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Anthropometric and genetic distance between Gonds of Central India.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 65:291-304 (1984)
Anthropometric and Genetic Distance Between Gonds
of Central India
URMILA PINGLE
Indian Statistical Institute, Calcutta-700 035, India
KEY WORDS Anthropometric measurements, Genetic polymorphisms, Genetic distance, Cluster analysis
ABSTRACT
A morphological and genetic study was undertaken on five
Gondi-speaking populations of Central India (Andhra Pradesh and Maharashtra States). There has been no systematic biological study on this large Dravidian-speaking tribal group, 4 million in number, amounting to 13% of the total
tribal population of India. Data was collected on 16 anthropometric measurements and seven genetic markers (blood groups, hemoglobin, G6PD and plasma
protein polymorphisms) on the Raj Gonds, Kolams, Manne, Koyas and Plains
Maria Gonds. Various genetic distance measures such as Mahalanobis’s D2
and Nei’s and Sanghvi’s measures and cluster analysis techniques were used
to determine the relationship between these groups based on anthropometrics
and genetic variables. The statistical analysis revealed the Gonds to be a
heterogenous group in both morphology and genetic characteristics. The morphological and genetic distances between these five groups when projected
graphically revealed that the spatial distribution of these Gonds generally
corresponds to their present geographical distribution. However, the actual
relationships among each of the Gond populations show differences when based
on these two biological variables, the possible reasons for this being discussed
in the paper. The emphasis of this study is on the importance of geographical
proximity in producing morphological and genetic similarity between populations, brought about by a short distance a s well as similar geographical factors
(such as soil, terrain, flora, etc.) drawing these populations together under a
common ecocultural umbrella.
Gonds are one of the largest tribal populations of Central India, roughly 4 million in
number, amounting to about 13% of the total
tribal populations of India. “Gondwana” the
area of their distribution stretches into Madhya Pradesh, Maharashtra, and Andhra Pradesh States. The Gondi language belongs to
the Dravidian group of languages. The Gonds
had kingdoms in Deogarh on the foothills of
Satpuras at Mandla (Madhya Pradesh) and
Chanda (Maharashtra) from the 14th to the
18th centuries AD until they were defeated
by the Marathas. The social and anthropological aspects of the Gonds of Andhra Pradesh have been studied over a period of 40
years by Furer-Haimendorf (1979) who considers the Gonds neither racially, culturally,
nor linguistically a homogenous group. However, there has been no systematic biological
0 1984 ALAN R. LISS, INC.
study on Gonds and the relationship betwen
various subsections of this large linguistic
tribal group. Hence, the present study was
undertaken on five major Gondi-speaking
populations of Andhra Pradesh and Maharashtra.
AREA OF DISTRIBUTION AND BACKGROUND
OF GOND POPULATIONS
The area of study lies between 18”40’and
19”40’ north and 78’45’ and 80’45’ east,
approximately (Fig. 1). The geology and
hence the physical characteristics of the soil
differ from west to east; the western portion
Received July 7, 1982; accepted June 27,1984.
Urmila Pingle’s present address is 1-69, Habshiguda, Hyderabad-500 007, Andhra F’radesh, India.
-7-
-Rivers
2
4
6
8
Toluk
vllloge
and s t r e o m s
R o i i w o y line
Plains Moria Gond a n d
H i l l Marro Gand village
M
A
Koyo v i l \ a g e ( M i x e d )
Plains M o r i a Gond v i l l a g e
M a n n e a n d Koya v i i l o g e [ M i x e d )
K o y o willoge
Manne
RojGond o n d K o l o m v i l l a g e ( M i x e d )
RojGond villoge
Kolom village
headquarlers
K E Y
MI LES
L
l
l
i
l
0
Taluk
Tolu k
Adllobob
U t nur
H
R
S
T
U
A
2
4
6
8
MILES
LLLU
0
Lokshetf pet Toluk
A
S
A
T
E
Sultona b o d
T
Sirpur
Monthint
Toluk
ond Chcnnur Toiukr
M AMARASTRA
Fig. 1. Map showing distribution of all Gond villages (tribewise).
A
4s1fobod Toluk
Sironcha Toluk
SO0
ulLC1
u
293
ANTHROPOMETRIC AND GENETIC DISTANCE AMONG GONDS
of the field area is mountainous and covered
by black cotton soil of Deccan trap. The tribal
populations selected here are the Raj Gonds
and Kolams of Utnoor and Asifabad taluks*
(Andhra Pradesh). Though these two populations live in close proximity to each other
they have distinct geomorphological territories. The Kolams, who until recently were
shifting cultivators, live on the hilltops where
the soil is thin and suitable only for cultivation of primitive millets. The Raj Gonds however cultivate the valley bottoms near
streams where there is a thick layer of black
cotton soil suitable for jowar and cotton cultivation with a plough. The total Kolam population in Adilabad district is about 9,386
whereas the Raj Gonds have a total population of 89,150.
East of these trap hills, the plains of Gondwana sedimentary soils form the valley of
Pranitha and Godavari rivers. The predominant tribal populations of these soils, the
Koyas and Manne, belonging to Chinnur taluk (Andhra Pradesh), were selected for the
study. The Koyas and Manne are approximately 7,000 each in both taluks. East of the
river Prahnitha and mainly in the valley of
the Indravati river live the main subgroup of
Gonds, the Marias, Mwias, and Bisonhorn
Marias. This is a country of dense forests and
plains and hills. The sample population selected are the Maria Gonds living mainly in
the Sironcha taluk of Chanda district (Maharashtra). This group is divided into two
subgroups, the Plains Maria and Hill Maria
Gonds. The Hill Marias have a low population density of four persons per km2 compared to the Plains Maria Gonds who are
more numerous, their population density
reaching 14 persons per km2 (Census of India, Maharashtra State, 1961),having a total
population of 33,189.
BACKGROUND OF THE GONDS
Raj Gonds
The basis of social structure of the Raj
Gonds is a system of four exogamous phratries (saga) each of which is subdivided into
a number of named clans (pari) (F'urer-Haimendorf, 1979). The most distinctive feature
of the Gond religion is the cult of the phratry
and clan deities, described by the generic
term Persa pen or Great God. The frequency
of consanguineous marriages is high as referred to in Table 1(Pingle, 1981).
Kolams
Most Kolams speak Kolami, which is also
a Dravidian language closely related to
Gondi. In their own language the Kolams
refer to themselves as Kolavar but in Gondi
they are called Pujari Purer-Haimendorf,
1979).The Kolams also speak in Gondi to the
Gonds and Pardhans (bards to the Gonds).
The Kolams, like the Gonds, are organized
into exogamous clans and most of these have
names identical with those of certain Gond
clans. These clans are grouped in phratries
corresponding to the Gond system. Kolams
and Gonds consider themselves related communities and Kolams eat freely in the houses
of Gonds and many Gonds partake without
hesitation of the food of the Kolams (FurerHaimendorf, 1979). The data on consanguineous marriages and marriage distance is
given in Table I.
Manne
The Manne are a Telugu-speaking subgroup
of the Kolams and have forgotten Kolami under the dominance of Telugu culture and have
now become a separate endogamous group,
not intermarrying with the Kolams. They
have a marriage system similar to that practiced by the rest of the Gonds.
Koyas
The Koyas speak Koye, a language described by Grierson (1906) as a dialect of
Gondi. Most of them however have now forgotten the language and speak Telugu, which
they have adopted from their neighbors. They
have a phratry system similar to that of the
TABLE 1. Consanguinity and marriage distance pattern among gonds
Tribes
Kolam
Koya
Manne
Plains Maria
Gonds
Raj Gonds
Total
marriages
Consanguineous
marriages (%)
Average
inbreeding
coeffkient
Marriage distance (km)
Village
endogomy
Mean f SE
Median
(%I
129
180
136
116
88 (68.2)
52 (28.9)
39 (28.7)
32 (27.6)
0.043
0.018
0.018
0.017
17.65 f 1.47
9.62 f 0.77
8.62 k 0.91
13.66
1.36
13.42
6.86
4.40
9.44
29.8
36.7
45.1
27.2
131
39 (29.8)
0.019
14.42 +_ 1.97
5.29
39.1
294
U. PINGLE
Gonds and some of their clan Gods, which
are still in Bastar, are brought down on clan
festivals to the East Godavari district.
Maria Gonds
The Maria Gonds speak Gondi and are distributed in Sironcha taluk of Chanda district
(Maharashtra). A supplement on the Maria
Gonds of Chanda has been written by Grigson (1949).Their Bastar neighbors, called the
Marias, are a separate endogamous group.
Most of the Marias were practicing shifting
cultivation some 50 years ago and some of
them still do so on top of Abujhmar hills (Hill
Maria Gonds). They had a Gotul system,
which is a youth club for both unmarried
girls and boys now more or less disappeared
in Chanda but is still practiced among the
Marias of Bastar (Elwin, 1947). The Godgrouping appears similar to that of the Raj
Gonds and may be a recent importation to
the Maria Gonds of Chanda (Grigson, 1949).
However, the Bastar Marias and Murias
have neither God-grouping nor Chuddur
Penk.
MATERIALS AND METHODS
Data on anthropometric measurements and
blood samples were collected on 100 individuals consisting of 50 males and 50 females
from each of five Gond populations. In addition, a n extra 50 male samples for each of
the above groups was collected for the purpose of screening for G6PD deficiency and
hemoglobin and transferrin variants. Villages were selected to represent as far as
possible the different areas of distribution of
the tribe. Since most of the villages were
situated along the main streams, villages
were selected so as to represent each of the
main streams thus covering a representative
proportion of the tribal groups in the area
selected for study. All available adults in the
villages were covered avoiding as far as possible (as consanguinity was high in these
groups) relatives and rejecting members of
the same family. Information on age, marital
status, relatedness of spouse, place of birth of
spouse, etc., were also collected on each of
the individuals.
For each selected individual, after the measurements were taken, blood samples were
collected by means of finger prick into heparnized microcapillary tubes.
Biochemical methods
Microtechniques were followed for blood
grouping in ABO, MN, and RkO,Dsystems.
The antisera were obtained from HaMtins
Institute, Bombay, and the techniques followed were those advocated by this institute.
Screening for G6PD deficiency was done by
the brilliant cresyl blue (BLB) dye decolorization technique of Motulsky and CampbellKrant (1960).
Separation of serum proteins like haptoglobin, transferrin, albumin, and hemoglobin
were achieved by polyacrylamide electrophoresis. The method of Clark (1964)was adopted
for separation of serum proteins on a 7%
acrylamide gel. A 5.5% polyacrylamide gel
was used with the buffer system of Dietz et
a1 (1971) for hemoglobin separation. Sickle
cell test was carried out on a heterozygote
Hb AS sample using 2% sodium metabisulphite solution and then observing the red
cells under a microscope for sickling after
15-20 minutes.
Anthropometric techniques
Sixteen anthropometric measurements
were carried out and the methods followed
were according to Human biology, I.B.P.
Handbook No. 9, Guide to field methods (Weiner and Lourie, 1969).The following were the
measurements carried out: 1)stature, 2) upper arm length, 3) fore arm length, 4) bigonial
diameter, 5) morphological face height (nasion-gnathion), 6) nose height or length, 7)
nose breadth, 8)head length, 9) head breadth,
10)biocromial diameter, 11)bi-iliac diameter,
12) right upper arm circumference, 13)right
triceps skinfold, 14) right subscapular skinfold, 15)weight, and 16)bizygomatic breadth.
Bizygomatic breadth, however, was measured as the minimum distance between zygomatic arches rather than the maximum
distance as in I.B.P. methods.
STATISTICAL METHODS
Scrutiny of data
The scrutiny of raw data for the presence
of “outliers,” that is, observations that appear to be inconsistent with the remainder of
that set of data, is a n important and essential
step to valid analysis of the data. Since the
presence of these outliers is likely to distort
the inference process, as a first step, in order
to detect the outlier, a “descriptive analysis”
of the data was carried out. This involves 1)
frequency distribution in narrow class intervals and drawing histograms; 2) computing
measures of locations like mean, mode, median, and measures of dispersion; 3) recording maximum and minimum values; and 4)
ANTHROPOMETRIC AND GENETIC DISTANCE AMONG GONDS
computing measures of skewness and kurtosis.
Outliers were suspected if 1)there are gaps
in the histogram and the extreme observations are far removed from the nearest neighbors; 2) if the variation in some samples is
larger than in the others for the same character; 3) if there is a general consistency in
measures of skewness and kurtosis but in
some cases the values are discordant.
Transformation of a variable to make the
transformed variable more approximately
normal is generally advocated when there
are strong reasons to believe that the original variable is not inherently a normally distributed one. However, in this case, the
variables all represent standard physical
measurements that are known to be reasonably normally distributed and the observed
deviations from normality could therefore be
attributed to a small number of extreme values, i.e., outliers mostly due to recording errors (21 in a total of 10,000). This view is
further supported by the fact that, after deletion of these outliers, the variables under
consideration exhibited very closely the structure of normal distribution. In the case of
measurements such as skinfold thickness
measurements and weight, the distribution
to start with is not normal, the skewed distribution being due to inherent variability.
These measurements were not used for multivariate analysis.
PRELIMINARY ANALYSIS
Anthropometric data
Out of the 16 measurements only 10 were
selected for further analysis. Of the six that
were rejected, the upper arm and forearm
length are highly correlated with stature and
hence redundant, whereas the rest, consisting of soft tissue measurements like midarm
circumference, triceps skinfold, subcapular
skinfold, and weight showed a skewed distribution and therefore were not suitable for
multivariate analysis. Also, they are highly
affected by environmental factors and not
suitable for comparative analysis. The following tests were carried out on all measurements: 1)variances within tribes and tests of
homogeneity for both females and males; 2)
tests for homogeneity of a set of correlation
coefficients (Rao, 1965); 3) equality of variances and covariances for males and females
by likelihood ratio test and using x2 approximation according to Box (1949). Only the ten
measurements mentioned above were found
295
to have nearly the same variances for all the
groups.
A similar correlation matrix and the covariance matrices were found to be equal
within both male and female groups. Hence
only these measurements were used for multivariate analysis.
Analysis of variance was carried out on
each of the selected measurements between
female groups and male groups and for sexual dimorphism.
Genetic data
The gene frequency and tests of homogeneity and consistency were applied for seven
biochemical markers. Albumin protein was
found to be invariant and was not used for
comparative analysis. The gene frequency for
the ABO system was estimated by maximum
likelihood method, and tests for departure
from Hardy-Weinberg equilibrium and heterogeneity were carried out according to Rao
(1965). The test for consistency and homogeneity was also carried out on MN system
according to Rao (1965). x2 test for heterogeneity of gene frequencies between the five
Gond populations was carried out in the other
genetic markers.
Genetic distance measures
Mahalanobis’s D2:This is the best measure
for metric data provided that some assumptions are satisfied, the two most important
being homogeneity of the variances, and correlations and normality of the multivariate
distribution. This is given by
(Xj-Xk)
w-’(Xj-Xk) =
Dj2k (Mahalanobis’s D2)
where W is the pooled estimate of the within
population dispersion matrix common to the
jth and kth populations. D2 given above is a
measure of generalized “sum of squares of
differences” among the mean values,
weighed approximately to take into account
the internal correlations among the variables, and is a measure of overlap between
populations and is thus more appropriate to
study affinities or dissimilarities between individuals in different populations.
Nei’s distance: Nei (1972) taking into account the effect of genetic polymorphism
within populations defined the normalized
identity of genes between populations as
equivalent to protein identity. He then related it to the accumulated number of gene
296
U. PINGLE
differences per locus, now called genetic distance. This measure of genetic distance, according to Nei, has several advantages over
those proposed by Cavalli-Sforza and Edwards (1967), Balakrishnan and Sanghvi
(1968),and others: 1)It is related to Malecot’s
coefficient of kinship in a simple way; 2) it
measures the accumulated number of gene
substitutions per locus; 3) if the rate of gene
substitution per year is constant it is linearly
related to evolutionary time.
G2c of Balakrishnan and Sanghvi: The
same fundamental assumption in Mahalanobis’s D2 is followed, that is, though the populations may differ in their mean gene
frequencies all of them have the same dispersion matrices.
H-clust, a hierarchial clustering program
written by Stephen C. Johnson of the Bell
Telephone Laboratories, New Jersey, was
used to give clustering based on both these
techniques. The representations of these
analyses in the form of dendograms or trees
were made as they have a n advantge of being
readily interpretable as conventional taxonomic hierarchies. Subgraphs were projected
at different levels of clustering based on the
complete linkage method. This helps in visualizing the spatial distribution of the populations.
RESULTS
Morphology
The mean values for various measurements in male and female groups are given
Cluster analysis
in Tables 2 and 3. Analysis of variance
In many empirical fields, there is a n in- showed F values were high for eight of the
creasing interest in identifying the group- 11 measurements showing significant differings or clusterings of the object under study ences between the five tribal groups in both
that best represent certain empirically mea- sexes (Table 4).There was also a significant
sured relations of similarity. The problem difference between sexes within each tribal
then is discovering if there is any structure group for each of these 11 measurements.
(i.e., natural arrangement of the object into Except for head breadth, no other measurehomogenous groups) inherent in the data. ment showed a difference in sexual dimorThe two clustering techniques used in this phism between the five tribal groups.
study are 1) single linkage by Sneath and
x2 test for homogeneity on D2 matrices for
Sokal(l973)or minimum method by Johnson males and female Gonds (Table 5) showed
(1967) or 2) complete linkage or maximum that D2 values were significantly different
method (Johnson, 1967).
between the five groups in both sexes.
Single linkage method: In this method a n
Two clustering techniques, the single linkoperational taxonomic unit (OTU) that is a age and complete linkage methods were carcandidate for a n extant cluster has similarity ried out on D2 values and showed a near
to that cluster equal to its similarity to the correspondence in relationship between the
closest member within the cluster. These five Gond groups in both sexes (Figs. 2 and
connections between OTUs and clusters and 3). However, the relationship between males
between two clusters are established by sin- and females did not show close correspondgle links between pairs of OTUs. This proce- ence or consistency when either of the clusdure frequently leads to long, straggly tering techniques was followed. In the males,
the Koyas cluster with Manne and Raj
clusters.
Complete linkage method: This is the di- Gonds, the Kolams and Plains Maria Gonds
rect antithesis of the single linkage tech- joining separately in that order, whereas in
nique. An OTU that is a candidate for the females two clusters are formed, one conadmission to a n extant cluster has similarity sisting of Kolams and Manne and the other
to that cluster equal to its similarity to the of Koyas with Plains Maria Gonds. After
furtherest member within the cluster. Where these two clusters join each other, the Raj
two clusters join, their similarity is that ex- Gonds join separately. In both sexes the
isting between the furtherest pair or mem- Manne (MN) lie intermediate, connecting the
bers one in each cluster. The method will Kolams (KL1) and Raj Gonds (RG1) with
generally lead to tight, hyperspherical, dis- Koyas (KY1)and Plains Maria Gonds (PM).
crete clusters that join others only with diffi- The position of the Koyas (KY1) however is
culty and at related low overall similarity different in the two sexes shifting nearer to
values. Clusters by this method are compact Manne (MN) in the males and away from
and more structured, showing more taxa and Manne towards Plains Maria Gonds (PM) in
the females.
more ranks.
137.308
184.173
118.481
103.314 (-1)
109.442
49.288
1613.038
315.058
262.510 (-1)
351.615
252.471 (-1)
226.308
46.731
75.481
96.771
39.308
0.781
0.726
0.691
0.709
0.318
0.620
7.719
2.257
1.746
2.307
1.722
2.350
0.970
1.977
1.612
0.442
SE
Manne (56)
X
141.073 (- 1)
184.839
120.018
103.115 ( - 1)
111.673
49.607
1607.607
311.018
264.500
353.607
258.036
228.071
49.286
70.625
94.852
38.054
0.606
0.793
0.787
0.871
0.815
0.571
8.554
2.354
2.317
2.313
1.677
1.978
1.109
1.983
1.364
0.364
-
SE
Koya (51)
141.843
186.922
118.647
101.900
108.843
47.529
1593.118
306.608
255.961
357.078
256.569
236.000
49.118
75.588
98.708
38.196
X
-
0.602
0.889
0.767
0.729
0.819
0.624
7.633
. .~~
2.835
2.226
2.127
1.934
2.065
1.542
1.978
1.362
0.324
SE
142.185
186.241
118.019
104.906
110.963
47.944
1606.648
305.778
256.778
351.111
264.481
243.741
58.113 (-1)
86.226 (-1)
105.509
39.185
0.664
0.834
0.747
0.698
0.917
0.419
7.873
2.185
1.799
2.115
2.011
2.577
2.061
3.230
1.795
0.401
Plains Maria Gonds (54)
X
SE
-
139.481
187.148
119.259
102.096
110.736
47.815
1628.278
313.630
265.241
355.759
254.333
238.444
48.148
78.333
103.170
37.481
X
0.491
0.647
0.726
0.750
0.799
0.510
8.757
2.126
1.944
2.067
1.699
2.282
1.615
2.368
1.828
0.415
SE
Raj Gonds (54)
(xand SE) of anthropometric measurements after omitting outliers (number indicated in bracket)
135.260
177.340
113.280
96.043
103,120
44.560
36.580
1490.500
287.938 (-2)
235.959 (-1)
311.380
245.420
210.660
66.400
70.400
83.653
X
-
'Abbreviations as in footnote to Table 2.
UAL
LAL
BAB
BIB
MAC
TFF
SFF
Wt
St
HL
BzB
BgB
TFL
NL
NB
HB
Character
Kolam (50)
1.811
1.803
2.195
2.914
2.511
1.310
SE
0.763
0.884
0.682
0.696
0.707
0.476
0.301
6.646
2.102
1.880
X
137.160
180.660
111.440
94.042
101.553
42.760
35.160
1505.540
293.840
240.040
321.000
247.000
215,000
62.200
69.260
82.531
-
136.479 (-1)
177.167 (-1)
113.204
95.596 (-1)
105.522
45.878
35.388
1503.551
289.479 (-1)
283.306
320.490
250.755
23.347
58.673
69.898
81.449
X
-
SE
0.650
0.745
0.662
0.688
1.014
0.499
0.342
6.614
1.848
2.080
1.751
2.117
2.281
2.019
2.140
1.483
X
135.125
180.042
111.396
96.721 (-1)
104.128
43.271
35.875
1500.833
286.813
233.958
317.478 (-2)
254.688
225.146
80.000
87.500
88.479
~
SE
0.725
0.892
0.725
0.728
0.910
0.471
0.407
9.201
1,883
1,804
2.135
1.870
2.535
3.332
2.600
1.680
134.459
181.054
115.297
94.118
104.056
44.946
36.027
1515.216
287.784
238.432
312.081
247.838
222.622
75.676
78.514
87.943
X
-
SE
0.618
0.954
0.872
0.936
0.883
0.520
0.424
7.164
2.479
1.684
2.923
2.168
2.920
3.520
4.132
1.669
anthropometric measurements after omitting outliers (number indicated in bracket)
Raj Gonds (37)
Plains Maria Gonds (48)
Manne (49)
SE
0.609
0.779
0.855
0.862
0.817
0.436
0.369
8.521
2.113
2.073
1.727
1.929
2.141
2.997
2.815
1.580
of
Koya (50)
T A B L E 3. Female tribals: mean and standard error
'HB, head breadth; HL, head length; BzB, bizogmatic breadth; BgB, higonial diameter; TFL, morphological face height; NL, nose length; St, stature; UAL, upper arm length; LAL,
forearm length; BAB, biacromial diameter; BIB, bi-iliac diameter; MAC, midarm circumference; TFF, triceps skinfold; SFF, subscapular skinfold; Wt, weight; NB, nose breadth.
HB'
HL
BzB
BgB
TFL
NL
St
UAL
LAL
BAB
BIB
MAC
TFF
SFF
Wt
NB
C h ilri rt.er
Kolam (52)
X
TABLE 2. Male tribals:
8
-J
co
N
z:
8
z
c:
5z
?-
3m
2
=!
n
m
5
z0
?-
E2
w
0
!s
?-
298
U. PINGLE
TABLE 4. Analvsis o f variance findiuidual anthrooometric characters): F statistics
Characters
HB
HL
BzB
%?
NL
NB
St
BAB
BIB
Wt
Between
male tribals
F (4,482)
Between
female tribals
F (4,482)
Sex difference
within tribals
F (5,482)
Sexual dimorphism
between tribes
F (4,482)
9.3106* *
2.5922*
1.0165
2.8285*
1.6866
3.4251**
4.6603**
2.6680*
1.4555
6.0459**
7.7744 * *
2.5778*
4.8178**
3.8753**
2.0975
2.4057*
5.3792**
2.0819
1.0683
4.2209**
3.3935**
4.3188**
29.0532**
33.4196* *
34.4140**
50.4718**
31.3000**
31.6999**
25.7829**
89.6933**
181.9257**
9.6177**
46.0932**
3.8416 * *
0.2814
1.6832
0.2658
0.1042
1.2634
1.6448
1.3948
2.0438
0.2774
0.3535
'Abbreviations as in Table 2
* P i.05.
** P < .01.
TABLE 5. D 2matrix based on anthropometrics in Gonds (males and
females)
Kolam
Kolam
Koya
Manne
Plains
Maria Gonds
Rai Gonds
SINGLE
-
KL1
PY
Kova
Plains
Manne Maria Gonds Rai Gond
1.8831 0.8991
2.5188
1.6306
1.5878 0.8756
3.0084 1.5379 1.3564
1.5741
1.3567
1.5132
-
1.2068
2.2054
2.2286
2.3750
1.4313
2.7960
-
1.3700 1.2776
LINKAGE
KYl
YN
COMPLETE
R G t
-
1.0
-
t.2
-
1.2
-
1.4
0.2
0.4
0.6
0.8
1.4
i d 6
1.8
2 4 0
2'2
-
0.2
0.4
0.6
0.0
1.0
PY
KL1
-
-
1.6
-
1.0
-
2'0 2.2
-
2.4
-
2.6
-
2.0
-
3.0
3.2
3.4
L INKACE
K 1 1
-
Fig. 2. D2 among Gond males (cluster analysis).
MN
RGI
299
ANTHROPOMETRIC AND GENETIC DISTANCE AMONG GONDS
SINGLE
-
KY4
PY
LINKAGE
KL4
YN
COMPLETE
RO4
0.4
-
0.6
-
0.6
0.0
-
0.8
0.2
0.2
0.4
1.0
-
1.2
-
1.2
I. 4
-
1.4
1.6
-
1.6
1.8
-
1.8
1.0
. 2 0 -
2.0
2 2 -
2.2
-
-
-
2.6
2.8
-
2.8
-
3.0
-
-
3.4
R01
-
2.6
3.4
PY
-
-
3.2
LINKAGE
-
2.4
-
KY4
-
-
3.2
YN
-
2.4
3 0 -
KL4
-
-
Fig. 3. D2 among female Gonds (cluster analysis).
Subgraphs at various levels of clustering
based on complete linkage method were
drawn for both males and females separately
(Figs. 4 and 5). These help in showing the
multivariate spatial distributions of these
groups, which in both sexeg generally correspond with their geographical distribution.
The Manne lie intermediate linking the Raj
Gonds and Kolams with the Koyas and
Plains Maria Gonds.
Genetic analysis
All the groups are significantly different
from each other in the ABO gene frequency
distribution. The B gene frequency is highest
in Kolams, Raj Gonds, and Manne (Table 6).
x2 test was not carried out in the distribution of the Rh blood group types because of
the low frequency of Rh negative allele in
some of the groups. The Rh negative allele is
highest in Raj Gonds and Kolams (Table 6)
and lowest in Koyas.
All groups show heterogeneity among each
other for MN system (P < 0.05). The N gene
frequency is lowest in the Manne and Plains
Maria Gonds (Table 6).
The Gond populations were significantly
heterogenous for both hemoglobin and G6PD
deficiency. The hemoglobin S gene frequency
was highest in Plains Maria Gonds and Raj
Gonds. The highest G6PD deficiency is found
among the Raj Gonds and Plains Maria
Gonds (Table 6). These groups also have a
high hemoglobin S gene frequency, which is
consistent with the fact that the grou s were
and are in areas hyperendemic for jaalciparum malarial infection.
These groups are however homogenous for
haptoglobin protein. The haptoglobin 1 gene
frequency is low in these populations ranging from 0.06 to 0.13, comparable to most
Indian populations (in whom it is very low)
(Table 6).
The x2 test showed significant heterogeneity for transferrin types between all Gond
populations at the P < 0.05 level. The slow
variant D transferrin (subtyping for D variant not done), which is rare in other populations in India, is highest in Kolams, Manne,
Raj Gonds, and Koyas (Table 6) and low in
Plains Maria Gonds, showing a genetic cline
from west to east with a decrease in D transferrin in the eastern groups. Another Dravidian-speaking group, the Oraons, have also
been found to have a high D gene frequency
(Kirk et al., 1962; Mukherjee et al., 1975).
This D gene may prove a useful marker in
differentiating the Dravidian-speaking from
U.PINGLE
300
Level 1.5
Level < 1.0
KL1
0L
K l t
MN
Level 2.0
RG1
PM
Level 2 2.5
RG1
PM
Fig. 4. D2 among male Gonds (complete subgraphs)
forming separate endogamous entities. In addition, this tribal endogomy has a territorial
basis, each of these populations being located
in distinct geomorphological areas (Fig. 1).
This is further substantiated by linguistic
and cultural differences between them.
When projected graphically, the morphological and genetic distances between these
groups correspond generally to their present
geographical distribution. The Manne lie intermediate, connecting the other four populations. The Raj Gonds and Plains Maria
Gonds lie farthest away from each other a t
two opposite poles. However, the morphological and genetic relationships between the
Gond populations differ among themselves.
The morphological relationship is the less
consistent, the cluster patterns differing by
sex. The Koya females are nearer the Manne
and Raj Gond and further from the Plains
Maria Gond females. The Kolams also vary
by sex; the males are removed from all the
groups, but the females are very close to the
Manne females. This variability by sex in
morphological relationship is difficult to inDISCUSSION
terpret and may be in part due to differences
Statistical analysis showed that the Gonds in sex dimorphism and the greater degree of
are a heterogenous group, morphologically environmental influence on morphological
and genetically different from each other, characteristics.
the Munda-speaking tribals, as well as from
South Indian tribals in whom the frequency
of D gene is found to be very lwo.
Nei’s and Sanghvi’s distance measures
were computed based on gene frequencies of
seven biochemical characters (Table 7).
The two methods of clustering based on
Nei’s and Sanghvi’s genetic distance matrices showed close correspondence in the relationships among the five tribal groups
(Figs. 5 and 6). The Raj Gonds (RG1) and
Kolams (KL1) cluster together, whereas the
other three, the Plains Maria Gonds (PM),
Manne (MN) and Koya (KYl), form a separate cluster.
Complete subgraphs based on both these
distance measures also showed a close correspondence in the relationship and spatial distribution of the five groups. The Manne are
a n intermediate group connecting the Koyas
and Plains Maria Gonds with Kolams and
Raj Gonds (Fig. 7). The Manne are closer to
Koyas and Plains Maria Gonds than to Kolams and Raj Gonds.
HP
1
2.1
2.2
Transferr i n
C
CD
D
G6PD
Normal
Deficient
S
0.0352
S
1.0000
0.0000
0.0403
D
0
0.9597
C
Hp2 0.9270
Hpl 0.0730
0.9648
0.2143
n
A
0.7857
m
149
137
12
0
0
13
76
132
10
0
62
30
6
MN
M
MN
N
Hb
A
AS
0.7575
0.2425
D
96
6
d
0.1748
0.3527
0.4725
p
q
r
17
44
25
16
ABO
A
B
0
AB
Rh
D
d
Obs.
freq.
125
6
187
4
0
0
20
135
213
15
0
75
29
5
168
1
56
39
57
17
Obs.
freq.
0.1789
0.8211
0.9231
0.0769
0.2452
0.1814
0.5734
0.9924
0.0076
0.0105
0.9895
Hp2 0.9355
H p l 0.0645
Hbs 0.0329
HbA 0.9671
n
m
D
d
p
q
r
Koyas
Gene
frequency
142
0
136
6
0
0
15
83
0.0130
0.9870
0.1188
0.8812
0.8620
0.1380
0.2099
0.2347
0.5554
0.0211
D
1.0000
0.0000
0.9789
C
Hp2 0.9427
H p l 0.0573
S
A
n
151
4
0
m
18
3
d
D
p
q
r
Manne
Gene
frequency
80
103
2
7
32
36
30
Obs.
freq.
7
106
159
3
0
0
18
78
0
140
27
86
13
4
101
2
28
26
40
9
0.0808
0.9191
0.1091
0.8981
0.8606
0.1393
0.1985
0.1865
0.6150
0.0093
D
0.9381
0.0619
0.9907
C
Hp2 0.9063
Hpl 0.0931
S
A
n
m
d
D
p
q
r
Plains Maria Gonds
Obs.
Gene
freq.
frequency
TABLE 6. PhenotvDe and gene freauencies for genetic markers in the Gond DoDulations
Kolams
Gene
frequency
92
13
135
8
0
0
14
63
IL20
20
0
61
27
3
83
7
24
34
21
12
Obs.
freq.
0.0714
0.9286
0.1813
0.8187
0.7211
0.2789
0.2226
0.2968
0.4806
D
C
0.9341
0.0659
0.0280
0.9720
Hp2 0.9091
Hpl 0.0909
S
A
n
m
D
d
p
q
r
Raj Gonds
Gene
frequency
s
w
2
s
0
0
2,
$5
u
b
m
0
2
2
r
(I)
El
c,
z2
22
5c3
E>
3
8
9
*
5
302
U. PINGLE
which is predominantly influenced by Telugu
culture. Unlike the morphologic, the genetic
characteristics are purely heritable and, not
being influenced by environment, provide a
finer tuning to population differences. The
genetic relationship also reflects more closely
the effect of geographical proximity in producing a greater genetic similarity within
these two clusters of Gond populations.
The emphasis of the present study based
on quantitative techniques is on the importance of geographical proximity in producing
morphological and genetic similarity between populations, which is brought about
The relationship based on genetic characters however is different from the morphological relationship and corresponds closely to
the degree of geographical contiguity between the populations. The Raj Gonds and
Kolams form a close cluster, with Manne,
Koyas, and Plains Maria Gonds forming a
second cluster. The Raj Gonds and Kolams
live in close proximity to each other on the
highlands consisting of black cotton soil,
which is contiguous with Marathwada,
whereas the Manne, Koyas, and Plains Maria
Gonds lie on the east in the valley of the
Pranitha-Godavari and Indravathi rivers,
TABLE Z Genetic distance in Gonds (Nei and SanehuiJ
Kolam
Koya
Manne
Plains Maria
Gonds
Raj Gonds
Kolam
Koya
Manne
Plains
Maria Gonds
Raj Gonds
-
0.5375
0.2837
0.4559
0.0889
0.0641
0.0365
0.0548
-
0.0956
0.0119
0.0182
0.1877
0.1457
0.0088
-
0.6238
0.3536
0.3537
0.0604
0.0325
0.0389
-
0.0095
Nei’s distance
COMPLETE
‘001-
101
KLL
PY
-
LINKAGE
YN
101
.Wl-
-
-
-
.ooa-
.005 -
*005
-003
-
-
-009.O2 -
-007
*04
‘06
.08
-
-
cI
SINGLE LINKAGE
111
.007
-
.w1.02
-
.04
-
-
.08 .06
Sanghvi’s
distance
KLL
PY
-
Fig. 5. Nei’s genetic distance among Gonds (cluster
analysis).
YN
KYI
303
ANTHROPOMETRIC AND GENETIC DISTANCE AMONG GONDS
COMPLETE
RG1
KL1
PY
LINKAGE
MN
SINGLE LINKAGE
KV1
RG1
KL1
PM
MN
L
T
Fig. 6. Sanghvi's genetic distance among Gonds (cluster analysis).
0.01 L.v.1
MN<Kvl
0 02
L.V.1
CY
0 . 0 6 4 Llr.1
0 . 0 4 Lev.1
.009
RG1
Fig. 7. Nei's genetic distance among Gonds (complete
subgraphs).
KY!
U. PINGLE
304
by a closer distance a s well a s similar geographical factors (such as soil, terrain, flora,
etc.), drawing these populations together under a common ecocultural umbrella. The
closer the geographical distance between
populations, the closer the genetic as well as
morphological similarity between them. This
is more so in populations such as these who,
due to having adopted a settled agricultural
occupation, are less mobile. This low mobility in turn produces a smaller marriage distance (distance between place of birth of
spouses) thus restricting the spacial distribution of genes to a smaller area.
ACKNOWLEDGMENTS
I am deeply grateful to Professor C.R. Rao,
F.R.S., Jawaharlal Nehru Professor, Indian
Statistical Institute and at present Visiting
Professor, Department of Mathematics and
Statistics, University of Pittsburgh, for his
guidance throughout the work and for having given me all facilities for analysis of the
data in his department a t the University of
Pittsburgh.
I am indebted to Mr. N.V. Raja Reddy,
Bheeemaram (Andhra Pradesh), for his constant and sustained encouragement and advice throughout the field work.
I am thankful to Professor C. von FurerHaimendorf, Emeritus Professor, School of
Oriental and African Studies, University of
London, for his valuable comments on the
conclusions of my study.
LITERATURE CITED
Balakrishnan, V, and Sanghvi, LD (1968) Distance between populations on the basis of attribute data. Biometrics 245359-865.
Box, GEP (1949)A general distribution theory for a class
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Cavalli-Sforza, LL, and Edwards, AWF (1967) Phylogenetic analysis: Models and estimation procedures. Evolution, 21,550-570.
Census of India (1961) District Census Handbook,
Chanda, Director, Government Printing and Stationery, Maharashtra State, Bombay.
Clark, JT (1964) Simplified ‘disc’ polyacrylamide gel
electrophoresis. Ann N.Y. Acad. Sci. 121t305.
Dietz, AA, Lubrano, T, and Rubinstein, HM (1971) Haemoglobin and haptoglobin determination by disc electrophoresis. Clin. Biochem., 4.59.
Elwin, V (1947) Muria and their Ghotul. Bombay: Oxford University Press.
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Delhi.
Grierson, G (1906) Linguistic Survey of India, Vol. IV,
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Motulsky, AG, and Campbell-Krant, JH (1960) Population genetics of GGPD deficiency of the red cell. In BS
Blumberg (ed): Proceedings Conference Genetic Polymorphism and Geographic Variations in Disease. New
York: Grune and Stratton.
Mukherjee, BN, Das, SK, and Sharma, DP (1975)Serum
protein and red cell enzyme polymorphism in Oraon
tribe, India. Ann. Hum. Biol. 2t201-204.
Nei, M. (1972) Genetic distances between populations.
Am. Naturalist 106t283-292.
Pingle U (1981) Ph.D. thesis: “Morphological and Genetic composition of Gonds of Central India: A Statistical Study” (unpublished) submitted to Indian
Statistical Institute, Calcutta.
Rao, CR (1965) Linear Statistical Inference and Its Applications, 2nd edition. New York: John Wiley and
Sons.
Sneatb, PHA, and Sokal, RR (1973) Numerical Taxonomy. San Francisco: W.H. Freeman and Co.
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Blackwell Scientific Publications.
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