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Population studies on Southwestern Indian Tribes. IV. The Zuni

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Population Studies on Southwestern Indian Tribes
IV. THE ZUNI
P. L. WORKMAN,' J. D . NISWANDER, K. S . BROWN AND
W . C. LEYSHON
Human Genetics Branch, National Institute of Dental Research, National
Institutes of Health, Bethesda, Maryland 20014
KEY WORDS
Amerindians.
Genetic differentiation . Kinship . Distance . Zuni
ABSTRACT
The Zuni Indians of west-central New Mexico have been relatively
isolated since their foundation by an amalgamation of individuals from different
southwestern cultural areas during the Regressive Pueblo period (c.1200-1350
A.D.). Genetic analysis revealed a high frequency of blood type B in both young
(0.06) and old (0.05) Zuni, but at 14 other blood group and serum protein loci,
allelic frequencies including A (0.011) and Rh negative (0.001) were generally
similar to those of other relatively unmixed southwestern Indian tribes. Consideration of Zuni history and demography since Spanish contact in 1540, together
with genetic analyses, suggest that the high B frequency probably derives from
intermixture with a small number of B, Rh positive non-Indians in the early post
contact period. Genetic differentiation among four southwestern tribes, Zuni,
Pima, Papago and Maricopa, was summarized by kinship analysis. Approximately
70% of the inter-tribal genetic variation could be explained by the geographic
distances among these groups showing that isolation by distance has been the
most important factor in determining the pattern of regional genetic differentiation.
The Zuni Indians living in west-central
New Mexico appear to be an amalgamation of at least two indigenous southwestern Indian groups. Cushing (1896) first described Zuni mythology and language as a
fusion of distinct elements, and recent linguistic studies appear to confirm this (Newman, '64; Swadesh, '59, '67). The Zuni
language is quite distinct from other New
Mexico Pueblo languages and not related
to the Uto-Aztecan language of the Papago,
Pima and Hopi in Arizona.
In addition to language affiliations, reconstruction of the history of the southwestern tribes must also account for tribal
variation in cultural and biological factors.
As part of an ongoing study of biological
differentiation in the southwest we will restrict our focus in this paper to a genetic
characterization of the Zuni and to the pattern of genetic differentiation among the
indigenous southwestern Indian tribes. The
genetical analyses contribute to the larger,
yet unwritten, history through an understanding of the most probable patterns of
AM. J. PHYS. ANTHROP.,
41: 119-132.
mate exchange (which may or may not
have been accompanied by cultural or linguistic influences) and, by inference, on
the likelihood that different groups may
have descended from a common founder
population. In order to interpret the pattern of biological variation it is first
necessary to examine observations from
archaeology, ethnohistory and historical
demography.
HISTORICAL OBSERVATIONS
Prehistory to Spanish contact (I 540 A.D.)
Man is believed to have first entered the
southwest at least as early as 10,000 B . C .
In the northern region (Nevada, Utah, Colorado) the first men were primarily big
game hunters; their earliest known sites
contain flint tools and projectile points in
association with the bones of mammoth,
long-horned bison, and traces of fire. In
southern Arizona and New Mexico, however, the frequent occurrence of milling
'
Present address: Department of Anthropology, University of Massachusetts, Amherst, Massachusetts 01002.
119
120
P. L. WORKMAN, J. D. NISWANDER, K. S. BROWN AND W. C. LEYSHON
stones as well as bones, in the earliest sites,
suggests a culture more dependent upon
hunting and gathering. By 6000 B . C . men
were widely dispersed in small bands and
the heterogeneity among artifacts from
different sites believed to be contemporaneous makes it likely that cultural diversification accompanied the dispersion.
For several thousand years the southwestern peoples developed in relative isolation. Sites dating from 2500 B . C . show
remains of baskets, sandals, milling stones
and flint and bone tools. The entire southwest at that time appears to have been inhabited by small, seminomadic bands of
hunters and gatherers. Under such conditions, cultural differentiation could proceed without significant biological divergence. The genetic structure might be well
approximated by a model of partially isolated subgroups in which the amount of
intermixture varies with the travel time
between groups.
During this same period there arose in
the southern Mexican plateau a culture
whose contributions were to dominate all
further change in the southwest. The most
important development in Mexico was the
invention of horticulture and the domestication of squash, beans, maize and many
other plants. By about 1000 B.C. these agricultural developments had been introduced
into the southwest and the seminomadic
groups were generally replaced by larger,
sedentary agricultural settlements which
may have had both greater cultural isolation and a higher degree of endogamy.
Another, somewhat later, contribution from
Mexico was pottery which was introduced
into the region by about 200 B . C . After this
period we can trace the development of
three great culture areas: The Mogollon in
southern New Mexico and southeastern
Arizona; the Hohokam in southcentral Arizona; and, the Anasazi, centered on the
four corners area, i.e., the common border
of Utah, Colorado, Arizona and New Mexico. Each of these, of course, should not be
taken to represent a single Mendelian population but rather a collection of groups of
different sizes and degrees of isolation sharing numerous culture traits (burial practices, axe types, architectural style, pottery
type, etc.). In addition, especially by analysis of pottery variants, local differentiation within these culture areas can also be
recognized.
There appears to have been sufficient
cultural, and possibly biological, interchange among these areas so that between
500 A . D . and 1100 A . D . a generally uniform
style of life obtained throughout the southwest. From about 700 A . D . there was an
expansion of each of the cultures into new
sites. The main thrust of the Anasazi after
900 A . D . was an expansion into the Rio
Grande Valley, which, before then, had
been sparsely populated. During this
period the Anasazi began the construction
of the first large communal houses, or
Pueblos. One of the most famous, dating
from about 1000 A.D. was that at Chaco
Canyon in northwestern New Mexico. During the twelfth century, called the Great
Pueblo period, the Anasazi rose to its highest point. The Indians moved from their
small scattered settlements and congregated in the large pueblos, some of which
had between 500 and 1,000 rooms.
The thirteenth century, in marked contrast to the preceding period, is described
as the Regressive Pueblo period. The great
houses were gradually abandoned as was
most of the territory which had been occupied by the Anasazi. In general, the population withdrew to the Rio Grande Valley but
three somewhat isolated areas remained
occupied in the west in sites now occupied
by the Hopi, Zuni and Acoma tribes. A variety of reasons have been offered to explain
these events: overpopulation; disease; attacks by hostile bands of Utes and Athapascans moving down from the northern
plains; drought which made agriculture in
the area almost impossible; and, a shift
from dominant winter rain to heavier summer rain which inhibited spring planting
and led to increased erosion. Despite evidence for an almost total lack of rainfall
between 1276 and 1299 it is likely that all
of these factors may have been of some importance. At the same time many Mogollon
settlements in New Mexico were also abandoned. Throughout the southwest, the Indians congregated in large villages, or
groups of related villages to a large extent
both geographically and culturally isolated.
Many of the contemporary southwestern
Indian tribes trace their origin to this period.
In the Zuni area, between 700 A.D. and
1100 A . D . , the inhabitants were dispersed
in many small settlements often containing
only a single family. During the tweIfth
POPULATION STUDIES ON THE ZUNI INDIANS
century several large pueblos were built
and many small settlements were abandoned. A large pueblo at Atsinna, dating
from 1100, had more than 500 rooms. All
of the sites from this period demonstrate a
culture similar to that at Chaco Canyon,
called San Juan Anasazi. However, sites
dating after 1300 show a rather marked
change to a culture very much like that of
the Mogollon to the west and south. Sites
whose period of use spanned this period
show both Pueblo and Mogollon features.
For example, at Atsinna, the last structure
in the Anasazi style was probably constructed about 1300; rooms dating from a
later period are in the Mogollon style.
It is likely that the original inhabitants
were absorbed by northern groups of the
Mogollon, and not merely displaced, because Zuni language and mythology appears to be a fusion of two distinct cultural
traditions. Other groups may also have
contributed to the population in the area.
For example, there is good evidence for extensive Zuni-Hohokam trade (Bandelier,
1892). During the general Pueblo withdrawal, migrants from the north probably
settled at Zuni where water was available
throughout the period, and, successful
farming may have been possible. Thus, by
1400, the Zuni appear to have been a rather heterogeneous mixture of southwestern
peoples. However, the majority were probably of Mogollon descent and even in the
late nineteenth century, the Zuni made
ceremonial visits to shrines in the west and
southwest, territory formerly occupied by
the Mogollon.
Between 1300 and 1400, many additional
Zuni settlements were abandoned and the
Zuni withdrew into six large villages well
isolated from their nearest neighbors at
Acoma. The Acoma Indians congregated on
a single mesa some 75 miles from the nearest Zuni village: prior to the fourteenth
century, Zuni and Acoma settlements had
been only 25 miles apart. Although there
was probably some contact and mate exchange with Acoma and with the Hopi to
the west, the Zuni appear to have remained
relatively isolated until Spanish contact in
1540. The distribution of Indian groups
in the southwest just prior to that time is
shown in figure 1.
The archaeological observations summarized here are based primarily on the
work of Reed (‘49, ’50, ’55), Wendorf and
121
Reed (’5.3, Woodbury (‘56) and Rinaldo
(‘64).
Spanish contact to the present: In 1540,
the Spanish explorer, Coronado, accompanied by some 30 soldiers and a number
of Nahuatl speaking Indians from Jalesco
and Sonora in Mexico, sought the legendary “Seven Cities of Cibola.” “Shibola”
was, in fact, Zuni, and Coronado, guided
by a Zuni residing with the Pima, found
six villages of adobe instead of seven cites
of gold. The Zuni were partially subdued
and when Coronado left, there were at least
three Indians from Mexico known to have
married into the tribe (Bandelier, 1892).
This contact provided the first opportunity
for gene flow with Spaniards; other contact
with Mexican Indians, however, might have
come earlier.
Throughout the following 200 years, the
Zuni resisted Spanish secular and religious
influence. The first Catholic church was
established at Zuni in 1628 but priests
were killed in 1630, 1670, and at the time
of the general Pueblo revolt against Spanish domination, in 1680. Again, in 1703,
the Spanish resident policeman was killed.
Each time major conflict with the Spanish
appeared likely, the Zuni moved to the top
of their sacred mesa which rises 1,000 feet
above the surrounding plain. The mesa has
water and grass and could be easily defended by very few men since only one narrow, steep trail permitted access to the top.
From 1680-1692 and in 1703 the mesa
became the home of the Zuni, and overall,
the tribe spent several years in protected
isolation. During this period, the smaller
villages were effectively abandoned and the
majority of the Zuni lived in a central
pueblo, Zuni.
These experiences enabled the Zuni to
retain cultural, and, probably, biologcal
integrity to a far greater degree than most
other southwestern Indians. With the exception of the Hopi to the west, the Zuni is
the only Pueblo group in which Spanish
is not spoken and in which Spanish influences are noticeably absent. The retention
of so much of their own culture encouraged
numerous ethnological studies including
those by Cushing, 1896; Stevenson, ’04;
Kroeber, ’19; Bunzel, ’32; Benedict, ’35;
and Eggan, ’50.
Today, the Zuni pueblo has been relatively modernized. On reservation employment includes farming and jewelry manu-
122
P. L. WORKMAN, J. D . NISWANDER, K . S. BROWN AND W . C. LEYSHON
Fig. 1
Culture areas of the southwest 1500 A.D.
facture and the first phase of an industrial
complex providing more technical jobs has
been completed. However, the desire to retain their cultural identity persists most
strongly. Adair and Vogt (‘49) in a study
of returning veterans noted that “As the
village has become more modern in its
technology, antagonism to white culture
has become increasingly apparent, especially to those values which threaten Zuni
religion.”
On the whole, ethnohistorical observations suggest that the Zuni are likely to be
biologically quite similar to the heterogeneous ancestral population which appears
to have been established during the fourteenth century. There is no evidence for
significant intermixture with non-Indians
and intermarriage with other Pueblo Indians has never been frequent.
DEMOGRAPHY
In order to use contemporary gene frequencies for inferences about relations
among southwestern tribes, it is also necessary to consider the possibility of fluctuations in population size, which might have
resulted in substantial genetic drift.
Estimates of the population size at Zuni
prior to Spanish contact vary between
3,000 and 6,000 (Cushing, ’20). For example, according to Bandelier (1892), at the
time of contact there were six villages with
a total of about 3,000 inhabitants. The
largest village, Hawikah, had between 200
and 300 families; the smallest had 30 to 60
families. As shown in table 1 and figure 2,
the population appears to have declined
slowly until it reached a low point of 1,200
in 1864. Since that time there has been a
steady rise, which has accelerated rapidly
in the past two decades.
The decline and slow return to precontact numbers must have involved considerable loss due to epidemics. As recently
as the winter of 1898-1899 about 250 died
from smallpox (HrdliEka, ’08). However,
major improvements in medical facilities,
123
POPULATION STUDIES ON THE ZUNI INDIANS
TABLE 1
TABLE 2
Estimates of Zuni population size f r o m 1540
to 1968
Age and sex distribution of Zuni born prior to
Year
1540
1680
1790
1809
1850
1864
1879
1890
Size
Year
Size
1900
1910
1920
1930
1940
1950
1963
1968
3000
2500
1935
1598
1500
1200
1500
1547
J a n . 1,1969
Age
1523
1640
1813
1952
2205
2922
5011
6007
Source of data: 1540 (Bandelier, 1892); 1680 (Cushing, ’20);1790-1840 (AnnualReportof thecommissioner
of Indian Affairs, 1867); 1850-1940 (Annual Reports of
the Commissioner of Indian Affairs); 1950 (Bureau of
Indian Affairs); 1963 (Public Health Service); 1968
(present study).
Male
Female
Total
70 of
Total
1 4
5 9
10-19
20-29
30-39
40-49
50-59
60439
70
364
503
746
438
300
239
187
140
137
321
488
778
497
345
184
159
88
93
685
991
1524
935
645
423
346
228
230
11.4
16.5
25.4
15.6
10.7
7.0
5.8
3.8
3.8
Total
3054
2953
6007
100.0
+
ical of the population for many years. The
1921 and 1941 estimates included males
and high fertility without high child mor- and females in ratios 1,040/823 and 1,245/
tality, have resulted in a current growth 1,007, respectively. In an early medical
rate of over 5% per year.
study, Fleming (‘24) was “surprised to find
The age and sex distribution of the pop- many men of approximately 70 years of age
ulation, based on the Zuni family register leading lives of extreme physical activity.”
as of spring 1968, is given in table 2. A Studies on the distribution of hyperglycesmall proportion of the 6,007 Zuni may mia, to be published elsewhere, suggest
actually be living off the reservation. Most that the life style of the Zuni, which inindividuals live in the main village at Zuni, cludes considerable sex differences in activbut there are also small farming settle- ity and diet, may be responsible for the
ments at Ojo Caliente, Tekapo, Pescado, observed two to three times greater inciand Nutria. As expected in a rapidly in- dence of hyperglycemia in Zuni women.
creasing population, the population is, on Women over 60 have a much lower average
the average, quite young. Over 53% are blood glucose level than women aged 35less than 20 years old. The data of table 2 60, and it is possible that a greater mortalshow an atypical sex ratio pattern. In the ity of females related to hyperglycemia,
early years, first males (1-9 years) and obesity, and hypertension may account, in
then females (10-35) predominate. How- part, for the predominance of males among
ever, more adults over 35 are male and the the older Zuni. Analyses of the medical obdifference is quite large for those over 60. servations on the Zuni will be presented in
This distribution appears to have been typ- detail elsewhere.
6000
-
ferential fertility in populations of this size
may result in rather marked changes in
gene frequencies. HrdliEka (‘08) noted that
few women were sterile and that most families had several children. Thus any effect of differential fertility would be more
likely due to variation in the reproductive
rates of males.
5000 W
N
V, 4000-
z
5
2
a
x
30002000-
1000 I
I
I
I
I
GENETIC ANALYSES
124
P. L. WORKMAN, J. D. NISWANDER, K. S. BROWN AND W. C. LEYSHON
tensive physical examinations including
detailed dental and ophthalmological examinations were done on over 450 Zuni over
20 years of age. Urine samples were obTABLE 3
Distribution of Zuni “full bloods” i n the sample
population b y nge and sex
Age
% sample
of Zuni in
agegroup
Male
Female
Total
Total
Zuni
13-19
20-29
30-39
4049
50-59
60-69
a70
72
50
26
13
18
18
22
137
130
49
39
39
22
27
209
180
75
52
57
40
49
997
935
645
423
346
228
230
21.0
19.3
11.6
12.3
16.5
17.5
21.3
Total
219
443
662
3804
17.4
tained for glucose and albumin screening.
Briefer examinations of about 200 schoolchildren were conducted. This provided observations on 662 Zunis for genetic analysis
and for comparative studies of adaptation
and health status in southwestern Indian
tribes.
Blood was drawn for serological analyses,
two hours after ingestion of a 75 gm glucose equivalent carbohydrate load. Blood
samples were divided into three aliquots;
one with ACD as an anticoagulant, a second containing fluoride as an enzyme inhibitor and a third clotted sample was obtained without additives for enzyme studies.
Blood glucose values were determined by
the Southwestern Field Studies Section of
the National Institute of Arthritis and
Metabolic ~
i
~
~
TABLE 4
Distribution of Zuni phenotypes
Phenotype
No.
quency
ABO
0
A1
B
576
15
71
0.870
0.023
0.107
MNSs
MMSS
MMSs
MMss
MNSS
MNSs
MNss
NNSS
30
156
199
4
77
154
2
4
36
0.045
0.236
0.301
0.006
0.116
0.233
0.003
0.006
0.054
CcDEe
CCDee
ccDEE
CcDee
ccDEe
CCDEE
CCDEe
CcDEE
ccdEe
198
329
39
30
11
1
48
5
0.299
0.497
0.059
0.045
0.017
0.001
0.073
0.008
1
0.001
PI ( + )
Pi(-)
533
129
0.805
0.195
Duffy
Fya+
Fya -
618
44
0.934
0.066
Kell
kk
kkP
656
6
0.991
0.009
Lewis
Lea-
662
1.000
Diego
DibDib
DiaDib
DiaDia
631
31
0
0.953
0.047
NNSs
NK S b
Rh
P
Fre
Fre
System
0.000
System
Phenotype
+
No.
quency
Kidd
Jka
Jka -
336
326
0.508
0.492
Cerumen
W-(wet)
ww (dry)
160
298
0.349
0.651
Haptoglobin
1-1
1-2
2-2
279
310
68
0.425
0.472
0.103
Group
specific
1-1
1-2
2-2
485
157
13
0.740
0.240
0.020
Red cell
Acid phosphatase
PAPA
PBPB
32
210
418
0.048
0.318
0.633
Transferrin
TfCTf‘
TfCTf 6 O - l
TfBO-1
530
121
8
0.809
0.185
0.006
Albumin
AL~AL
A
639
A L ~ A L ~ ~ 16
~
0.976
0.024
Phosphoglucomutase: PGMl
1-1
1-2
2-2
434
203
025
0.655
0.307
0.038
PAPB
~
125
POPULATION STUDIES ON THE ZUNI INDIANS
TABLE 5
Zuni gene frequencies
System
ABO
Allele
Frequency
0
At
B
0.933
0.011
0.055
MS
Ms
NS
Ns
0.213
0.546
0.020
0.221
CDe
cDE
cDe
CDE
0.704
0.219
0.033
0.043
System
Rh
0.001
cde
Frequency
Jk "
0.296
0.704
W
W
0.193
0.807
Haptoglobin
HPI
HP~
0.661
0.339
Group
specific
Gcl
Gc~
0.860
0.140
Red cell
Acid phosphatase
PA
PB
0.208
0.792
Tr an sferr i n
TfC
TfBO-1
0.902
0.099
Albumin
A L ~
ALM@X
0.987
0.012
PGMI'
PGMiZ
0.809
0.191
Kidd
jkb
Cerumen
MNSs
Allele
,
P
p1
PZ
0.559
0.441
Duffy
FYa
Fyb
0.742
0.258
Kell
k
kP
0.995
0.005
Lewis
Le
Lea
1.000
0.000
Diego
Dia
Dib
0.023
0.977
Red cells were typed for A, Al, B, M, N ,
S, s, U, P, C, C " , E, c, e, K, K p n , K P b ,
k, Lea, Fy", Jk", Di" and Dib . Haptoglobin albumin, transferrin and group specific serum protein types were determined
by polyacrilamide gel electrophoresis according to methods previously described
(Brown and Johnson, '70). Red cell acid
phosphatase and phosphoglucomutase genotypes were determined by the methods of
Karp and Sutton ('67) and Spencer, Hopkinson and Harris ('64) respectively.
The recent ancestry of the Zunis in the
sample was determined and 95% could
be putatively described as full-blooded.
The analysis in the present paper is restricted to the sample of full-blooded Zuni
as described in table 3. Data on the Gc, Hp,
AL and Tf systems has been reported by
Brown and Johnson ('70) but, for completeness, we present all of the phenotypes and
gene frequencies in tables 4 and 5, respectively.
Ten codominant systems permit estimation of Fi = 1 - Hi/2piqi where Hi is
the observed proportion of heterozygotes
and pi, qi = 1 - pi are the frequencies
for the ith allele. A s pointed out by Harpending, Workman and Grove ('73), if a
population is not highly endogamous, migration effects alone will cause the expected value of this estimate to be negative.
As shown in table 6 , the mean of the Fi is
0.0019 which provides support for the asTABLE 6
Estimates of F and corresponding chi-square
tests for codominant loci
System
or locus
MN
ss
cc
Ee
Ac.P.
Di
PGM
HP
Gc
Tf
mean (Fi)
Fi
x2 = F2N
0.0295
- 0.0005
0.0689
- 0.0057
0.0328
- 0.0240
0.576
0.0002
3.141
0.022
0.711
0.380
0.043
1.791
0.005
1.071
0.0081
- 0.0522
0.0028
- 0.0404
0.0019
126
P. L. WORKMAN, J. D. NISWANDER, K. S. BROWN AND W. C. LEYSHON
sumption that the Zunis are relatively
isolated. Chi-square tests for each locus,
comparing observed frequencies with these
predicted by the Hardy-Weinberg Law, are
given by x ( ~ )=~ F2N and, as also shown
in table 6, none of the chi-squares is significant.
Genetic stratification of the Zuni population might on a priori grounds, be either
spatial or temporal. The Zuni pueblo can
be divided into five areas corresponding to
locations separated by the Zuni River and
by the major roads through the pueblo.
These subdivisions are actually recognized
by the Zuni for administrative aspects of
their health care service. Since the Zuni
are matrilineal, live in extended family
units, and are subdivided into clans, there
might be opportunities for local genetic
differentiation. However, Kroeber ('19)
found no association between clan and
residence location and it appears that new
households are established without regard
to the location of other extended family
members. Genetic analyses by contingency
chi-square tests showed no significant differences among the five locations and it
seems that spatial differentiation can be
assumed to be absent.
Because of the recent rapid increase in
population size, the possibility of genetic
changes due to drift or differential fertility
should be considered. The Zuni sample was
sufficiently representative of the different
age groups that one can compare gene frequencies in the youngest group (age 13-19;
N = 208) and the oldest group (age 3 60;
N = 89. Table 7 shows such a comparison
for several of the genes studied. No large
differences can be seen, and none were
significantly different, so the possibility of
age stratification can be rejected. Thus, the
pooled Zuni frequencies can be assumed
to be representative of the contemporary
genome, and, indeed, to be a good approximation to the frequencies at least in the
past century.
The most striking observation in tables
4 and 5 is the high frequency of type B in
the ABO system. Evidence for non-Indian
intermixture in southwestern Indians usually is taken from the A1 and r (cde) genes.
Here, however, the frequencies of A1 and
r are 0.011 and 0.001 which suggests very
little non-Indian gene flow. On the other
hand, B, assumed absent in American Indi-
TABLE 7
Genefrequencies i n the y o u n g e s t (1 3-1 9 ; N =208)
and oldest ( 260; N = 289) individuals i n t h e
Zuni sample
Gene
13-19
60+
Gene
13-19
60+
P1
Fya
0.58
0.58
0.74
Hpl
Gc1
R1 (CDe)
R2 (cDE)
RO(cDe)
Rz (CDE)
0.65
0.87
0.67
0.81
0.70
0.24
0.02
0.04
Jka
Dib
B
PA
0.75
0.26
0.02
0.06
0.20
0.35
0.04
0.05
0.21
0.66
0.24
0.05
0.05
ans, has a frequency of 0.055 overall and
a frequency of 0.05 in individuals over 60
years old. If intermixture were the source
of the B gene, it is not likely to be due primarily to recent intermixture. In other
southwestern tribes, excluding Athapascans, only one report of a similar high B
frequency has been reported. Allen and
Schaeffer ('35) found B = 0.054 in Tewa
Indians at the Taos, New Mexico Pueblo;
however, in that group, A, had a frequency
of 0.192 and the evidence for intermixture
is clear.
In order to test for stratification with respect to the B gene the frequencies of genes
at other loci were determined for individuals with type 0 and type B. The more recent the intermixture, the more likely type
B individuals would have gene frequencies closer to Caucasian values, assuming
B to be derived from Caucasian gene flow.
No differences were found among B and 0
individuals and we must conclude that if
B came in by gene flow, it is of quite distant origin. The high frequency of B alone
among the non-Indian alleles suggests the
possibility that very few non-Indians (B, Rh
positive), and perhaps only a single individual, may have introduced the B allele
into the population. The demographic history of the Zuni would easily permit the
subsequent differential fertility and/or drift
required to elevate the B frequency to its
current level. Alternatively, postulating the
presence of B in the original founder population, while not impossible, has no support in any other study of southwestern or
Mexican Indian tribes.
GENETIC DIFFERENTIATION AMONG
SO U T H W E ST E R N I N D I A N TRIBES
The southwest contains by far the greatest number of American Indians within the
continental United States, including both
127
POPULATION STUDIES ON THE ZUNI INDIANS
Athapascans (Navajo and Apache) who migrated into the area as early perhaps as the
twelfth century and indigenous tribes
whose ancestors may have been in the region for 5,000 to 10,000 years. The current
reservation population sizes are given in
table 8. Most of the genetic observations on
these tribes were summarized by Matson,
Burch, Polesky, Swanson, Sutton and Robinson ('68). There are very few observations
on the Athapascans and the Yumans, and
even fewer on the Pueblos. Since we are
here concerned with biological differentiation among the indigenous groups, there
appear to be adequate genetic data only
for the Maricopa (Matson et al., '68), Pima
(Matson et al., '68). Papago (Niswander,
Brown, Iba, Leyshon and Workman, '70)
and the Zuni. Consequently, the analysis
to be presented is to be viewed only as a
preliminary study of genetic differentiation
in the southwest. The gene frequencies reported for these tribes are given in table 9.
Gene frequencies in all four tribes are
available only for the ABO, MNSs, Rh, P,
Fy and Jk systems. Using these loci, an
analysis of genetic variation among the
tribes can be carried out as follows. For k
alleles over s populations we form the ( s x
k) matrix V with elements
(PSk
- Pk)
&k
(1 - ck),
where Pk is the weighted mean gene frequency and Psk is the frequency of the
kth allele in the Sth population. Then R =
(l/k) VV' is an ( s X s) matrix whose diagonal elements, rii , describe what has been
termed local kinship (Morton, Yee, Harris
and Lew, 'i'l), that is, the deviation of the
ith population from the mean gene frequencies; larger values of rii indicate
greater relative isolation. The weighted
sum, S w ir ii, an estimate of mean kinship
within the total population, is an index
of the total genetic heterogeneity among
populations, where w denotes the subpopulation sample size relative to the total
sample. A correction for sampling error,
- (1 - h)/2N , where h is the fraction of
the census population sampled, and N T is
the total over all samples, must be added
to Hw ir ii. Here we use the hypergeometric
correction, assuming h = 0.10, to permit
inference on the contemporary gene pool
(Harpending and Jenkins, '73; Workman,
Harpending, Lalouel, Lynch, Niswander
and Singleton, '73) rather than binomial
sampling to draw inferences on the parental gene pool as done by Morton et al. ('71)
for kinship bioassay. The corrected mean
kinship, called here R St (rather than the
usual notation, F st, in order to avoid confusion with F taken as a probability), can
also be obtained from the average value of
the scaled gene frequency variances so that
,
where p i and up' are the appropriately
weighted mean and variance of the ith
allele.
The matrix R with corrected diagonal
elements is given in table 10. The Zuni
and Maricopa have much higher values of
r i i indicating greater genetic isolation. The
off diagonal elements of R, r indicate the
TABLE 8
Estimated Indian population on or adjacent to Reservations i n Arizona and New Mexico.
(Bureau of Indian Affairs estimates.for Sept., 1968)
Tribe or group
Estimated
population
Pima
Papago
10,240
6,210
Yuman
5,180
Apache
san Carlos
Fort Apache
Jicarilla
Mescalero
Navaho
4,650
6,000
1,500
1,690
13,840
1 19,500
Tribe or group
Estimated
population
Zuni
5,040
Hopi
6,080
Acoma
Laguna
E. Keresan
Tanoan
Towa
Tewa
N. Tiwa
S. Tiwa
Total Pueblo
1,920
2,880
2,400
1,380
2,400
1,270
2,210
25,580
128
P. L. WORKMAN, J. D. NISWANDER, K. S. BROWN AND W. C. LEYSHON
TABLE 9
G e n e f r e q u e n c i e s in t h e Z u n i , P i m a , Papago a n d Maricopa I n d i a n s
Gene
Zuni
+ Az
Pima
1
Maricopa
Pap ago 3r4
0.011
0.055
0.933
0.041
0.000
0.959
0.063
0.003
0.934
0.000
0.000
1.000
MS
MS
NS
NS
0.213
0.546
0.020
0.221
0.274
0.427
0.091
0.208
0.346
0.444
0.080
0.130
0.294
0.347
0.049
0.310
RO (cDe)
Rl (CDe)
RZ (cDE)
R z (CDE)
r (cde)
0.033
0.704
0.219
0.043
0.001
0.039
0.581
0.361
0.019
0.000
0.022
0.626
0.330
0.022
0.000
0.030
PI
0.559
0.742
0.296
0.023
0.661
0.860
0.538
0.709
0.359
0.029
0.543
0.533
0.762
0.476
-
-
0.494
0.820
0.360
0.028
0.452
0.832
0.099
0.031
0.037
-
0.020
0.005
0.227
-
AI
B
0
;;:
Dia
Hpl
Gc1
~fB0-l
A
W
L
~
~
~0.012
0.193
0.208
0.809
PA
PGM1'
5
0.607
0.317
0.046
0.000
-
Present study; N = 662.
Matson et al. ( ' 6 8 ) ; N = 909.
Niswander et aL ('70); N = 709.
4 Brown and Johnson ('70).
5 Matson et al. ( ' 6 8 ) ; N = 124.
1
2
3
TABLE 10
R m a t r i x f o r f o u r s o u t h w e s t e r n I n d i a n p o p u l a t i o n s a b o v e the diagonal; Sanghvi's
G2
b e l o w t h e diagonal
(1 1
I
(1) Zuni
0.0205
(2)
(4 )
(3)
- 0.0080
- 0.0087
(2) P i m a
0.0083
(3) Papago
0.0109
0.0044
0.0080
(4) Maricopa
0.0119
0.0055
0.0092
Mean kinship =
- 0.0048
0.001
0.0034
- 0.0042
I
0.0208
0.0109.
relationships between pairs of populations.
A genetic distance between populations i
and j could be taken as the squared Euclidean distance d $ = r i i
rjj - 2
rij . However, interdependencies among
genes suggest a more appropriate genetic
distance would be given by G2 (Sanghvi,
'53) or B2 (Balakrishnan and Sanghvi, '68).
The corrected values of R st per gene are
given in table 11. Only the B allele gives
values which appear to be much larger
than the mean. The high value of B at Zuni
+
TABLE 11
E s t i m a t e s of R,t p e r g e n e , corrected for
s a m p l i n g error
Gene
Rst
Gene
Rst
0.0122
0.0359
0.0051
0.0119
0.0121
0.0137
0.0132
R1 (cDe)
R1 (CDe)
R2 (cDE)
R z (CDE)
0.0008
0.0097
0.0151
0.0037
0.0018
0.0105
0.0066
~
A1
B
0
MS
MS
NS
Ns
Mean kinship ( E s t ) =
P
Fy"
Jk a
0.0109.
~~
POPULATION STUDIES ON THE ZUNI INDIANS
alone is also indicative of nonrandom differentiation for that allele. The low value
for RO (cDe) indicating very little differentiation may be due to chance or, perhaps,
to stabilizing pressures on this allele. No
interpretation of this observation can be
made with the present data.
The Indian populations in the southwest
are located in discrete sites at which year
round water is available. Intermixture
among these tribes might be related to the
distance separating them as already demonstrated for subdivisions within the Papago by Workman and Niswander ('70)
and Workman et al. ('73). The effect of isolation by distance can be studied given the
geographical distance between tribes and
the R matrix. Using the longitude and latitude of the main population centers of the
reservations (Sells and Sacaton for the Papago and Pima) as coordinates, the geographical locations can be plotted directly
in two-dimensions. A two-dimensional reduction of the R matrix can also be ob-
129
tained using the first two eigenvectors of
the centroid adjusted R matrix following
Lalouel('73). Although some information is
lost by the procedure, the first two eigenvectors maximize the dispersion of points
in two-dimensions. The proportion of the
total sum of squares retrieved by this reduction can be measured by f (A) = (A,
Az) I ZA i, where h are the eigenvalues of
the adjusted matrix. For the matrix of
table 8, f (A) = 0.914; that is, almost all
information is in the two-dimensional reduction. The two-dimensional reduction
can then be rotated to best fit with the geographical coordinates as suggested by Lalouel ('73). A measure of the goodness of
fit of the rotated projection of the R matrix
to geographic coordinate is given by Carrol's measure of disagreement, C , =
0.30 or, equivalently by the correlation between the coordinates of the configurations, 0.85 (see Lalouel, '73, for details).
The resulting projection is shown in figure 3.
+
goographic location
gonotic prodiction
Fig. 3 Two-dimensional reduction of the R matrix rotated to best fit with the geographical
locations of four southwestern Indian tribes (shaded areas labelled by tribe, delimit present
reservation boundaries).
130
P. L. WORKMAN, J. D. NISWANDER, K. S. BROWN AND W. C. LEYSHON
MI
0
4
Fig. 4 Two-dimensional reduction, plotting alleles in four southwestern Indian tribes using
first two eigenvectors of the S matrix.
The positions of the Zuni, Pima and
Papago based on the information on the R
matrix appear very close to their present
geographical locations which their ancestors inhabited for many centuries. The
Maricopa, today located just south of the
Pima reservation, were moved to that place
about 1890 from the upper Colorado River,
much to the northwest. The genetic prediction places them much closer toward
their ancestral location as should be expected if geographical location is an important
aspect of genetic differentiation. As also
seen in table 9, the largest genetic distance
is between the Zuni and Maricopa as would
be expected given their geographical distance in earlier years.
An alternative representation using all
the genetic markers given in table 9 was
made using Sanghvi’s G2, weighting the
contribution from each k-allelic locus by
(k - 1). That is,
G2.. = L [ Z G Z / ( k - l ) ]
U
L
where L is the number of loci.2 The G values so obtained are also given in table 10.
The fit by G2 did not differ substantially
from that by R and need not be presented
here. The close agreement of geographical
location and the location predicted by the
genetic model indicates that distance has
been a major factor in determining the
amount of gene flow among tribes.
Another reduction of the genetic data
which is also useful, plots the gene frequencies as points on axes directly comparable to the plot of the populations. The
genetic variation can be partitioned either
This differs slightly from formulation used for these
data by Sofaer et al. (Am. J. Phys. Anthrop., 37: 357366, 1962), in which each allele is weighted equally.
Although the results appear trivially different we prefer
the weighted formula because it gives slightly higher
correlation of thedistances in two dimensions with those
in the original n-space.
POPULATION STUDIES ON THE ZUNI INDIANS
by populations or by alleles, as was shown,
for example, in formula (1) for the derivation of R S t . That is, instead of obtaining
a dimensional reduction of R = l / k VV',
we consider S = (l/s)V'V which is a (k X
k) matrix; describing variation and covariation among allelic frequencies. Since both
R and S have the same set of nonzero
eigenvalues (see discussion by Harpending
and Jenkins, '73) the dimensions of any reduced space are the same whether we plot
populations or alleles. The two-dimensional
plot of the genes used in this analysis is
given in figure 4. By comparison of the
location of genes and populations in figures 3 and 4 we see that Zuni is typified
by its frequencies of B, R* and Ms; Maricopa by the frequencies of J k a , 0 and Ns,
and the Papago and Pima are characterized by their R 2 , A, NS, and MS frequencies
(see also the data in table 9). Note that
RO (cDe) which differentiates none of the
populations ( R s , = 0.0008) plots in the
middle of the array.
These two-dimensional representations
appear to provide the simplest and most
easily interpreted reduction of the data.
They do not use all of the information but
if, as in the present case, geographical
considerations are of major importance,
then a two-dimensional representation will
contain most of the information.
The analysis presented here indicates
that the Zuni have been relatively isolated
within the region, despite a general agreement of gene frequencies at most loci including the presence of the Albumin Mexico variant in all the tribes. Differences
such as presented by the B allele, indicate
for the Zuni, a generally low degree of gene
flow with the tribes included in this comparison at least since the time of contact
with the Spanish. The Zuni also have a
high frequency of albinism (1:240) which
has changed very little over the past 90
years (Witkop Niswander, Bergsma, Workman and White, '72), and only the neighboring Hopi have a similar frequency of
albinism indicating, perhaps, a degree of
Zuni-Hopi intermarriage. Since Brown,
Hanna, Dahlberg and Strandskov ('58),
found no evidence for B in a small sample
of Hopi, Tewa, the exchange between these
tribes may also pre-date the introduction of
B into the Zuni. When genetic studies of
the other Pueblos are available it is hoped
131
that some of the questions raised, but not
answered, in this present study can be resolved.
ACKNOWLEDGMENTS
The authors are indebted to the Zuni
people, their tribal leaders and the staff
of the Public Health Service Indian Hospital at Zuni, particularly Dr. Errett E. Hummel, Jr., Service Unit Director. We also
wish to acknowledge the technical support
of Mr. L. L. Taylor. This work was supported in part by grant lROlHG06003 from the
United States National Institutes of Health
while the senior author was at the Population Genetics Laboratory, University of
Hawaii, Honolulu.
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