 (among 5 individuals, i.e. 2.4%), which together con-

0
0

0
0
0
0

0
0

0
tributed 14.2% of total haplogroup. The subclade O1 was the
third major haplogroup found among the
Maharashtra, Rajasthan, Tamil Nadu,

studied population.
0.1978
0.3403

0.0024
0.0016
0.0075
0.1505

0.2463
0.7844

0.2766
0.0947

0.0993
0.0144
0.0159
[Indian]

0.159

0.456
India

Haplogroup L
0.0015

0.0201
0.0042

0.0742
0.0296
0.0001

0.1887

0.0234
0.0831

0.0051
[Indian]

0.003
India

The haplogroup L is linked to the process of expansion of

0

0
0
0

agriculture associated with human evolution (Thanseem

et al. 2006). Some studies linked this haplotype as the ori-
0.0388

0.0555
0.0686

0.4152
0.4197
0.6312

0.0055
0.0086
0.0504
0.0056
0.0201
[Indian]
0.021
India

ginal founding member of the great Indus Valley civilisation

0
0
0

(McElreavey and Quintana-Murci 2005; Sengupta et al. 2006).

From the distribution point of view, the L haplogroup is
Madhya Pradesh,
India [Indian]

widely dispersed in the North Eurasian region as well as

0.0002
0.0048

0.0481
0.0178

0.4078
0.8156

0.0042
0.0146
0.0025
0.0751
0.0036
0.0187

among selected Siberian groups, whereas lower frequency is

0
0
0

seen among East Asians, populations from the Middle East,

and among Europeans (Karafet et al. 2002, 2008). A study
Jharkhand, Karnataka,

indicated that the frequency of haplogroup L gradually

0.1937
0.2149

0.0001
0.0005
0.0004
0.2931
0.0538

0.6334

0.0014
0.0233
0.0202
0.1061
0.0105
0.0265
[Indian]

0.008
India

increases from North to South in the Indian continent (Singh

et al. 2018). In the studied population, haplotype L was
observed among 10 individuals (4.7%), whose ancestry might
0.0924
0.2182

0.1213

0.0002

0.0013
0.0147
0.0116
0.0949
0.0052
0.0049
[Han] [Bhotra] [Indian] India [Indian] [Indian]

0.017
India

0.01

be linked to the founding members of the great Indus Valley

0
0
0

civilisation.
India and Kashmir,

0.0052

0.0001
0.0008

0.0218

0.0578
0.0352
0.0339

0.0295
0.0498
0.0229
0.0852
0.0057
0.0711
Jammu

0.001

Haplogroup C
0

Haplogroup C is a non-African haplogroup, highly concen-

Odisha, Assam,

0.0235
0.1729

0.0615
0.0437

0.0574

0.0266
0.0056
0.0255

0.0382
0.0334
0.0279
0.1612

0.0419
0.005

trated in Asia and Oceania (Wells et al. 2001; Lell et al. 2002;


0

Hammer et al. 2006; Regueiro et al. 2006; Hudjashov et al.

0.0296 0.1018
0.0664 0.1484

0.0882 0.0995 0.1253

0.0593 0.0646 0.0705
0.0966 0.0991 0.1058

0.1633

0.0676
0.2505
0.1097
0.1665

2007). It is less frequently found in Europe and America,

India

0.149

whereas it is completely absent in the African region

0
0

0
0

(Underhill et al. 2000; Cruciani et al. 2002; Lell et al. 2002;

0.1239

0.1062
0.1123

0.2216
0.0512
0.1274
China

0.082

p values are shown above the diagonal and Rst values below it.
0.07

Zegura et al. 2004; Singh Malhi et al. 2008). In the present

0
0

study, the subclade C2 was observed among 9 individuals

Beijing,

0.0092
0.1151
0.0209
0.0795

0.1019
0.1103
0.0711
0.2297
0.0561
0.1235
Bangladesh China
[Bangladeshi] [Han]

(4.3%). The distribution of subclade C2 was restricted to

0
0

Oceania and its neighbouring areas, except the Australian

0.0827
0.0813
0.0938
0.0104
0.0462

0.0063

0.0129

0.0194

0.0029
0.1093

0.0059

continent (Zhong et al. 2010). This indicated the patrilineal

Dhaka,

0.006

0.012

0.002

affinity of the studied population to Oceania and its sur-

0

rounding region in their evolutionary history.

Nayagarh,
Odisha,

0.0925
0.0933
0.0973
0.0309
0.0542

0.0068
0.0165

0.0212
0.0307
0.0121

0.0157
0.0202
India

0.013

0.086
0.02

Haplogroup J

Uttar Pradesh, India [Indian]

West Bengal, India [Indian]

Historically, the Indian landmass encountered different demic

Maharashtra, India [Indian]

Uttarakhand, India [Indian]

Tamil Nadu, India [Indian]
Jharkhand, India [Indian]
Karnataka, India [Indian]

Rajasthan, India [Indian]

Nayagarh, Odisha, India

diffusion events from West Asia as well as other regions of

Madhya Pradesh, India
Odisha, India [Bhotra]

Jammu and Kashmir,

Assam,India [Indian]
Beijing, China [Han]
Dhaka, Bangladesh

the world, that probably incorporated J2 haplotype in the

[Bangladeshi]

India [Indian]

continent (Sahoo et al. 2006; Vahia et al. 2017). Literature

China [Han]
Population

[Indian]

also suggests that the two subclades J2a and J2b are inter-
linked with the demic diffusion of cultivators in Eurasia and
North Africa from the Mesopotamia region (i.e. Syria and
 ANNALS OF HUMAN BIOLOGY 11

Figure 2. Neighbor-joining (NJ) tree based on pairwise Rst values between the studied population and compared populations from YHRD.

Figure 3. Principal coordinate analysis (PcoA) plot based on pairwise Fst values between the studied population and compared populations generated after output
of YHRD analysis.

Iraq) during the Neolithic period (Al-Zahery et al. 2003; Di observed among 3 individuals (1.4%) in the present
Giacomo et al. 2004; Semino et al. 2004). Among the current investigation.
study population, the two subclades J2a and J2b were asso-
ciated with 5 (2.4%) and 8 (3.8%) individuals, respectively.

Haplogroup I
Haplogroup I is a European type haplogroup, originating
before the beginning of the last glacial phase (International
Society of Genetic Genealogy 2013). The subclade I2a was
Haplogroup Q
Studies indicate that the haplogroup Q is seen in low fre-
quency among the population of Pakistan and India, and
absent in the South Indian region (Karafet et al. 2008; Singh
et al. 2018). Though the studied population was confined to
12 M. PANDA ET AL.

Figure 4. Heat map based on pairwise Fst values between the studied population and compared populations generated after output of YHRD analysis.

Figure 5. Haplogroup distribution among the male population of Nayagarh, Odisha obtained using Whit Athey’s Y-DNA Haplogroup Predictor.

Figure 6. Frequencies of observed subclades of different Y-chromosomal haplogroups.
the Eastern-Indian region, haplotype Q was seen with low
frequency, associated with only 3 individuals (1.4%).
Haplogroup G
Findings of a previous study specified that the subclade G2a
is absent among the population of Eastern India (Singh et al.
2018). But the present study reported the presence of G2a in
the studied population with very low frequency i.e. seen
among 2 individuals (0.9%).
Haplogroup E
The two subclades of Haplogroup E, E1b1a and E1b1b were
found among 2 individuals each and together contributed
1.8% of the total haplogroup.
Haplogroup T
This haplogroup was earlier recognised as Haplogroup K2.
Haplogroup T is distributed in Eurasia and North America,
Near East, Arabia (especially Iraq), northeastern Africa and
the Horn of Africa region. Among the southern and eastern
Indians, the haplogroup T has been observed with a fre-
quency of <14% (Sahoo et al. 2006; Sengupta et al. 2006). In
the population of Nayagarh, Odisha, we reported the hap-
logroup T as the least frequent haplogroup, seen in 1 indi-
vidual only (0.5%).
ANNALS OF HUMAN BIOLOGY 13
Discussion
In the present study, a total of 236 unrelated males from
Nayagarh, Odisha, were genotyped for 23 Y-STRs using
PowerPlexR Y23 System (Promega). A total of 44 different
alleles were observed with GD values that ranged from
0.346303642264695 to 0.943263430048837 and with DC of
0.945. The phylogenetic and inter-population analyses indi-
cated that the close genomic affinities existed between the
studied population and the Indian population of Jharkhand
and Madhya Pradesh as well as with the Bangladeshi popula-
tion of Dhaka. The principal haplogroups of the studied
population were reported as R1a, H1 and O1. The data gath-
ered in the present study will be useful for different statis-
tical calculations and frequency estimation for enquiries
related to forensic case-work. The present work is the first
comprehensive study of the male population of Odisha state
with extensive 23 Y-STRs. Our study findings indicate that
the Y-STR panel used could be efficient in population genetic
analysis, anthropological studies, and for forensic case work
among the studied population. The study outcomes proved
the efficiency of using a multiplex system for forensic, popu-
lation genetic and anthropological studies for the present
population. This study also enriches the existing Indian Y-
chromosome haplotype database.
Acknowledgement
We are thankful to the volunteer donors for providing the blood sam-
ples for the present study and to Promega (India) for providing multi-
plex kit used in the study.
14 M. PANDA ET AL.
Ethical approval
Prior approval regarding the study protocol, was received from the
Institutional Ethics Committee of Dr. H. S. Gour Vishwavidyalaya (A
Central University), Sagar M.P., India, vide letter no. DHSGV/IEC/2021/19/
1 dated 10.08.2021. Simultaneously, the ethical norms of the declaration
of Helsinki (World Medical Association) (Rickham 1964) were followed
during the present study.
Disclosure statement
No potential conflict of interest was reported by the author(s).
Funding
The author(s) reported there is no funding associated with the work fea-
tured in this article.
ORCID
Ramkishan Kumawat http://orcid.org/0000-0002-7794-9615
Gyaneshwer Chaubey http://orcid.org/0000-0003-2899-3852
Pankaj Shrivastava http://orcid.org/0000-0002-1647-2390
References
Adnan A, Ralf A, Rakha A, Kousouri N, Kayser M. 2016. Improving empir-
ical evidence on differentiating closely related men with RM Y-STRs: a
comprehensive pedigree study from Pakistan. Forensic Sci Int Genet.
25:45–51.
Alam S, Ali ME, Ferdous A, Hossain T, Hasan MM, Akhteruzzaman S.
2010. Haplotype diversity of 17 Y-chromosomal STR loci in the
Bangladeshi population. Forensic Sci Int Genet. 4(2):e59–e60.
Al-Zahery N, Semino O, Benuzzi G, Magri C, Passarino G, Torroni A,
Santachiara-Benerecetti A. 2003. Y-chromosome and mtDNA polymor-
phisms in Iraq, a crossroad of the early human dispersal and of post-
Neolithic migrations. Mol Phylogenet Evol. 28(3):458–472.
Athey TW. 2005. Haplogroup prediction from Y-STR values using an
allele-frequency approach. J Genet Geneal. 1:1–7.
Athey TW. 2006. Haplogroup prediction from Y-STR values using a
Bayesian-allele-frequency approach. J Genet Geneal. 2(2):34–39.
Ballantyne KN, Kayser M. 2012. Additional Y-STRs in forensics: why,
which, and when. Forensic Sci Rev. 24(1):63–78.
Basu A, Mukherjee N, Roy S, Sengupta S, Banerjee S, Chakraborty M, Dey
B, et al. 2003. Ethnic India: a genomic view, with special reference to
peopling and structure. Genome Res. 13(10):2277–2290.
Behura NK, Mohanty KK. 2006. Readings in social anthropology. India:
Dominant.
Bhasin MK, Walter H. 2001. Genetics of castes and tribes of India. India:
Kamla-Raj Enterprises.
Carracedo A,
Butler JM, Gusm~ao L, Linacre A, Parson W, Roewer L,
Schneider PM. 2013. New guidelines for the publication of genetic
population data. Forensic Sci Int Genet. 7(2):217–220.
Cruciani F, Santolamazza P, Shen P, Macaulay V, Moral P, Olckers A,
Modiano D, et al. 2002. A back migration from Asia to sub-Saharan
Africa is supported by high-resolution analysis of human Y-chromo-
some haplotypes. Am J Hum Genet. 70(5):1197–1214.
Debnath M, Palanichamy MG, Mitra B, Jin J-Q, Chaudhuri TK, Zhang Y-P.
2011. Y-chromosome haplogroup diversity in the sub-Himalayan Terai
and Duars populations of East India. J Hum Genet. 56(11):765–771.
Di Giacomo F, Luca F, Popa L, Akar N, Anagnou N, Banyko J, Brdicka R,
et al. 2004. Y chromosomal haplogroup J as a signature of the post-
neolithic colonization of Europe. Hum Genet. 115(5):357–371.
Dogan S, Babic N, Gurkan C, Goksu A, Marjanovic D, Hadziavdic V. 2016.
Y-chromosomal haplogroup distribution in the Tuzla Canton of Bosnia
and Herzegovina: A concordance study using four different in silico
assignment algorithms based on Y-STR data. Homo. 67(6):471–483.
Emmerova B, Ehler E, Comas D, Votrubova J, Vanek D. 2017. Comparison
of Y-chromosomal haplogroup predictors. Forensic Sci Int: Genet
Suppl Ser. 6:e145–e147.
Excoffier L, Lischer HE. 2010. Arlequin suite ver 3.5: a new series of pro-
grams to perform population genetics analyses under Linux and
Windows. Mol Ecol Resour. 10(3):564–567.
Excoffier L, Smouse PE, Quattro JM. 1992. Analysis of molecular variance
inferred from metric distances among DNA haplotypes: application to
human mitochondrial DNA restriction data. Genetics. 131(2):479–491.
Ghosh A. 1990. An Encyclopaedia of Indian Archaeology. Netherlands:
BRILL.
Gusmao L, Butler J, Carracedo A, Gill P, Kayser M, Mayr W, Morling N,
International Society of Forensic Genetics, et al. 2006. DNA
Commission of the International Society of Forensic Genetics (ISFG):
an update of the recommendations on the use of Y-STRs in forensic
analysis. Int J Legal Med. 120(4):191–200.
Hammer Ø, Harper DA, Ryan PD. 2001. PAST: paleontological statistics
software package for education and data analysis. Palaeontol
Electronica. 4(1):9.
Hammer MF, Karafet TM, Park H, Omoto K, Harihara S, Stoneking M,
Horai S. 2006. Dual origins of the Japanese: common ground for
hunter-gatherer and farmer Y chromosomes. J Hum Genet. 51(1):
47–58.
Haplogroup T-M184 j Project Gutenberg Self-Publishing - eBooks j Read
eBooks online. [accessed 2021 Apr 22]. http://self.gutenberg.org/art-
icle/whebn0009238638/haplogroup%20t-m184.
Heraclides A, Bashiardes E, Fern
andez-Dom
ınguez E, Bertoncini S,
Chimonas M, Christofi V, King J, et al. 2017. Y-chromosomal analysis
of Greek Cypriots reveals a primarily common pre-Ottoman paternal
ancestry with Turkish Cypriots. PLoS One. 12(6):e0179474.
Hudjashov G, Kivisild T, Underhill PA, Endicott P, Sanchez JJ, Lin AA,
Shen P, et al. 2007. Revealing the prehistoric settlement of Australia
by Y chromosome and mtDNA analysis. Proc Natl Acad Sci. 104(21):
8726–8730.
Iacovacci G, D’Atanasio E, Marini O, Coppa A, Sellitto D, Trombetta B,
Berti A, Cruciani F. 2017. Forensic data and microvariant sequence
characterization of 27 Y-STR loci analyzed in four Eastern African
countries. Forensic Sci Int Genet. 27:123–131.
Imam J, Reyaz R, Singh RS, Bapuly AK, Shrivastava P. 2018. Genomic por-
trait of population of Jharkhand, India, drawn with 15 autosomal STRs
and 17 Y-STRs. Int J Legal Med. 132(1):139–140.
Indian Genome Variation Consortium. 2008. Genetic landscape of the
people of India: a canvas for disease gene exploration. Journal of
Genetics. 87:3–20.
International Society of Genetic Genealogy. 2013. Y-DNA Haplogroup
Tree 2013, Version: 8.89 Date: 31 December 2013 [accessed 2021 Apr
22]. http://www.isogg.org/tree/.
Jobling MA, Tyler-Smith C. 2000. New uses for new haplotypes the
human Y chromosome, disease and selection. Trends Genet. 16(8):
356–362.
Karafet TM, Mendez FL, Meilerman MB, Underhill PA, Zegura SL, Hammer
MF. 2008. New binary polymorphisms reshape and increase resolution
of the human Y chromosomal haplogroup tree. Genome Res. 18(5):
830–838.
Karafet TM, Osipova LP, Gubina MA, Posukh OL, Zegura SL, Hammer MF.
2002. High levels of Y-chromosome differentiation among native
Siberian populations and the genetic signature of a boreal hunter-
gatherer way of life. Hum Biol. 74(6):761–789.
Kruskal JB. 1964. Nonmetric multidimensional scaling: a numerical
method. Psychometrika. 29(2):115–129.
Kumar A, Kumar R, Kumawat R, Mathur B, Shrivastava P, Chaubey G,
Yadav RK. 2020. Genetic portrait study for 23 Y-STR loci in the popu-
lation of Rajasthan, India. Int J Legal Med. 134(5):1691–1693.
Kumawat RK, Shrivastava P, Shrivastava D, Mathur GK. 2020. Molecular
diversity of 23 Y-STR genetic markers in the population of Rajasthan,
India. Meta Gene. 24:100694.
Kwak KD, Jin HJ, Shin DJ, Kim JM, Roewer L, Krawczak M, Tyler-Smith C,
Kim W. 2005. Y-chromosomal STR haplotypes and their applications
to forensic and population studies in east Asia. Int J Legal Med.
119(4):195–201.

Lang M, Liu H, Song F, Qiao X, Ye Y, Ren H, Li J, et al. 2019. Forensic
characteristics and genetic analysis of both 27 Y-STRs and 143 Y-SNPs
in Eastern Han Chinese population. Forensic Sci Int Genet. 42:
e13–e20.
Lell JT, Sukernik RI, Starikovskaya YB, Su B, Jin L, Schurr TG, Underhill PA,
Wallace DC. 2002. The dual origin and Siberian affinities of Native
American Y chromosomes. The. Am J Hum Genet. 70(1):192–206.
Li L, Yu G, Li S, Jin L, Yan S. 2016. Genetic analysis of 17 Y-STR loci from
1019 individuals of six Han populations in East China. Forensic Sci Int
Genet. 20:101–102.
Lowery RK, Herrera K, Uribe G, Reguiero M, Herrera RJ. 2013. Sub-popula-
tion structure evident in forensic Y-STR profiles from Armenian geo-
graphical groups. Leg Med. 15(2):85–90.
Majumder PP. 1998. People of India: biological diversity and affinities.
Evol Anthropol. 6(3):100–110.
McElreavey K, Quintana-Murci L. 2005. A population genetics perspective
of the Indus Valley through uniparentally-inherited markers. Ann Hum
Biol. 32(2):154–162.
Nayagarh District Government of Odisha j District Administration Portal j
India. [accessed 2021 Apr 16]. https://nayagarh.nic.in/.
Nayagarh District Population Census 2011–2021. Orissa literacy sex ratio
and density [accessed 2021 Apr 16]. https://www.census2011.co.in/
census/district/409-nayagarh.html.
Nothnagel M, Fan G, Guo F, He Y, Hou Y, Hu S, Huang J, et al. 2017.
Revisiting the male genetic landscape of China: a multi-center study
of almost 38,000 Y-STR haplotypes. Hum Genet. 136(5):485–497.
Oppenheimer S. 2012. Out-of-Africa, the peopling of continents and
islands: tracing uniparental gene trees across the map. Philos Trans R
Soc Lond B Biol Sci. 367(1590):770–784.
Papiha SS. 1996. Genetic variation in India. Human Biol. 68(5):607–628.
Petrejc
ıkov
a E, Carnogursk
a J, Hronsk
a D, Bernasovsk
a J, Boronov
a I,
Gabrikov
a D, Bo ^zikov
a A, Macekov
a S. 2014. Y-SNP analysis versus Y-
haplogroup predictor in the Slovak population. Anthropol Anz. 71(3):
275–285.
Prinz M, Boll K, Baum H, Shaler B. 1997. Multiplexing of Y chromosome
specific STRs and performance for mixed samples. Forensic Sci Int.
85(3):209–218.
Quintana-Murci L, Krausz C, McElreavey K. 2001. The human Y chromo-
some: function, evolution and disease. Forensic Sci Int. 118(2–3):
169–181.
Regueiro M, Cadenas AM, Gayden T, Underhill PA, Herrera RJ. 2006. Iran:
tricontinental nexus for Y-chromosome driven migration. Hum Hered.
61(3):132–143.
Rickham P. 1964. Human experimentation. Code of ethics of the world
medical association. Declaration of Helsinki. Br Med J. 2(5402):
177–177.
Roewer L, Andersen MM, Ballantyne J, Butler JM, Caliebe A, Corach D,
D’Amato ME, et al. 2020. DNA commission of the International
Society of Forensic Genetics (ISFG): recommendations on the inter-
pretation of Y-STR results in forensic analysis. Forensic Sci Int Genet.
48:102308.
Roewer L, Croucher PJ, Willuweit S, Lu TT, Kayser M, Lessig R, de Knijff P,
Jobling MA, et al. 2005. Signature of recent historical events in the
European Y-chromosomal STR haplotype distribution. Hum Genet.
116(4):279–291.
Sahoo S, Singh A, Himabindu G, Banerjee J, Sitalaximi T, Gaikwad S,
Trivedi R, et al. 2006. A prehistory of Indian Y chromosomes: evaluat-
ing demic diffusion scenarios. Proc Natl Acad Sci. 103(4):843–848.
Satapathy KC. 2018. Human diversity study in Odisha: an overview.
Education. 6(1):6–14.
Semino O, Magri C, Benuzzi G, Lin AA, Al-Zahery N, Battaglia V, Maccioni
L, Triantaphyllidis C, et al. 2004. Origin, diffusion, and differentiation
of Y-chromosome haplogroups E and J: inferences on the neolithiza-
tion of Europe and later migratory events in the Mediterranean area.
Am J Human Genet. 74(5):1023–1034.
ANNALS OF HUMAN BIOLOGY 15
Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow C-ET,
Lin AA, et al. 2006. Polarity and temporality of high-resolution y-
chromosome distributions in India identify both indigenous and
exogenous expansions and reveal minor genetic influence of Central
Asian pastoralists. Am J Hum Genet. 78(2):202–221.
Shi H, Dong Y, Wen B, Xiao C-J, Underhill PA, Shen P, Chakraborty R,
et al. 2005. Y-chromosome evidence of southern origin of the East
Asian-specific haplogroup O3-M122. Am J Hum Genet. 77(3):408–419.
Shrivastava P, Jain T, Trivedi VB. 2017. Haplotype data for 17 Y-STR loci
in the population of Madhya Pradesh. India. Forensic Science
International: Genetics. 26:e31–e32.
Singh M, Sarkar A, Nandineni MR. 2018. A comprehensive portrait of Y-
STR diversity of Indian populations and comparison with 129 world-
wide populations. Sci Rep. 8(1):15421.
Singh Malhi R, Gonzalez-Oliver A, Schroeder KB, Kemp BM, Greenberg
JA, Dobrowski SZ, Smith DG, et al. 2008. Distribution of Y chromo-
somes among native North Americans: a study of Athapaskan popula-
tion history. Am J Phys Anthropol. 137(4):412–424.
Srivastava A, Kumawat R, Dixit S, Kaitholia K, Shrivastava D, Yadav VK,
Nigam K, et al. 2019. Genetic data for PowerPlex 21TM autosomal and
PowerPlex 23 Y-STRTM loci from population of the state of Uttar
Pradesh, India . Int J Legal Med. 133(5):1381–1383.
Thanseem I, Thangaraj K, Chaubey G, Singh VK, Bhaskar LV, Reddy BM,
Reddy AG, Singh L. 2006. Genetic affinities among the lower castes
and tribal groups of India: inference from Y chromosome and mito-
chondrial DNA. BMC Genet. 7(1):1–11.
Trivedi R, Sahoo S, Singh A, Bindu GH, Banerjee J, Tandon M, Gaikwad S,
et al. 2008. Genetic imprints of pleistocene origin of indian popula-
tions: a comprehensive phylogeographic sketch of Indian Y-chromo-
somes. Int J Human Genet. 8(1–2):97–118.
Underhill PA, Myres NM, Rootsi S, Metspalu M, Zhivotovsky LA, King RJ,
Lin AA, et al. 2010. Separating the post-Glacial coancestry of
European and Asian Y chromosomes within haplogroup R1a. Eur J
Hum Genet. 18(4):479–484.
Underhill PA, Shen P, Lin AA, Jin L, Passarino G, Yang WH, Kauffman E,
et al. 2000. Y chromosome sequence variation and the history of
human populations. Nat Genet. 26(3):358–361.
Vahia MN, Yadav N, Ladiwala U, Mathur D. 2017. A diffusion based study
of population dynamics: prehistoric migrations into South Asia. PLOS
One. 12(5):e0176985.
Wang C-C, Li H. 2013. Inferring human history in East Asia from Y chro-
mosomes. Investig Genet. 4(1):11.
Wells RS, Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva I, Blue-Smith
J, Jin L, et al. 2001. The Eurasian heartland: a continental perspective
on Y-chromosome diversity. Proc Natl Acad Sci. 98(18):10244–10249.
Yan S, Wang C-C, Li H, Li S-L, Jin L, Ziegle J, Swamikrishnan P, The
Genographic Consortium, et al. 2011. An updated tree of Y-chromo-
some haplogroup O and revised phylogenetic positions of mutations
P164 and PK4. Eur J Hum Genet. 19(9):1018–1018.
Zegura SL, Karafet TM, Zhivotovsky LA, Hammer MF. 2004. High-reso-
lution SNPs and microsatellite haplotypes point to a single, recent
entry of native American Y chromosomes into the Americas. Mol Biol
Evol. 21(1):164–175.
Zeng Z, Rowold DJ, Garcia-Bertrand R, Calderon S, Regueiro M, Li L,
Zhong M, Herrera RJ. 2014. Taiwanese aborigines: genetic heterogen-
eity and paternal contribution to Oceania. Gene. 542(2):240–247.
Zhong H, Shi H, Qi X-B, Duan Z-Y, Tan P-P, Jin L, Su B, Ma RZ. 2011.
Extended Y chromosome investigation suggests postglacial migra-
tions of modern humans into East Asia via the northern route. Mol
Biol Evol. 28(1):717–727.
Zhong H, Shi H, Qi X-B, Xiao C-J, Jin L, Ma RZ, Su B. 2010. Global distri-
bution of Y-chromosome haplogroup C reveals the prehistoric migra-
tion routes of African exodus and early settlement in East Asia. J Hum
Genet. 55(7):428–435.