American
Indian Demographic History and Cultural
Affiliation: A
Discussion of Certain Limitations on the Use
of mtDNA and Y
Chromosome Testing
By
Peter N.
Jones
The Bäuu
Institute
PO Box 4445
USA
pnj@bauuinstitute.com
1
Abstract
Mitochondrial
DNA (mtDNA) and Y chromosome studies have been used increasingly over the
last 20 years by
anthropological geneticists and others to reconstruct the peopling of the
Americas as well
as to infer American Indian cultural affiliation and demographic histories.
While the
promise of this method is great, there are several problems inherent in some of
its
current uses.
These limitations are discussed concerning the following six currently accepted
methods:
interpretation of coalescent times as times of origin; 2) the current uses of
haplogroups;
3) sample sizes;
4) use of language groups to define population groups; 5) use of contemporary
American Indian
reservations to infer prehistoric tribal history; and 6) a combination of these
to
determine
American Indian population history, historic migrations, and demographic
history.
This paper
concludes that caution must be exercised in claiming too much for the method.
Instead, it is
recommend that it be used in conjunction with other established sources of data
such as oral
history, ethnography, linguistics, and archaeology when attempting to
reconstruct
American Indian
cultural history or in determining the cultural affiliation of groups.
Key words
Genetic
anthropology, haplogroups, demographics, cultural affiliation, American Indians
2
Introduction
Anthropological
genetics is one of the more recent sub-fields in the study of American
Indian cultural
affiliation and demographic history, dating back some 60 years (see Matson and
Schrader 1933;
Matson 1938). However, in the last 20 years advances in technology have
allowed
anthropological geneticists to explore the origin of modern humans, the size and
geographic
origin of human populations, as well as the possibility of finding ethnic or
even
individual
people’s homelands. Recent articles using genetic data have claimed to link
Europe/Western
Asian populations with North American Indian populations as having recent
common ancestry
(Brown et al. 1998), identified a single wave of migration for the peopling of
the New World
(i.e., Bianchi et al. 1997; Easton et al. 1996; Merriwether et al. 1995) as
opposed
to several
(i.e., Karafet et al. 1997), and concluded that some American Indian tribes
recently
moved into a
geographic area (Kaestle 1997; Kaestle and Smith 2001) despite contrary evidence
from oral
history and archaeology. Some of the most publicized uses of genetic
anthropology in
recent years
concern the question of the peopling of the Americas, and in compliance with the
Native American
Graves Protection and Repatriation Act (NAGPRA), as seen in such cases as
the Spirit Cave
Mummy and the Kennewick Man repatriation controversies. In these two
examples the
situation is complicated by the great antiquity of the skeletons, 9,415+/-25
years
ago and
8410+/-60 years ago, respectively (Napton 1997; Chatters 2000). The potential
benefits
this new genetic
research has to offer are vast and highly valuable. Such benefits include a
better
understanding of
the genetic and evolutionary factors that influence populations; an
understanding of
maternally transmitted diseases such as blindness, epilepsy, dementias, cardiac
and skeletal
muscle diseases, diabetes mellitus, and movement disorders; the development of
new metabolic
and genetic therapies for mitochondrial diseases; and a better understanding of
3
the geographic
origin of anatomically modern humans, to name just a few. However, caution is
urged in
applying the methods and results of present genetic research to the use of
American
Indian cultural
affiliation and demographic histories. This paper examines six weaknesses
inherent in
current uses of genetic anthropology that attempt to resolve questions of
demographic
history and
prehistoric cultural affiliation: 1) interpretation of coalescent times as times
of origin;
2) the current
uses of haplogroups; 3) sample sizes; 4) use of language groups to define
population
groups; 5) use of contemporary American Indian reservations to infer prehistoric
tribal history;
and 6) a combination of these to determine American Indian population history,
historic
migrations, and demographic history.
I. Coalescent
Times as Times of Origin
A first problem
confounding the uses of mtDNA (mitochondrial deoxyribonucleic acid)
and Y
chromosomes to infer American Indian demographic histories is in interpreting
the
coalescent times
of genes as times of origin for the population. Although tracing the genealogy
of mtDNA
theoretically can lead to a single common ancestor, this is not evidence that
the
human population
went through a period when only one breeding population was alive and
reproducing.
Tracing the coalescent times leads to one ancestor of a unilineally transmitted
set
of markers, but
the descendents of the original mtDNA will have had haplotype frequencies that
differed among
themselves, resulting in a biased sample of the total historic population when
using coalescent
times. This is so because working back in time is does not allow one to take
into account the
various branches of diversity that the historic population had, but only can
detect
the lineal
history of the specific marker being coalesced. Three primary assumptions
arising
4
from the use of
coalescent times (Hoelzer et al. 1998; Hudson 1990; Templeton 1993; Wolpoff
1999) that have
been employed in understanding American Indian demographic history are:
A) gene
coalescence is a regular process of mutation accumulation in neutral systems,
and therefore
can be timed like a regularly ticking clock with an acceptable range of
error,
B) American
Indian populations were isolated from each other after they originated or
migrated to the
Americas, and
C) the history
of particular gene systems is the history of the specific populations in
which they are
found.
Prior to the
historic period, and especially before the formation of reservations beginning
in the 1850s,
many American Indian groups were highly mobile autonomous entities, covering
large areas of
land. Similarly, many American Indians practiced a high degree of spousal
exchange and
intergroup marriage among other groups in order to solidify trade arrangements
and political
alliances. Some of these exchanges took place well over 500 miles from where the
group has been
historically recorded to inhabit. Examples of these trade centers are the large
Native fisheries
of the Northwest Coast such as The Dalles, Celilo Falls, and the Lillooet River
Fishery
(Schuster 1998; Stern 1998; Hayden 1992) where groups from the Northwest Coast,
Plateau,
Northern Plains, and Great Basin regions gathered. Other examples can be found
from
the
archaeological record that show similar large regional centers that may have
acted as
gathering and/or
redistribution centers such as Chaco Canyon in the US southwest (Lekson
2000) and Monte
Alban, San Jose Mogote, Tlapacoya, and Tlatilco in central Mexico (Flannery
and Marcus
1994). In fact, as Walker (1998:5-6) has noted for the Plateau peoples, “It is
clear
that Plateau
peoples were and remain highly inter-active maintaining extensive intergroup
5
connections as
well as extensive linkages with the Plains, Northwest Coast, and Great Basin
groups.
Connections with Subarctic groups are evident in the northern reaches of the
Plateau of
Canada.”
One important
requirement in the coalescence theory is the use of random samples of
genes from the
population under study. However, most studies have not used random samples,
but instead have
used convenience samples obtained from diabetic studies, rheumatic studies,
and AIDS
studies, as well as other studies (Jones, in preparation; also see below). As
Donnelly
and Tavare
(1995: 418) point out, “In practice, genetic data are typically obtained from
convenience
samples rather than proper random samples. There is an obvious danger that such
data may contain
individuals who share relatively too much ancestry on the relevant timescales.
The extent to
which application of coalescent (or traditional) methods to such convenience
samples may be
misleading remains an open, and potentially serious, question.” Furthermore,
most studies
rely on the idea that American Indians came over in small groups (usually
thought
to have occurred
as part of one to three migration waves; see Dillehay 2000 to cite one recent
work) across the
Bering Land Bridge in prehistoric times. If this is the case, coalescence times
will be shorter
because smaller populations in the past are more likely to share ancestors
(Donnelly and
Tavare 1995: 410), and thus lead to an accelerated time of origin for American
Indians and thus
not truthfully demonstrating the occupational time depth American Indians have
in the Americas.
Furthermore,
departures from random mating due to inbreeding, assortative mating, or
population
stratification can lead to non-random association between genotypes and further
complicate the
interpretation of the data and coalescent times. One such example is the
well-
documented
moiety and clan system among the Tlingit peoples of southern Alaska. Among the
6
Tlingit,
marriage was always with a member of one’s opposite moiety, and preferably with
a
member of the
father’s clan and house (De Laguna 1975). Therefore, the Tlingit as well as many
other American
Indian groups of the Northwest Coast and other regions practiced a highly
selective,
non-random form of mating that could influence the genetic data (see Handbook
of
North
American Indian series published by the Smithsonian Institution). There is
also a growing
body of evidence
suggesting that there could have been various forms of admixture between
American
Indians, Japanese, and Russians during the last 500 years (Quimby 1985; van
Stone
1984; Boyd
1999), not to mention known examples of admixture during the historic period
with
trappers, fur
traders, explorers, and other Europeans. Likewise, Karafet et al. (1997)
concluded
that because of
the presence of the 1T haplotype (a Y chromosome combination haplotype [see
next section for
a discussion of haplotypes]) in both northeastern Siberia and the Americas, the
possibility of
historic and prehistoric back-migration is extremely likely. Similar studies
have
also noted the
possibility of gene transfer or the “hitch-hiking theory” among American Indian
and Asian
populations (Bianchi et al. 1997; Bradman and Thomas 1998; Hudson 1990). Because
population-coalescence
times are frequently a result of the fusion of several of the ancient
phylogenetic
clusters and not the age of individual populations (Watson et al. 1997), faulty
results may be
reported. It is evident that neither American Indians nor specific American
Indian
groups were ever
isolated populations and that the history of a contemporary group’s genes are
not a specific
history of that American Indian population. Therefore, using gene coalescent
times as
possible times of origins for American Indians can lead to spurious conclusions,
for
there is no
evidence that American Indians were ever: 1) part of a neutral system that can
be
timed like a
regularly clicking clock, 2) were isolated from each other or from Asian
populations,
and 3) that the
current genes systems found in a particular population fully represent the
diversity
7
and history of
that population.
II. Current
Uses of Haplogroups
Although it has
been noted that limitations exist when studying only one gene (Chen et
al. 2000;
Karafet et al. 1997; Mountain and Cavalli-Sforza 1997), most studies still rely
on only
one gene and its
alleles because of the ease in identifying differences in a restricted location
on
that gene,
especially in non-recombining genes such as mtDNA. The allele sequences that
are
studied are
called haplotypes, which for American Indians presently fall into five
recognized
haplogroups (A,
B, C, D, and X), and have been used in most studies concerning American
Indian
population genetics.
One of the
current limitations with the uses of haplogroups for inferring American Indian
cultural
affiliation is that there is the possibility of discovering new haplotypes as
more tribes are
studied and
techniques develop (Easton et al. 1996; Karafet et al. 1997; Schurr et al. 1990;
Smith
et al. 1999). By
testing only for known haplotype frequencies, it is likely that other haplotypes
will go
undetected, resulting in spurious conclusions from simplified haplotype
frequencies.
Along with the
possibility of new haplotypes being discovered, it is known that many
prehistoric
American Indian
groups were not stationary and the use of within-local-population frequencies
for the genetic
sequences, highly affected by each population’s specific recent demographic
history, and
thus when relying on the use of haplogroups, scientists will probably
underestimate
the nucleotide
diversity of American Indians as a whole (Bonatto and Salzano 1997). Therefore,
the differing
results between CR (control region) sequences and RFLP (restriction fragment
length
polymorphism) data cannot be explained either by sample size or attributed to
the
different ways
in which the haplotype frequencies were treated, but are more probably due to
the
8
different
populations or regions of the mtDNA studied. Furthermore, the only changes
introduced in
genes are point mutations, insertions, and deletions (with insertions and
deletions
being rare in
comparison to point mutations). This means that each of the four possible
founding
lineage clusters
can be thought of as containing the founding lineage haplotype plus a collection
of that
lineage’s descendants. However, as has been noted for the Y chromosome, the
original Y
chromosome can
eventually die out, shifting time, haplotype frequency, or relationships
(Bradman and
Thomas 1998) and can result in faulty data when comparing present American
Indian tribal
frequencies to those of ancient American Indian haplotype frequencies. As
Bradman and
Thomas (1998) pointed out using the insertion of the YAP (Y chromosome alu
polymorphism)
indel (insert) on the Y chromosome, descendents of individuals after only one
generation may
not carry the same Y chromosome alleles. It is possible that a descendent of the
individual who
first acquired the YAP indel may lose that indel, yet still remain a descendent
of
that
individual. This is also possible with mtDNA, where a father’s son or daughter
will not
carry the
genetic information of that person’s father’s mother. By only looking at
specific
alleles,
mutations, insertions, and deletions can be viewed as coming from discontinuous
populations.
Likewise, “the combination of a decrease in the effective population size and
genetic
hitch-hiking may have been the cause producing a single variety of Y-chromosomes
in
the earliest
ancestors of extant Amerindians,” (Bianchi et al. 1997: 87) which would result
in
faulty results
in determining affiliation between American Indian groups. Similarly, because
the
mitochondrial
genome undergoes no recombination, the 16,569-bp genome behaves
evolutionarily
as a single locus. As MacEachern (2000: 358) recently notes, “In particular, it
appears that
there may be significant variability in selection mechanisms on the genome
itself
and in the
mitochondria and in rates of phylogenetic versus intergenerational mtDNA
mutation
9
that are only
now being appreciated (Gibbons 1998; Parsons, Muniec, and Sullivan 1997).”
Therefore,
inferences from any one such locus lack robustness (Pamilo and Nei 1988). As
noted
above, because
of the potential inaccuracy in using a constant molecular clock, estimates of
mutation rates
are going to be imprecise (Donnelly and Tavare 1995; Hoelzer et al. 1998).
Because of the
high mobility of American Indian groups in the prehistoric, along with examples
of intergroup
marriage and non-random mating, there is ample reason to believe that the
genetic
history of
American Indians is much more complex then the current five haplogroup
frequencies
lead us to
believe.
III. Sample
Size
Many of the
discrepancies and much of the unreliability of the data employed in
American Indian
genetic studies lies in the sample sizes of the populations used. Variations in
population size
are commonly attributed to bottlenecks and the so-called founder principle in
which a
population encounters a severe reduction in size or a few individuals colonize a
new area
resulting in a
small selection of gene frequencies as compared with the original population.
However, an
important complication that makes it impossible to determine census size of a
prehistoric
human group as a direct estimate of the effective population size is that human
populations
have overlapping generations. Rogers and Jorde (1995: 1-36) have shown that the
only sense in
which sequence diversity can be employed as a measure of age is as an estimation
of the time
during which a particular population has expanded after experiencing a severe
bottleneck.
This is because we are dealing with alleles (haplotypes), and not with distinct
populations. In
fact, the error variance increases with time and the earliest observations are
the
most precise.
Computer simulations that suggest that the four major haplogroups found among
10
American
Indians underwent a bottleneck followed by a large population expansion may be
questioned.
These simulations are based primarily on the analysis of CR sequences from
haplogroup A
and do not take into account haplogroups B, C, D, and X (as well as the
possibility
of future
haplogroups being discovered). Similarly, although most studies on the problem
of
dating the
original occupation of the Americas have used sequence diversity as a measure of
age,
few have
investigated whether their samples met the very stringent assumptions required
by this
practice
(Bonatto and Salzano 1997: 1417). Furthermore, Bonatto and Salzano (1997: 1417)
have also noted
that studies using RFLPs found that haplogroup B had a much lower diversity
than the other
three (A, C, D) which would lead to inaccurate computer simulations. Based on
this, the
current dates from mtDNA and Y chromosome studies contending that American
Indians arrived
in the “New World” around 35,000 years ago can be questioned (Bonatto and
Salzano 1997;
Brown et al. 1998). This number is actually the time during which American
Indians
theoretically experienced an expansion after a bottleneck. However, it is
unknown if this
bottleneck took
place in Asia, the generally accepted origin of American Indians, or in the
Americas after
their arrival, nor is it known what effects migrations and subsequent
bottlenecks
from disease
and other factors have on this time estimation. Therefore, the date of 35,000
years
ago could be
the time one group of American Indians entered the “New World” or when a group
experienced a
bottleneck in Asia and subsequently entered the Americas, or any number of other
possible
scenarios.
Another problem
with the current sample sizes being used is the actual numbers of
individuals
tested to infer the genetic makeup of the entire population. Typically, sample
sizes
range between
four and 30 individuals per tribal population; this is insufficient to detect
little
more than the
most common haplotypes in each population. Although it is necessary to have
11
genetic samples
from 50 males or 50 females of an individual population to accurately infer
genetic
demographic history, no study has done this (Wells 2000). The largest study to
date on
American
Indians dealt with 2,198 males from 60 global populations, including 20 American
Indian groups
(Karafet et al. 1999; this study relied on large amounts of data gathered from
previously
published reports, and thus could not correct for those sample sizes). However,
only
the Inuit
Eskimo and Navajo samples were over 50 at 62 males and 56 males respectively.
All
others ranged
from as high as 44 to as low as two individuals. It is unrealistic to assume
that one
can get an
accurate picture of a tribe’s genetic frequencies using only two males. In fact,
Weiss
(1994: 834)
suggests that we may not be able to distinguish loss of lineages after one
migration
or from
separate migrations from a common source population, thus further stressing the
critical
need for
adequate population sample sizes. A clear example of the importance of sample
size is
seen in Easton
et al.’s (1996) study and Torroni et al.’s (1993) study on the Yanomamo. In
Easton et al.’s
sample they detected both haplotypes X6 and X7, but in Torroni et al.’s sample
from a
neighboring village they did not detect any of these two haplotypes. As Ward et
al.
(1993) have
noted, a sample size of 25 will detect ~63 percent of the lineages in a tribe
with
normal
diversity. In tribes with extensive diversity a sample size of 25 individuals
will only
detect ~40
percent of the lineages and sample sizes of 70 or above are required to detect
two-
thirds of the
lineages. The fact that the majority of studies lack the required sample sizes
necessary to
detect even 63 percent of the lineages in a normally diverse tribe brings into
question many
of the results of these studies, especially when it has been noted that most
American Indian
tribes are believed to have a high level of diversity (Ward et al. 1993).
In the past, as
now, choice of mates is largely dictated by geographic, socioeconomic,
religious,
ethnic, and other constraints. This has the effect of subdividing and
stratifying the
12
gene pool of a
population in very complex ways. Likewise, migration is also difficult to
reconstruct
from mtDNA and Y chromosomes. The most meaningful measure of migration from
a genetic point
of view is obtained by taking the generation as the time unit. Measuring the
distribution
between birthplaces of parent and offspring theoretically can yield a
statistical
measure of
migration. However, this method works only for a continuous model in which the
population is
constant, and is not entirely satisfactory when the population is highly
clustered as
is believed
most prehistoric American Indian populations were (Cavalli-Sforza and Bodmer
1971:433). A
similar limitation in using such data to infer migrations is that exchange
between
non-neighboring
clusters is frequent enough among American Indians to violate the rules of the
simplest
stepping-stone models (Cavalli-Sforza and Bodmer 1971:433).
Another aspect
of human DNA confounding many of the current uses of this data to
reconstruct
hypothetical demographic histories is that human mtDNA variation is high.
Likewise,
genetic variation within populations is much greater than between
populations
(Walpoff
1999:551). What this means is that mtDNA evolution, and possibly the evolution
of
other genetic
systems, is not the same as the evolution of particular populations. As Scozzori
et
al. (1999) have
noted, groups or tribes thought to have descended from a common ancestor more
than 10,000
years ago may have lost even their shared-by-descent portion of their gene pool
and
can no longer
be detected as affiliated through genetic analysis. Likewise, population
specific
mutations and
the gene trees inferred from these sequences are generally inconsistent with
historic and
prehistoric population affiliation. Page and Charleston (1990) have identified a
method for
visualizing and quantifying the relationship between a pair of gene and species
trees
that constructs
a third, reconciled tree. Reconciled trees use a more critically optimal method
for
mapping the
combined history of genes and populations. However, even this more accurate
13
method of
depicting gene and population trees has limitations such as allele phylogenies
and
horizontal
transfer, neither of which has been addressed in studies concerning American
Indian
demographic
history. In fact, many of the polymorphisms observed for mtDNA probably
predates
population separations (Mountain and Cavalli-Sforza 1997) and would not be
useful in
constructing
genetic, population, or reconciled trees. Mitochondrial DNA or Y chromosome
lineages are
not human populations. In order to estimate the significance of variation of
gene
frequencies
between groups, it is necessary to estimate how large a sample must be in order
to be
representative
of the group. This can only be accomplished if an accurate estimate of the real
variation to be
expected in the gene frequencies is possible. This estimation is valid only for
genes without
dominance, in which case genes can be counted. However, if people in the sample
from a given
tribal village or town are closely related, a single source of variation may
greatly
inflate the
estimate of variance between populations (Cavalli-Sforza and Bodmer 1971:422).
Multivariate
analysis, or the use of more than one trait or gene, which is presently the most
commonly
employed method of analysis, poses more difficult problems in that one must
determine the
maximum number of genes possible for each population in order to be accurate.
Unfortunately,
many authors have tested only a small set of markers on one gene (univariate)
for
their studies
(Cavalli-Sforza et al. 1994: 22), combining their data with those of others to
result
in several sets
of markers to arrive at their multivariate analysis. Not only have limited
numbers
of markers been
studied and subsequently combined with other studies (which was noted above),
but the
mutation rate for insertions and deletions on those markers is unknown.
14
IV. Language
Not only have
sample sizes of groups or tribes being tested been inadequate, but most
studies have
relied on the use of controversial linguistic phyla in order to place their data
into
objective,
quantifiable groups. However, as several papers have pointed out, not only do
the
correspondences
between languages and populations differ (i.e., Barbujani 1997; Karafet et al.
1999; Scozzari
et al. 1999; Schurr et al. 1999), but there is no agreed upon set of linguistic
phyla
for American
Indians (Greenberg et al. 1986; Greenberg 1987; Bateman et al. 1990; O’Grady et
al. 1989;
Ruhlen 1987, 1994). Most studies use several linguistic phyla that are subject
to
serious
criticism, such as Altaic s.l., Austric s.l., Indo-Pacific, Amerind
(sensu Greenberg 1987),
and Na-dene
s.l., which are in turn awarded equal status as more accepted phyla from other
parts
of the world
such as Sino-Tibetan, Indo-European, and Dravidian (Bateman et al. 1990).
Furthermore, it
has been noted that “given 56% correspondence between linguistic phyla and
population
aggregates at the coarse level of resolution, 11% correspondence at the fine
level, and
the poor
integrity of both superphyla, the parallelism between the genetic and linguistic
entities
does not strike
us as especially ‘remarkable’” (Bateman et al. 1990:7). Likewise, “there is an
important
problem of time scales involved in this work, since at this point neither
genetic nor
linguistic
research can lay claim to chronometric techniques comparable in precision to
those
used by
archaeologists and historians” (Pluciennik 1995:44-45). Languages do not change
at
specific rates
and therefore using contemporary linguistic phyla to extrapolate prehistoric
population
groups is ill-founded. For example, in 1995 there were approximately 209 native
North American
languages still spoken, close to only half the number that existed five hundred
15
years earlier
(Goddard 1996). Similarly, of the Eastern Algonquian languages, only seven were
spoken in 1970
out of a total of 20 from 200 years earlier (Goddard 1978).
Many of the
researchers conducting genetic tests have noted the discrepancies between
linguistic
phyla and genetic phyla. Scozzari et al. (1999) concluded that geography is a
better
method for
identifying affiliation then linguistics. Likewise, Poloni et al. (1997)
concluded that
genetic data is
more accurate and useful for distinguishing between linguistic phyla than
between
populations in
the same language family. Finally, Schurr et al. (1999) noted that populations
on
the Kamchatka
peninsula were genetically similar based on geography but quite divergent when
compared to
linguistically related groups. Therefore, the use of linguistic phyla may be
useful
when studying
the differences between language phylas (i.e., between Na-Dene and
Eskimo-
Aluet), but not
as useful when studying groups within the same language phyla (i.e., Yakama and
Nez Perce). Not
taking into account the current discrepancies between American Indian
language phylas
can lead to several different conclusions depending on how the linguistic and
genetic data
are combined. For example, Karafet et al. (1997) found that Y chromosome
markers did not
agree with the linguistic phyla proposed by Greenberg et al. (1986) for the
peopling of the
Americas. However, in a later study using different Y chromosome markers
Karafet et al.
(1999) did agree with the linguistic phyla proposed by Greenberg et al. (1986).
Other studies
have arrived at similarly contradictory conclusions (see Schurr et al. 1999;
Poloni
et al.
1997).
To use current
American Indian languages as a baseline for prehistoric American
Indian genetic
affiliations and population groups seems presumptuous. Until linguistic
specialists
agree upon the classifications of American Indian languages, they should not be
used
as a means of
inferring and objectifying prehistoric population groups.
16
V.
Contemporary American Indian Reservations and Demographic
History
As noted above,
the current sample sizes of most studies fall far short of a reasonable
number of
individuals being tested to be considered an accurate data set of the
population.
However,
besides the limitations arising from the small sample sizes, as well as those
discussed
concerning the
present use of linguistic phyla, there are even greater problems lying in what
the
studies
consider populations. Presently, studies concerning American Indian cultural
affiliation
and demographic
history test individuals from a reservation and combine their allele frequencies
to arrive at
the haplotype makeup of that population. Therefore, the researchers are using
contemporary
American Indian reservation demographics to arrive at a population that they
then
infer back into
prehistory. However, one of the primary problems with this method is that most
contemporary
American Indian reservations are not made up of a single group, but consist of
several
different groups of American Indians that prior to being forced onto
reservations were
autonomous
groups. For example, Merriwether et al. (1995) use samples from Haida, Dogrib,
and other
contemporary American Indian reservation groups which they consider as one
population
group. However, the Dogrib as a whole tribe were prehistorically made up of
several
different bands
that occupied a large area in the Northwest Territories, Canada, between the
Great Slave
Lake in the south to the Great Bear Lake in the north and from the lowlands on
the
east side of
the Mackenzie River to Contwoyto, Aylmer, and Artillery Lakes. The Dogrib are
known to have
had regular contact with the Bearlake Indians, the Slaveys, Chipewyans, and
occasionally
Eskimos (Helm 1981: 291). Likewise, the Haida, along with other Northwest Coast
tribes were
known to have traded slaves up and down the coast. The Haida traded slaves they
acquired from
the Kwakiutl with the Tlingit (Blackman 1990). Other such examples can be
found in the
studies by Smith et al. (1999), Karafet et al. (1999), Lorenz et al. (1996), and
Brown
17
et al. (1998)
that use contemporary reservation groups as prehistoric population groups. Such
contemporary
groups as the Yakama and Apache are good examples. The present Yakama
reservation in
Washington is made up of at least five different groups that were
prehistorically
independent
bands or groups (Schuster 1998). Similarly, there is still much disagreement
among
American Indian
specialists as to how many different Apache groups there were prior to the
arrival of
Euroamericans. Currently there are seven recognized Southern Apachean speaking
groups:
Chiricahua, Jicarilla, Kiowa-Apache, Lipan, Mescalero, Navajo, and Western
Apache.
However,
depending on “how much more extensive their territories are conceived to have
been
in the past
depends upon one’s view of claims that the Querechos, Vaqueros, Teyas, Janos,
Jocomes,
Mansos, Sumas, Cholomes, Jumanos, Cibolos, Pelones, Padoucas, and various other
groups named in
early Spanish and French records were Apacheans” (Opler 1983:368). Finally,
over the last
hundred years reservation populations have been greatly affected by outmarriage
with other
tribal groups and marriage with non-Indians. An example of this change can be
seen
in a study done
by Walker (1990; see also Walker 1972) for the Confederated Tribes of the
Umatilla Indian
Reservation (CTUIR). This study showed that in 1990 54 different tribes were
represented in
the blood of CTUIR individuals (see Table 1). Furthermore, one CTUIR
individual had
various amounts of Cayuse, Walla Walla, Umatilla, Nez Perce, Snohomis, and
non-Indian
blood, while another individual had Umatilla, Cayuse, Walla Walla, Yakama, Nez
Perce,
Quinault, Snoqualmie, Cascade, and non-Indian blood. It is evident that the
population
groups current
studies are using to infer American Indian cultural affiliation and demographic
history are not
acceptable. One cannot use contemporary allele frequencies from a few
individuals of
a contemporary American Indian reservation to arrive at an unequivocal haplotype
for that group,
either presently or prehistorically.
18
Table 1:
Blood Types found in CTUIR Tribal Members
Tribal
Affiliation
Geographic
Proximity
Geographic
Location
Alaskan
Distant
Arctic
Arikara
Distant
Plains
Assiniboin
Distant
Plains
Bannock
Distant
Great Basin
Blackfoot
Distant
Plains
Canadian
Distant
Cascade
Neighboring
Plateau
Cayuse
Official Blood
Line
Plateau
Cherokee
Distant
Plains
Cheyenne
Distant
Plains
Chippewa
Distant
Plains/Subarctic
Chocktaw
Distant
Southeast
Cochiti
Distant
Southwest
Coeur d’Alene
Neighboring
Plateau
Colville
Neighboring
Plateau
Cowichen
Distant
Coast
Cowlitz
Neighboring
Plateau
Cree
Distant
Plains/Subarctic
Crow
Distant
Plains
Flathead
Neighboring
Plateau
Grande Ronde
Distant
Coast
Hopi
Distant
Southwest
Klamath
Neighboring
Plateau
Klickitat
Neighboring
Plateau
Kootenai
Neighboring
Plateau
Laguna
Distant
Southwest
Lummi
Distant
Coast
Makah
Distant
Coast
Modoc
Distant
Coast
Muckleshoot
Distant
Coast
Navajo
Distant
Southwest
19
Nez Perce
Neighboring
Plateau
Ottawa
Distant
Northeast
Paiute
Distant
Great Basin
Palus
Neighboring
Plateau
Pawnee
Distant
Plains
Puyallup
Distant
Coast
Quinault
Distant
Coast
Sac and Fox
Distant
Plains
Seminole
Distant
Southeast
Shoshone
Distant
Great Basin
Siletz
Distant
Coast
Sioux
Distant
Plains
Snohomish
Distant
Coast
Spokane
Neighboring
Plateau
Tulalip
Distant
Coast
Walla Walla
Official Blood
Line
Plateau
Warm Springs
Neighboring
Plateau
Wasco
Neighboring
Plateau
White Mountain
Apache
Distant
Southwest
Winnebago
Distant
Plains
Wishram
Neighboring
Plateau
Yakama
Neighboring
Plateau
A further
problem in the use of contemporary American Indian reservations can be found
in the use of
ancient DNA (aDNA). Several reports have used aDNA to construct ancient
populations
that are then compared to present American Indian reservation populations. These
studies are
plagued by many of the limitations already noted such as sample size,
demographic
histories, the
possibility of mutation addition or deletion, and other factors. Kaestle (1997)
attempted to
compare an ancient population from western Nevada to those of contemporary
tribal
populations in the region through haplogroup frequencies. However, Kaestle’s
ancient
20
population
spanned 5000 years in time. A genetic sample dating to 5905+/-125 years BP
cannot
be considered
part of the same population as a genetic sample dating to 860+/-75 years BP
without also
automatically designating contemporary American Indians as part of that
population.
“Ancient DNA samples are not populations in the traditional sense of the term.
The
individual
specimens that constitute aDNA samples may span several centuries and even
geographic
space. Thus they are the equivalent of sampling an individual every few
generations
to characterize
a continuous population” (O’Rourke et al. 2000). This is especially true when
these samples
are then compared to contemporary American Indian reservations that each have
their own,
unique demographic history. Likewise, the use of aDNA models to reconstruct a
population
assume (or cannot accurately model) that these ancient populations were somewhat
isolated, and
that these populations did not practice forms of intergroup or outgroup
marriage.
However, as
previously noted for the Northwest Coast and the Plateau regions, highly complex
forms of
intergroup marriage have been practiced for centuries. Complex forms of
intergroup
marriage have
also been documented for Great Basin tribes (see D’Azevedo 1986). Not only has
there been
extensive intergroup marriage within regions, but also between regions as noted
in
Walker (1972,
1990).
Finally, it
should be noted that many of the scientists conducting these studies acquire a
large
proportion of their blood samples or genetic material through convenience
samples.
Convenience
samples means that the blood samples or genetic material were not collected by
the
investigating
scientist, but instead through third parties. Many of these third parties
initially
acquired the
blood or genetic material for other reasons, such as diabetes testing. A review
of
the literature
has revealed that over a hundred institutions have allowed these scientists
access to
American Indian
blood, a lot of the time without the individual who gave the blood having any
21
knowledge of
this. Though there are many problems with this in and of itself, the point that
is
important in
the present discussion is that the scientists have no means of verifying the
actual
tribal
affiliation of the blood sample they are using. For example, when an individual
goes in for
diabetes
testing, they designate themselves and their tribal affiliation, through there
is no
guarantee that
this designation is correct, nor is there any knowledge of that individuals
family
genetic
history. This fact could greatly mislead the scientists into concluding various
tribal
haplotype
frequencies that may not be correct.
VI. Genetics
and American Indian Cultural Affiliation
The fact that
most studies have not addressed the above concerns is only part of the
present problem
with applying anthropological genetics to American Indian demographic history
and cultural
affiliation. In fact, no studies concerning American Indians have seriously
taken
into account
the demographic history of the last 500 years when Euroamericans arrived in the
“New World.”
Almost every contemporary American Indian tribe or group in the Americas has
experienced
severe epidemic diseases, depopulation, acculturation, and displacement from
their
native lands.
These factors have caused some tribes to disappear, others have experienced
population
fluctuations greater than 80 percent, and some have been displaced from their
home
land by
hundreds of miles (Boyd 1999; Dobyns 1983; Ehle 1988; Jones 2002). For example,
the
American
Indians of California numbered upwards of 310,000 during the eighteenth century,
but
by the turn of
the twentieth century the native population had dropped to 20,000 (Cook 1978).
Similar
population declines are known for the Plateau with a loss of approximately
20,000
Natives between
1805 and 1860 (Boyd 1990), the Northwest with a population decline from
approximately
200,000 Natives in 1774 to 40,000 in 1874 (Boyd 1990, 2000), as well as other
22
regions.
Furthermore, admixture with historic Europeans and Euroamericans such as fur
trappers,
explorers, African Americans, and earlier settlers must be accounted for; it is
well
known, though
not well documented, that many of these non-indigenous peoples married or
mixed with
American Indians.
Any mtDNA or Y
chromosome study that attempts to date the first appearance of a
particular
population in a certain geographical area as big as the Americas or as small as
the
Great Basin
should be based on extensive sampling, not only of the population under
consideration
but also of potential source populations and neighboring populations. A
phylogenetic
analysis can then theoretically identify the putative founder sequences. This
requires that
the mutations distinguishing these founders must be disregarded in order to set
the
evolutionary
clock to zero to coincide with the arrival of a particular population in the new
area.
Considerable
scientific care must be exercised in choosing realistic demographic models to
describe the
process adequately. For example, current models that assume constant population
size and random
mating would be unrealistic in most situations for the following reasons: 1)
arrival in a
new area is likely to trigger subsequent population expansion for many
generations;
2) for a
population on the move, the demographic pressure is more relaxed, allowing
population
sizes to
fluctuate rather freely, without statistically conforming to an expected
(time-dependent)
size; 3) over a
period of tens of thousands of years, environmental conditions often change
drastically,
obviously influencing the effective population sizes; and 4) ongoing gene flow
from
neighboring
populations will inevitably distort the estimation of nucleotide diversity, and
thus
the coalescence
time estimate (Forster et al., 1996: 944).
23
Conclusion
Although mtDNA
and Y chromosome studies can provide insights on America Indian
origins and
prehistoric relationships, they should be used with caution. Mitochondrial DNA
and
Y chromosome
studies are in their infancy. Because of the various limitations listed above,
as
well as a lack
of correlation between anthropological genetic data, archaeological data,
ethnographic
data, and oral tradition, these studies should be viewed as inchoate and
requiring
further
investigation and support from the other fields of anthropology. Current
controversies
surrounding
Spirit Cave Man and Kennewick Man should not be resolved only by mtDNA or Y
chromosome
testing. The mtDNA and Y chromosome data for American Indians, as well as
many other
regions throughout the world, have serious limitations. However, because of the
claimed
authoritative validity of these studies there is great danger that they will
convince non-
specialists of
the validity of the hypothesized associations between American Indian groups.
The
non-specialists
would be ill-advised to rely on the claims put forth by mtDNA and Y
chromosome
inferred American Indian studies because they take little account of the vast
majority of
ethnographic, linguistic, historical, and archaeological research. However,
archaeologists,
ethnologists, linguists, and historians should take note of some of the
inferences
currently being
made by anthropological geneticists. It is believed that mtDNA and Y
chromosome data
offer an opportunity for discourse among our often disparate fields, allowing
us to achieve a
greater understanding of American Indian cultural affiliation and demographic
history.
However, further studies should gain tribal approval before using tribal blood,
especially when
claims are subsequently made denying various affiliations between that tribe’s
blood and other
groups or ancient skeletons.
24
References
Cited
Bailliet, G.,
F. Rothhammer, F.R. Carnese, C.M. Bravi, and N.O. Bianchi. 1994. Founder
Mitochondrial
Haplotypes in Amerindian Populations. American Journal of Human Genetics,
54:27-33.
Barbujani,
Guido. 1997. DNA Variation and Language Affinities. American Journal of Human
Genetics,
61:1011-1014.
Bateman,
Richard, Ives Goddard, Richard O’Grady, V.A. Funk, Rich Mooi, W. John Kress, and
Peter Cannell.
1990. Speaking of Forked Tongues: The Feasibility of Reconciling Human
Phylogeny and
the History of Language. Current Anthropology, 31(1):1-13.
Bradman, Neil
and Mark Thomas. 1998. Why Y? The Y Chromosome in the Study of Human
Evolution,
Migration, and Prehistory. Science Spectra, 14:32-37.
Bianchi, N.O.,
G. Bailliet, C.M. Bravi. 1994. Peopling of the Americas as Inferred Through The
Analysis of
Mitochondrial DNA. Brazil Journal of Genetics, 18:661-668.
Bianchi,
Nestor, Graciela Bailliet, Claudio Bravi, Raul Carnese, Francisco Rothhammer,
Veronica
Martinez-Marignac, and Sergio Pena. 1997. Origin of Amerindian Y-Chromosomes as
Inferred by the
Analysis of Six Polymorphic Markers. American Journal of Physical
Anthropology,
102:79-89.
Blackman,
Margaret B. 1990. Haida: Traditional Culture. In Handbook of North American
Indians,
Vol. 7: Northwest Coast. Edited by Wayne Suttles. Washington D.C.:
Government
Printing
Office.
Bonatto, Sandro
L. and Francisco M. Salzano. 1997. Diversity and Age of the Four Major
mtDNA
Haplogroups, and Their Implications for the Peopling of the New World.
American
Journal of
Human Genetics, 61:1413-1423.
Boyd, Robert.
1990. Demographic History, 1774-1874. In Handbook of North American Indians,
Vol. 7:
Northwest Coast. Wayne Suttles ed. Washington D.C.: Government Printing
Office.
Boyd, Robert.
1999. The Coming of the Spirit of Pestilence: Introduced Infectious Diseases
and
Population
Decline among Northwest Coast Indians, 1774-1874. Seattle: University of
Washington
Press.
25
Brown, Michael
D., Seyed H. Hosseini, Antonio Torroni, Hans-Jurgen Bandelt, Jon C. Allen,
Theodore G.
Schurr, Rosaria Scozzari, Fulvio Cruciani, and Douglas C. Wallace. 1998. mtDNA
Haplogroup X:
An Ancient Link between Europe/Western Asia and North America? American
Journal of
Human Genetics, 63:1852-1861.
Cavalli-Sforza,
L.L., and W.F. Bodmer. 1971. The Genetics of Human Populations. San
Francisco: W.H.
Freeman and Company.
Cavalli-Sforza,
L. Luca, Paolo Menozzi, and Alberto Piazza. 1994. The History and Geography
of Human
Genetics. Princeton, New Jersey: Princeton University Press.
Chatters, James
C. 2000. The Recovery and First Analysis of an Early Holocene Human
Skeleton from
Kennewick, Washington. American Antiquity, 65(2):291-316.
Chen, Yu-Sheng,
Antonel Olckers, Theodore G. Schurr, Andreas M. Kogelnik, Kirsi Huoponen,
and Douglas C.
Wallace. 2000. mtDNA Variation in the South African Kung and Khwe- and
Their Genetic
Relationship to Other African Populations. American Journal of Human
Genetics,
66:1362-1383.
Cook,
Sherburne. 1978. Historical Demography. In Handbook of North American
Indians, Vol.
8:
California. Robert F. Heizer ed. Washington D.C.: Government Printing
Office.
D’Azevedo,
Warren L. 1986. Handbook of North American Indians, Vol. 11: Great Basin.
Washington
D.C.: Government Printing Office.
De Laguna,
Frederica. 1975. Matrilineal Kin Groups in Northwestern North America. In Volume
1 of
Proceedings: Northern Athapaskan Conference, 1971. A. McFadyen Clark, ed.
Canada.
National
Museum of Man. Mercury Series. Ethnology Service Papers 27. Ottawa.
Dillehay, Tom
D. 2000 The Settlement of the Americas: A New Prehistory. Basic Books: New
York.
Dobyns, Henry.
1983. Their Number Became Thinned: Native American Population Dynamics in
Eastern
North America. Knoxville: University of Tennessee Press.
Donnelly,
Peter, and Simon Tavare. 1995. Coalescents and Genealogical Structure Under
Neutrality.
Annual Review of Genetics, 29:401-421.
Easton, Ruth
D., D. Andrew Merriwether, Douglas E. Crews, and Robert E. Ferrell. 1996.
mtDNA Variation
in the Yanomami: Evidence for Additional New World Founding Lineages.
American
Journal of Human Genetics, 59:213-225.
Ehle, John.
1988. Trail of Tears: The Rise and Fall of the Cherokee Nation. New York:
Anchor
Books.
26
Flannery, Kent
V., and Joyce Marcus. 1994. Early Formative Pottery of the Valley of Oaxaca,
Mexico.
Memoirs of the Museum of Anthropology, University of Michigan, No. 27.
Forster, Peter,
Rosalind Harding, Antonio Torroni, and Hans-Jurgen Bandelt. 1996. Origin and
Evolution of
Native American mtDNA Variation: A Reappraisal. American Journal of Human
Genetics,
59:935-945.
Gibbons, A.
1998. Calibrating the molecular clock. Science, 279:28-29.
Goddard, Ives.
1978. A Further Note on Pidgin English. International Journal of American
Linguistics,
44(1):73.
Goddard, Ives.
1996. Introduction. In Handbook of North American Indians, Vol. 17:
Languages.
Ives Goddard
ed. Washington D.C.: Government Printing Office.
Greenberg,
Joseph H. 1987. Language in the Americas. Stanford University Press, California.
Greenberg,
J.H., C.G. Turner II, and S.L. Zegura. 1986. The Settlement of the Americas: A
Comparison of
the Linguistic, Dental, and Genetic Evidence. Current Anthropology,
27:477-497.
Hayden, Brian
editor. 1992. A Complex Culture of the British Columbia Plateau: Traditional
Stl’atl’imx
Resource Use. Vancouver: UBC Press.
Helm, June.
1981. Dogrib. In Handbook of North American Indians, Vol. 6: Subarctic. June
Helm editor,
Washington D.C.: Government Printing Office. Pp. 291-309.
Hoelzer, Guy
A., Joel Wallman, Don J. Melnick. 1998. The Effects of Social Structure,
Geographical
Structure, and Population Size on the Evolution of Mitochondrial DNA: II.
Molecular
Clocks and the Lineage Sorting Period. Journal of Molecular Evolution,
48:21-31.
Hudson, Richard
R. 1990. Gene Genealogies and the Coalescent Process. Oxford Surveys in
Evolutionary
Biology, 7:1-44.
Jones, Peter.
2002. Old World Infectious Diseases in the Plateau Area of North America during
the
Protohistoric: Rethinking Our Understanding of the “Contact” in the Plateau. In
circulation.
Kaestle,
Frederika. 1997. Molecular Analysis of Ancient Native American DNA From Western
Nevada.
Nevada Historical Society Quarterly, 40(1):85-96.
Kaestle,
Frederika, and David Glenn Smith. 2001. American Journal of Physical
Anthoropology,
115:1-12.
27
Karafet,
Tatiana, Stephen L. Zegura, Jennifer Vuturo-Brady, Olga Posukh, Ludmila Osipova,
Victor Wiebe,
Francine Romero, Jeffery C. Long, Shinji Harihara, Feng Jin, Bumbein
Dashnyam,
Tudevdagva Gerelsaikhan, Keiichi Omoto, and Michael F. Hammer. 1997. Y
Chromosome
Markers and Trans-Bering Strait Dispersals. American Journal of Physical
Anthropology,
102:301-314.
Karafet, T.M.,
S.L. Zegura, O. Posukh, L. Osipova, A. Bergen, J. Long, D. Goldman, W. Klitz,
S. Harihara, P.
de Knijff, V. Wiebe, R.C. Griffiths, A.R. Templeton, and M.F. Hammer. 1999.
Ancestral Asian
Source(s) of New World Y-Chromosome Founder Haplotypes. American
Journal of
Human Genetics, 64:817-831.
Lekson, Stephen
H. 2000. The Chaco Meridian: Centers of Political Power in the Ancient
Southwest.
Walnut Creek, CA: AltaMira Press.
Lewontin, R.C.
1971. The Apportionment of Human Diversity. In Evolutionary Biology, Vol.
6.
T.H. Dobzhansky
et al., eds. Pp. 381-398. New York: Appleton-Century-Crofts.
Lorenz, Joseph
G., and David Glenn Smith. 1996. Distribution of Four Founding mtDNA
Haplogroups
Among Native North Americans. American Journal of Physical Anthropology,
101:307-323.
MacEachern,
Scott. 2000. Genes, Tribes, and African History. Current Anthropology,
41(3)357-
384.
Matson, G.A.
1938. Blood groups and ageusia in Indians of Montana and Alberta. American
Journal of
Physical Anthropology, 24:81-89.
Matson, G.A.,
and H.F. Schrader. 1933. Blood grouping among the “Blackfeet” and “Blood”
tribes of
American Indians. Journal of Immunology 25:15-163.
Merriwether, D.
Andrew, Francisco Rothhammer, and Robert E. Ferrell. 1995. Distribution of
the Four
Founding Lineage Haplotypes in Native Americans Suggests a Single Wave of
Migration for
the New World. American Journal of Physical Anthropology, 98:411-430.
Mountain,
Joanna L., and L. Luca Cavalli-Sforza. 1997. Multilocus Genotypes, a Tree of
Individuals,
and Human Evolutionary History. American Journal of Human Genetics,
61:705-
718.
Napton, L.
Kyle. 1997. The Spirit Cave Mummy: Coprolite Investigations. Nevada
Historical
Society
Quarterly, 40(1):97-103.
O’Grady, R.T.,
I. Goddard, R.M. Bateman, W.A. Di Michele, V.A. Funk, W.J. Kress, R. Mooi,
and P.F.
Cannell. 1989. Genes and Tongues. Science, 243:1651.
28
O’Rourke, D.H.,
M.G. Hayes, and S.W. Carlyle. 2000. Spatial and Temporal Stability of mtDNA
Haplogroup
Frequencies in Native North America. In Human Biology, 72(1):15-34.
Opler, Morris
E. 1983. The Apachean Culture Pattern and Its Origins. In Handbook of North
American
Indians, Vol. 10: The Southwest. Edited by Alfonso Ortiz. Washington D.C.:
Government
Printing Office.
Page, Roderic
D.M., and Michael A. Charleston. 1990. Reconciled Trees and Incongruent Gene
and Species
Trees. DIMACS Series in Discrete Mathematics and Theoretical Computer
Science,
Vol. 00: 1-14.
Pamilo, P., M.
Nei. 1998. Relationships Between Gene Trees and Species Trees. Molecular
Biological
Evolution, 5(5):568-583.
Parsons, T., D.
Muniec, and K. Sullivan. 1997. A high observed substitution rate in the human
mitochondrial
control region. Nature Genetics, 15:363-68.
Poloni, E.S.,
O. Semino, G. Passarino, A.S. Santachiara-Benerecetti, I. Dupanloup, A.
Langaney,
and L.
Excoffier. 1997. Human Genetic Affinities for Y-Chromosome P49a,f/TaqI
Haplotypes
Show Strong
Correspondence with Linguistics. American Journal of Human Genetics,
61:1015-
1035.
Pluciennik, M.
1995. A perilous but necessary search: Archaeology and European Identities.
In
Nationalism
and archaeology: Scottish Archaeological Forum. Edited by J. Atkinson and J.
O’Sullivan, pp.
35-58. Glasgow: Cruithne Press.
Quimby, George
I. 1985. Japanese Wrecks, Iron Tools, and Prehistoric Indians of the Northwest
Coast.
Arctic Anthropology, 22(2):7-15.
Rogers, A., and
L. Jorde. 1995. Genetic Evidence on Modern Human Origins. Human Biology,
67:1-36.
Ruhlen, M.
1986. A Guide to the World’s Languages. Vol 1. Stanford: Stanford
University Press.
Ruhlen, M.
1994. Linguistic Evidence for the Peopling of the Americas. In Methods and
Theory
for
Investigating the Peopling of the Americas, Bonnichsen and Steele editors.
Center for the
Study of the
First Americans, Corvallis: Oregon. Pp. 177-188.
Santos,
Fabricio R., Arpita Pandya, Chris Tyler-Smith, Sergio D.J. Pena, Moses
Schanfield,
William R.
Leonard, Ludmila Osipova, Michael H. Crawford, and R. John Mitchell. 1999. The
Central
Siberian Origin for Native American Y Chromosomes. American Journal of Human
Genetics,
64:619-628.
29
Schurr,
Theodore G., Scott W. Ballinger, Yik-Yuen Gan, Judith A. Hodge, D. Andrew
Merriwether,
Dale N. Lawrence, William C. Knowler, Kenneth M. Weiss, and Douglas C.
Wallace. 1990.
Amerindian Mitochondrial DNAs Have Rare Asian Mutations at High
Frequencies,
Suggesting They Derived from Four Primary Maternal Lineages. American Journal
of Human
Genetics, 46:613-623.
Schurr,
Theodore G., Rem I. Sukernik, Yelena B. Starikovskaya, and Douglas C. Wallace.
1999.
Mitochondrial
DNA Variation in Koryaks and Itel’men: Population Replacement in the Okhotsk
Sea- Bearing
Sea Region During the Neolithic. American Journal of Physical
Anthropology,
108:1-39.
Schuster,
Helen. 1998. Yakima and Neighboring Groups. In Handbook of North American
Indians,
Vol. 12: The Plateau. Deward E. Walker, Jr., ed. Washington D.C.: Government
Printing
Office.
Scozzari,
Rosaria, Fulvio Cruciani, Piero Santolamazza, Patrizia Malaspina, Antonio
Torroni,
Daniele
Sellitto, Barbara Arredi, Giovanni Destro-Bisol, Gianfranco De Stefano, Olga
Rickards,
Cristina
Martinez-Labarga, David Modiano, Gianfranco Biondi, Pedro Moral, Antonel
Olckers,
Douglas C.
Wallace, and Andrea Novelletto. 1999. Combined Use of Biallelic and
Microsatellite
Y-Chromosome
Polymorphisms to Infer Affinities among African Populations. American
Journal of
Human Genetics, 65:829-846.
Smith, David
Glenn, Ripan S. Malhi, Jason Eshleman, Joseph G. Lorenz, and Frederika A.
Kaestle. 1999.
Distribution of mtDNA Haplogroup X Among Native North Americans.
American
Journal of Physical Anthropology, 110:271-284.
Stern,
Theodore. 1998. Cayuse, Umatilla, and Walla Walla. In Handbook of North
American
Indians,
Vol. 12: The Plateau. Deward E. Walker, Jr., ed. Washington D.C.: Government
Printing
Office.
Templeton, Alan
R. 1993. The “Eve” Hypotheses: A Genetic Critique and Reanalysis. American
Anthropologist,
95(1):51-72.
Torroni, A.,
T.G. Schurr, C-C Yang, E.J.E. Szathmary, R.C. Williams, M.S. Shanfield, G.A.
Troup, et al.
1990. Native American Mitochondrial DNA Analysis Indicates that the Amerind
and Nadene
Populations Were Founded by Two Independent Migrations. Genetics,
130:153-162.
Torroni, A.,
T.G. Schurr, M.F. Cabell, M.D. Brown, J.V. Neel, M. Larson, D.G. Smith, C.M.
Vullo, and D.C.
Wallace. 1993. Asian Affinities and Continental Radiation of the Four Founding
Native American
mtDNAs. American Journal of Human Genetics, 53:563-590.
VanStone, James
W. 1984. Exploration and Contact History of Western Alaska. In Handbook of
North
American Indians, Vol. 5: The Arctic. David Damas, ed. Washington D.C.:
Government
Printing
Office.
30
Walker, Deward
E., Jr. 1972. The Emergent Native Americans. Boston: Little, Brown and
Company.
Walker, Deward
E., Jr. 1990. A Study of CRUIR Enrollment and Population. General Council
and Board of
Trustees of the Confederated Tribes of the Umatilla Indian Reservation. Mission:
Oregon.
Walker, Deward
E., Jr. 1998. Introduction. In Handbook of North American Indians, Vol. 12:
The
Plateau. Deward E. Walker, Jr., ed. Washington D.C.: Government Printing
Office.
Ward, R.H.,
Alan Redd, Diana Valencia, Barbara Frazier, and Svante Paabo. 1993. Genetic and
Linguistic
Differentiation in the Americas. Proceedings of the National Acadamey of
Sciences,
USA,
90:10663-10667.
Watson,
Elizabeth, Peter Forster, Martin Richards, and Hans-Jurgen Bandelt. 1997.
Mitochondrial
Footprints of Human Expansions in Africa. American Journal of Human
Genetics,
61:691-704.
Weiss, K.M.
1994. American Origins. Procession of the National Academy of Science,
91:833-
835.
Wells, Spencer.
2000. Cavalli-Sforza Laboratory, Stanford University Web Page.
Http://163.1.230.14/eurasia/htdocs/anthgen.html Accessed July 17, 2000.
Wolpoff,
Milford. 1999. Paleoanthropology, 2
nd
Ed. New
York: McGraw-Hill.
Boulder, Co.
80306
|
