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Department of Biological Anthropology and Anatomy

Duke University
Durham, NC 27710

The Status of the Race Concept in Physical Anthropology

There are hereditary differences among human beings. Some of these differences have geographical correlates. Some genetic variants that produce physical or behavioral deficits occur significantly more often in some areas, or in some ethnic
groups, than in others. However, none of these facts provides any intellectual support for the race concept, for racial classifications, or for social hierarchies based on ethnic-group membership.
The geographical element of the race concept is important in theory but is widely ignored in practice since it does not
conform well to the facts of current human phenotype distribution. Much of the literature on supposed racial differences
involves such geographically meaningless exercises as studying differences among "races" by subdividing a sample of
North Americans. If races are defined as geographically delimited conspecific populations characterized by distinctive
regional phenotypes, then human races do not exist now and have not existed for centuries, [race, human variation, intelligence]

he concept of race is a divisive and emotionally

charged topic among physical anthropologists.
The history of the American Association of Physical Anthropologists' "Statement on Biological Aspects of
Race" (AAPA 1996) attests to our divisions on this issue.
The AAPA statement had its inception at the 1989 meetings of the American Association for the Advancement of
Science, where the Canadian psychologist J. Philippe
Rushton was invited to deliver a talk on his notorious racial theories (Rushton and Bogaert 1989). Some physical
anthropologists who happened to be present were appalled to hear Rushton's views propounded under the auspices of the AAAS. They felt that physical anthropologists, as the supposed scientific experts on matters of race,
ought to have been consulted before Rushton was given a
platform. Convinced by this incident that it was high time
for physical anthropologists to take an official stand
against scientific racism, they asked the Executive Committee of the American Association of Physical Anthropologists to establish a committee to work toward that
A working group set up under the direction of Sol Katz.
the AAPA's representative to the AAAS, submitted a
draft statement on race in 1992 to the AAPA Executive
Committee (Sirianni 1992). The Executive Committee revised it further and passed it along for approval to the Association's business meeting in Toronto in 1993. The
statement was rejected by a vote of 43 to 35, with 4 abstentions (Sirianni 1993). Nevertheless, the AAPA executive
was given permission to rewrite the statement and dis-

seminate the revision in the name of the Association. After

three years of additional discussion, debate, and revision,
the statement was finally approved by the Executive Committee and published in the December 1996 issue of the
American Journal of Physical Anthropology (AJPA).
Some AAPA members who spoke against the race
statement in Toronto were opposed to it on philosophical
grounds. The AAPA, they argued, simply had no business
making pronouncements of this sort. If the issues being
dealt with were matters of scientific fact, they should be
thrashed out in the scientific literature, not settled by passing resolutions at meetings. It was no more appropriate for
the AAPA to have an official position on the facts of human racial variation, these people insisted, than it would
be for it to have an official position on the phylogeny of
marmosets, or the diagnostic skeletal signs of syphilis, or
any other factual issue. And if the issues in question were
matters not of fact but of politics and morality, then physical anthropologists could say nothing more authoritative
about them than anyone else.
Although these objections to the "Statement on Biological Aspects of Race" had nothing to do with the issue
of race as such, the rejection of that statement at the 1993
AAPA business meeting also reflected substantive disagreements about race among biological anthropologists.1
Some of us, myself included, regard human races as oversimplified or nonsensical constructs (Brace 1964, 1996;
Goodman and Armelagos 1996: Harrison et al. 1977;
Keita and Kiftles 1997; Livingstone 1962: Marks 1995:
Molnar 1992; Montagu 1942a, 1942b). Others think that

American Anthropologist 100(3)651 -660. Copyright 1999. American Anthropological Association



VOL. 100, No. 3

races are real biological entities. In a 1985 survey, 365

physical anthropologists were asked whether they agreed
with the statement, "There are biological races within the
species Homo sapiens." Almost half of them {N = 181)
said they did. Almost as many (N = 148) said they did not
(Lieberman and Reynolds 1996).
A similar 1978 survey, which revealed a similar difference of opinion, showed that the positions that physical
anthropologists take on these issues tend to correlate with
their social status and cultural background (Lieberman
and Reynolds 1978). Scientists' attitudes toward the race
concept are probably also correlated to some degree with
their politics. Experience suggests that (as might be expected) physical anthropologists who reject the concept of
race tend to lean more to the political left than their opponents do. But the division between the two camps is not really a split between tender-minded liberal egalitarians and
tough-minded conservative elitists. To a surprising extent, physical anthropologists in both camps make similar
assertions, cite similar sources, and express similar fervent opposition to racist practices and beliefs. The difference between them is mainly one of emphasis. The findings that one group admits grudgingly and seeks out
reasons for disregarding are spotlighted by the other group
as the central facts that reveal the way things really are.
There is a case to be made for each side, and it is not hard to
find physical anthropologists who have questioned the
existence of races in one publication but have used racial
categories to structure their data in another.

The Case for the Race Concept

Those who defend racial taxonomies generally say they
are just one way of expressing the generally recognized
fact that human genetic variation is correlated with geography. For example, most of the world's people who have
very dark skin and woolly-textured, tightly curled hair live
in Africa south of the Tropic of Cancer. Although many
people who live elsewhere also meet this description, the
great majority of them are descended from people who
lived in sub-Saharan Africa. In some parts of the world,
immigrants from tropical Africa have simply disappeared
into the general population through interbreeding, but in
other areasfor instance, in North Americathey have
formed persistent ethnic groups with distinctive cultural
traditions and a tendency toward preferential mating
within the group. Within such ethnic groups, assortative
mating has maintained high frequencies of a recognizably
"African" facial appearance and of genetic variants that
occur with high frequencies in equatorial African populations (for example, the hemoglobin-S mutation associated
with sickle-cell disease). Defenders of racial taxonomy
argue that it is not unreasonable to think of such groups as
African-derived breeding populations, or to distinguish
them with labels like "African American" that reflect their



African ancestry, or to lump them together for some purposes with their parent populations in Africa as constituting a "Negroid" group.
Proponents of the race concept acknowledge that racial
classifications can be used to discriminate against people.
But because such classifications reflect certain facts of human biology, they can also be used justly and fairly to
serve benign ends. For example, doctors need to be alerted
to the elevated probability of sickle-cell disease in patients
of equatorial African ancestry. Forensic anthropologists
may be asked by the police to provide racial identifications to help in solving crimessay, to determine whether
a skeleton found in the woods could be that of an African
American murder victim. Because there are some skeletal
traits that occur more frequently among some North
American ethnic groups than among others, it is sometimes possible to answer such questions with a fair degree
of confidence. And because racially defined ethnic groupings are real and important elements in American culture,
we often need to recognize such groupings in investigating the interaction between culture and biology. For instance, if we wish to determine whether Black children
have been systematically exposed to higher environmental lead levels than Whites, we need to structure our
sample in terms of race (Schell 1997).

The Case against the Race Concept

Biological anthropologists who deny the value of racial
typology would grant all these points, but would insist that
racial categories are nevertheless biologically incoherent
and heuristically misleading. As one classic textbook of
human biology expressed it two decades ago, "Classifications of man into Mongoloids, Caucasoids, Negroids, etc.
undoubtedly express certain genuine features of human
variation but they do so in a crude and misleading way"
(Harrison etal. 1977:184).
Proponents of the race concept usually define races in
terms of the typical or average properties of regional human populations, as though racial categories were geographically delimited biological subspecies. Summarizing the definitions of "race" proposed by proponents of
human racial classifications over the past half-century,
Molnar (1992:23) notes that all such definitions stress the
concept of races as geographical entities: "Primarily, the
divisions are based on the sharing of a common territory or
space and [assume] that geography played some role in establishing boundaries until recent times."
But this is not how the concept of race is in fact employed in either common usage or the scientific literature.
Geography has little to do with the race concept in its actual application. Studies of "racial" differences often draw
their data from so-called Black, White, and Asian individuals bom in the same geographical region. For example, much of the published literature on supposed racial


differences in "intelligence" is based on data sampled

from native-born North Americans identified as representatives of different races.
If North American "Blacks," "Whites," 'Asian-Americans," "Amerindians," and so on are racially different,
then (since all these people inhabit the same geographical
region) races are not geographically distinct. And if these
people are not racially different, then (since the range of
their combined phenotypes encompasses roughly the
whole range of variation in the human species) races are
notphenotypically distinctive. Thereforeunless we decide to leave the modem populations of at least Australia,
North and South Africa, Oceania, large parts of Eurasia,
and the entire Western Hemisphere out of the human picturedefinitions of races as geographically delimited
populations marked by distinctive phenotypes cannot correspond to any current reality.
When advocates of racial taxonomy try to take these
facts into account, the results are predictably incoherent.
For example, in their classic textbook on race, Coon et al.
(1950) tried to deal with the modern populations of North
America by recognizing a "primary human race" (!) called
"North American Colored," which "includes all forms
from Forest Negro to the borderline of White, with all possible combinations. .. .American Indian is a third genetic
element in the mixture.... For purposes of racial taxonomy it might more reasonably be subdivided into Negro
and Mulatto, with the least Negroid extreme included with
the Northwest European Whites, into which body many of
its numbers have already been assimilated" (p. 131).
Obviously, a classificatory unit defined in these
strained and nebulous terms is not delimited by either
geography or genetics, and has no taxonomic utility or
biological meaning. Indeed, some proponents of racial
classification grant that such "racial" groups as "North
American Colored" are biologically meaningless. "Most
human populations today," writes Shipman (1994:B-1),
"are the result of a delightful and thorough admixture of
genes from many different groups. Even those with strong
ethnic identities are often a genetic mixture, including the
erroneously labeled race' of African Americans.... African Americans are not a race." But the same reasoning
applies to other ethnic groups of North America, including
"Whites," "Asian Americans," and "Native Americans."
It applies as well to similar groups in other regions that
have experienced large-scale immigration from outside
over the course of the past 300 years. When all these parts
of the world are omitted from our racial classifications,
there is not much left to classify.
More commonly, advocates of racial typologies try to
leave most of the inconvenient facts about modern human
populations out of the picture. Such proponents grant that
races are less distinct than they used to be. but insist that
"geography played some role in establishing boundaries
until recent times." The idea here is that racial phenotypes



were pretty well correlated with geography (with Negroids restricted to Africa, Caucasoids to western Eurasia,
Mongoloids to eastern Asia, and so on) until the era of
European colonialism, when massive population movements both voluntary (like the colonization of South Africa by Dutch settlers) and forced (like the initial colonization of the Americas by enslaved Africans, or of Australia
by deported English convicts) brought different races together in various parts of the world and produced racially
mixed populations that are not easy to classify. This artificially induced and unnatural commingling of different
races has muddied the original picture, but enough of the
human species remains in a relatively pristine condition to
enable us to reconstruct the original situation by studying
"primitive isolates" today in uncolonized parts of the
world like Amazonia, the Ituri rain forest, and Lapland. Or
so the story goes.
It is true that human populations in some parts of the
world were more uniform and distinctive a thousand years
ago than they are at present. But populations like those of
modem North America, with high levels of phenotypic
variability maintained partly by migration and gene flow
from elsewhere, are not a new phenomenon. Similar
populations have inhabited northern and southern Africa
and much of western, central, and southern Asia for centuries or millennia. It would have been just as futile an exercise to try to apply racial typologies to the highly variable
people of Egypt or India four thousand years ago as it is to
do so in the United States today. In such populations, "racial" types are polymorphic, like ABO blood-group phenotypes. It makes no more sense to classify the individuals
comprising these populations into racial categories based
on epidermal pigmentation, hair texture, or nose, lip, and
eyelid shape (the traits that loom largest in our racial typologies, probably because they are all visible in people's
faces) than it would to separate them into races on the basis
of their ABO phenotypes. In fact, it makes even less sense,
since ABO phenotypes are discrete, whereas "racial"
types in such populations are highly variable and intergrade imperceptibly with each other.
There are of course things to be learned about human
adaptations by trying to reconstruct the past distributions
of human phenotypes. Skin pigmentation furnishes a familiar example. The darkness of human skin in the Old
World appears to be inversely correlated with distance
from the equator. Populations that deviate from this general pattern can plausibly be interpreted as recent migrants
from higher to lower latitudes, or vice versa (Brace 1996).
The pattern probably reflects a long history of low-level
natural selection favoring dark skin in areas of high yearround insolation. We might not discern this pattern so easily if we used modern data uncritically and pretended not
to know that the presence of large numbers of pinkskinned people in the Transvaal and of black-skinned people in Canada is a relatively recent phenomenon.



VOL. 100, No. 3

But there are also things we can learn about human adaptations by looking at modem phenotype distributions.
European colonies were established throughout most of
the world during the eighteenth and nineteenth centuries.
In temperate-zone areasAustralia, New Zealand, North
America, southern South America, and South Africa
these colonies tended to expand and supplant or supplement the indigenous peoples; in tropical areas, they generally made less of an impact. Crosby (1986) suggests that
this differential success reflects European cultural adaptation to temperate-zone ecologies. However, it might also
reflect patterns of selection against European phenotypes
in equatorial climates. Perhaps pink-skinned variants of
Homo sapiens do not thrive in areas of high year-round insolation. To test this thesis, we need to look at recentpopulations, not at our reconstructions of populations that lived
a thousand years ago. Both past and present distributions
provide biological data on human adaptations and microevolution. It is a mistake to think that today' s human populations are somehow unreal, unnatural, or corrupted, or
that modem technology has freed them from selection
No matter whether we look at past or at present-day
populations, the use of racial categories in structuring our
samples hinders, not helps, our efforts to describe human
variation and explain its causes. If we want to frame or test
hypotheses about the adaptive significance of skin color,
the appropriate question to ask is not, "Do Negroids do
better than Caucasoids in some environments?" but
rather, "Do dark-skinned people do better than lightskinned people in some environments?" Since there is
considerable variation within, and overlap between, "Negroids" and "Caucasoids" with respect to skin color, analyzing the variation in terms of racial categories serves
only to blur the question and introduce irrelevant variables
into the data.
Similar criticisms apply to any attempt to use racial
categories in describing or analyzing human genetic variation. Since there are thousands of separate, independently assorting variable loci in the human genome, it
is highly unlikely a priori that variation at any particular
locus will covary with any other. We would therefore expect on theoretical grounds that a descriptive classification based on a small number of "racial" traits would be of
little use in summarizing the variation that occurs at any or
all of the other variable loci in the human genome. Empirical studies bear out this expectation. Even if we try to
backdate the evidence to A.D. 1500 by restricting our data
base to supposedly isolated aboriginal populations, the
geographic patterns of variation at most loci do not fit our
racial typologies. In the case of the ABO system, for example, the A allele reaches its highest frequencies in
southern Australia, Europe, northeastern Asia, and the
Arctic fringes of North America; the B allele in central
Asia and West Africa; and the O allele in the New World,



New Guinea, and northern Australia (Cavalli-Sforza

1996:171-173). Other genetic loci generally show patterns that are discordant with racial typologies, ABO allele distributions, and/or each other. These facts underlie
Lewontin's often-quoted (1972) conclusion that "only
6.3% of human diversity [is] attributable to race," and that
there is more genetic variation within "racial" groupings
than there is between them (cf. Templeton, this issue).

Are There Trends in the Use

of the Race Concept?
The debate between biological anthropologists who defend the concept of race and those who deplore it has been
going on for over half a century (Barkan 1992). It is not
clear whether this debate is moving toward a resolution.
Different observers have different perceptions of the
changing importance of the race concept. Some think that
the use of racial categories and concepts is a vanishing
relic of outmoded and discredited ways of thought in biological anthropology (Landau 1997; Sanjek 1994),
whereas others discern a recent resurgence of the race concept in skeletal biology, forensic anthropology, paleoanthropology, nutritional studies, and human genetics
(Goodman and Armelagos 1996; Lieberman and Jackson
As far as I know, the only empirical study bearing on
these claims was undertaken by Littlefield et al. (1982),
who concluded that the 1970s had witnessed a sudden and
widespread abandonment of racial classification in textbook presentations of human biology. To try to assess
similar trends in the primary scientific literature, I undertook a survey of the research articles published in the
AAPA's official journal, the American Journal of Physical Anthropology, from 1965 to 1996.1 surveyed the oddnumbered years plus the 12 issues from 1996 (the most recent complete annual file at the time of this writing). Of the
1,749 scientific articles contained in these 17 annual files,
810 (46%) dealt with some aspect of modem human variation and therefore might potentially have used racial
categories. I divided these 810 articles into those that utilized racial categories, either in structuring the sample or
analyzing the data, and those that did not. Papers were
scored as not involving racial categories if their human
subjects were grouped into classes defined strictly in
terms of geography ("Lithuanians"), genotype ("hemoglobin-S homozygotes"), phenotype ("obese people"),
ethnic self-identification ("Amish"), or any combination
of such criteria. "Racial categories," as defined here, include traditional racial taxa ("Australoids"), self-contradictory geographical descriptions ("Australian Europeans"), ethnic identifications inferred by the researcher by
just looking at people, and any groupings defined in terms
of supposed historical origins rather than observable characteristics. The following are examples of racial categories





140 -

120 -


Total humarwanaSon
C3--Tolal AJPA articles per year

E S 2 ! S E : 2 ~ I D " '
2 2 2 2 S & S S S S S S S S S S S
Figure 1. Annual publication rates in the American Journal ofPhysical Anthropology, 1965-1996. White squares signify total scientific
articles, and black diamonds, total articles on modem human variation.

found in these 810 papers: "Typical Mongoloid and Negroid collectives," "American Japanese males," "(American) subjects primarily of African origin," "Full-blooded
Papago Indians," and "Australians of pure Aboriginal ancestry." Some papers could have been scored either way,
and a different observer would no doubt come up with
somewhat different definitions and numbers, but the standards used here were at any rate applied fairly consistently.
In the sampled years, the number of articles about modern human variation published annually in the AJPA rose
almost every year from 1965 to 1983. Since then, it has declined steadily. This rise and fall partly reflects a parallel
but more erratic fluctuation in the number of articles on all
subjects published annually in the AJPA (Figure 1). But
even when we correct for this by expressing the number of
articles on human variation as a percentage of total AJPA
articles published in each sampled year, there still appears
to have been a long-term decline in the percentage of the
AJPA devoted to the topic of modern human variation
(Figure 2). This might be interpreted as betokening a decreasing interest in the whole subject of "race" broadly defined. However, it might also reflect the proliferation of
new journals that provide alternative publication venues
for articles on human biology and genetics.
Of the 810 articles on modern human variation published in the AJPA during the years sampled, 40.5% utilized racial categories. The annual tally of racial-category
articles as a percentage of all human-variation articles
fluctuates widely around a near-identical mean of

Figure 2. Papers on variation in modem human populations, as a

percentage of total AJPA articles, 1965-1996. The least-squares line
trends downward, although the correlation is not significant (Fr =

40.67%, with a standard deviation of 11.45%. There is no

discernible trend in these percentages (Figure 3).
In summary, the role played by racial taxonomy in the
study of modern human variation has apparently changed
little or not at all over the course of the past 30 years. In the
1990s, as in the 1960s, most researchers studying human
variation do not make use of the concept of race in gathering and analyzing their data; however, a consistently large
minority continue to do so. These figures suggest that







Figure 3. Use of racial classifications in analyzing or structuring data

in AJPA articles on modem human variation, 1965-1996. No trend is
discernible over this 32-year period.



VOL. 100, No. 3

neither the proponents nor the opponents of racial classification have any grounds for thinking that history is on
their side.

Biological Determinism, Biological

Superiority, and Race
Although physical anthropologists are divided over the
concept of race, they share a general conviction that human behavior is significantly channeled, constrained, and
determined in various ways by human biology. This conviction is not universally shared by anthropologists in
other subfields. I suspect that many social and cultural anthropologists would dismiss this idea as politically motivated "biological determinism." Some seem to regard any
claims abouthuman biology as racist or elitist, and to think
of eugenics and The Bell Curve (Herrnstein and Murray
1994) whenever they hear people talk abouthuman behavior in the same breath as biology or genetics.
There are some good historical reasons for these associations. Throughout Western history, the wealthy and
powerful have found it comfortable and expedient to overestimate the importance of heredity in explaining the differences between people, in order to try to reassure themselves and persuade others that the prevailing social
inequalities are just and natural. In most or all complex societies, the ascription of social status has been to some degree hereditary, and membership in low-status classes or
castes has been widely regarded as a matter of simple inheritance, as though poverty, ignorance, and powerlessness were dominant alleles at single genetic loci. Such
practices and assumptions have never been more broadly
applied, more widely accepted, more strongly upheld by
mistaken scientific expertise, and more productive of misery, injustice, and evil than in the case of the concept of
biological race. Physical anthropology has in the past provided more intellectual support for this concept and its attendant evils than many of its present practitioners realize
or care to remember (Blakey 1994, 1996; Marks 1995;
Montagu 1942b; Wolpoff and Caspari 1997). Much of the
animus that attaches to the concept of race among physical
anthropologists stems from its shameful history.
Still, I think that the current widespread suspicions concerning biological concepts in anthropology involve some
misapprehensions. In what follows, I hope to persuade
some skeptical colleagues in other subfields of anthropology that admitting the reality and significance of hereditary biological differences between individuals does not
compel us to think that races are real biological entities, or
to believe that the rich are wealthy because of their superior genes. If we can arrive at a consensus on these points,
perhaps we can agree at a minimum that "biological anthropology" is not a contradiction in terms.
People are animals. As such, we face the same fundamental biological constraints on our lives as other ani-



mals. To live, we need to breathe, assimilate food, and excrete wastes. If we stop doing any of these things, we die.
Eventually we die anyway, no matter what we do. Like
other sorts of animals, we also face particular, speciesspecific biological constraints. Salmon can breathe water
but cannot learn to play the piano. The reverse is true of
most human beings. Cultural innovations may someday
enable us to evade our present biological constraints, but
at the moment we are stuck with them.
People also face environmental constraints on their
lives. Although many of these are beyond human control,
a lot of them are imposed on us by other people. Mostof us
could bericher,wiser, kinder, more accomplished, healthier, and happier than we are if only we had spent our lives
in different environments. Almost every sort of human potential is limited by both environmental and biological
factors. I cannot learn everything there is to leam because
my brain and my lifespan are finite. This is a fact of human
biology, which would be true in any environment currently attainable. But by the time I die, I will have learned
even less than I might have given my brain and lifespan,
because of my choices and because of the constraints
placed on me by my environment. This is true for all people, no matter what their hereditary capacities are.
I hope that we can all agree that these are simple, obvious truths about the human condition. (They are also truths
about the salmon condition, the horse condition, and so
on.) It follows that it makes no sense to ask whether a particular capacity is in principle limited chiefly by heredity
or by environment. Everything is in principle 100% limited by both heredity and environment. The life of a concert pianist must begin with a fertilized human ovum; an
opossum ovum will not suffice, because of its biological
limitations. On the other hand, no matter what sort of a human ovum we start with, it cannot develop into a concert
pianist in most environmentssay, in Europe in 10,000
B.C., or in the womb of an opossum, or in a 10% solution of
formaldehyde. Asking whether piano-playing skill is primarily determined by heredity or by environment is therefore meaningless. The relative importance of heredity and
environment in producing the observed differences between people in this or any other trait depends on the relative variabilities of the two factors in any particular situation. If all people were raised in identical environments,
any differences among them not due to their own choices
would obviously be due to heredity. If they were genetically identical, all such differences would be due to environmental factors.
In the world as it is today, it seems clear that some of the
differences between people (say, the differences between
an infant with Tay-Sachs disease and its parents) are almost entirely determined by genetic factors. Others (say,
the differences between political liberals and conservatives) are as far as we know determined almost entirely by
environment and individual choices. Yet others (e.g.,


piano-playing ability) are probably determined by combinations of, or interactions among environment, heredity,
and choice. Although scientists often try to estimate the
relative contributions of these factors to the observed variation in various human traits, we need to remember that
such estimates are themselves dependent on environmental variables. Some people believe that they can
evaluate the relative contributions of environmental and
genetic factors to a trait by comparing that trait's variation
among identical twins (who are clones of each other) with
its variation among fraternal twins (who are genetically
different). But even this approach does not really allow us
to factor out environmental influences altogether, because
the environment determines the extent to which a given
trait is influencedby geneticfactors. For example, in acultural context where (say) redheaded people were stereotyped as stupid and ineducable and were accordingly neglected by their teachers, we would expect identical twins
to resemble each other more closely in their educational
attainments than fraternal twins, simply because identical
twins are more likely to have the same hair color than fraternal twins are. In such an environment, success in school
might be causally linked with genetic factors that would
not affect educational attainment in other environments.
Because the degree to which any trait is genetically conditioned depends on environmental circumstances, there
is no such thing as "heritability" in the abstract. To quote
Weizmann et al. (1996:192-193), "Heritabilities depend
on the specific genetic composition of the population and
the environmental circumstances experienced by that
population.... [They] cannot be generalized to other
populations or other environmental conditions."
What is true of heritability is also true of fitness. The
theory of natural selection entails that within any species,
some genetic variants are more fit than othersthat is,
there are nonrandom factors that make certain variants
more likely than others to leave copies in the gene pools of
succeeding generations. Not all evolutionary change is
driven by natural selection, and it is not always easy to distinguish variants favored by selection from those that
prosper due to mere coincidence. For example, average
human skin pigmentation may well have decreased from
the seventeenth through the nineteenth centuries A.D. as a
side effect of the great expansions and emigrations of
light-skinned European populations during the era of colonialism. At the moment, human pigmentation may be on
the upswing again due to higher population growth rates in
the tropical countries of Africa, Asia, and the Americas,
where average skin color is darker than it is in Europe.
There is no reason to suspect that these historical fluctuations reflect changing patterns of natural selection on human skin color.
Genetic variants favored by natural selection can be
properly described as biologically superior to others. But
such variants are superior only in relation to a specific en-



vironmental context, including the species itself, its population structure, and its relationship to and interactions
with all aspects of its circumstances. Again, there is no
such thing as generalized fitness in the abstract. For example, lizards bom without limbs are generally at a disadvantage, but there have been situations in the past where this
was not the casewhich is why there are snakes and limbless lizards in the world today. Likewise, people who are
born without limbs are generally at a disadvantage; but we
can imagine or create environments where they are just as
fit as anyone else, or even more so.
Our culture leads us to regard mental abilities as the
most important markers of human status. Bothour cultural
traditions and our own professions as scholars and teachers encourage us to lump all mental abilities together as a
single variable called "intelligence," to equate high "intelligence" with biological superiority, and to feel that people with exceptional mental abilities somehow deserve to
be at the top of the heap. When a man with a crippled body
becomes a great astrophysicist, we are awed and inspired
by his example. When an illiterate, inarticulate, and unreflective man becomes a great boxer, we are less impressed. If the stupid prizefighter makes ten times as much
money in the course of a year as the crippled astrophysicist, we regard it as a scandal. Because intellectuals tend to
value other skillful manipulators of symbols more highly
than they value skillful boxers, gardeners, hunters, or masons, most of the published debate concerning the supposed biological superiority of certain human populations
has centered around the issue of congenital average differences in "intelligence" between "races." While nobody
gets very excited if scientists suggest that Swedes are on
the average taller than Japanese for genetic reasons, everybody gets hot under the collar whenever someone
claims that Swedes are on the average smarter than Nigerians for genetic reasons. The difference between the two
responses is due in part to the fact that "intelligence" is a
notoriously dubious variable, which is far less clearly defined and less easily quantified than "height." But at a
deeper level, the difference reflects the different cultural
values that we attach to stature and IQ.
However we choose to define or subdivide "intelligence," it is an unpleasant fact that some genetic variants
make their possessors stupider than other people: that is,
they result in impaired mental abilities in all currently attainable human environments. Some of these genes are
known to be significantly more common in some human
populations and ethnic groups than in others. These two
facts suggest (but do not prove) that human populations
and ethnic groups may well differ congenitally in average
mental potential at birth. This conclusion sounds shocking. However, even if it is true, it turns out to be far more
innocuous and less interesting than either racists or egalitarians assume.



VOL. 100, No. 3

The example of Tay-Sachs disease will show why this

is so. Central and Eastern European JewsAshkenazim
historically tended to marry and mate mainly within their
own ethnic group. Endogamous mating within a relatively
small local population like this can be expected to reduce
genetic diversity. In such situations, damaging genetic
variants may by sheer chance accumulate and proliferate
more rapidly than they are being removed by natural selection. This appears to have happened in the Ashkenazic
population. One such variant that occurs in elevated frequencies in Ashkenazim is an ugly recessive lethal mutation that produces a disorder variously called infantile
amaurotic family idiocy, cerebral sphingolipidosis, type I
GM2 gangliosidosis, or Tay-Sachs disease. Heterozygotes
for this mutation (children who inherit it from one parent,
but not from die other) are perfectly normal. But homozygotes (children who inherit it from both parents) never
grow up. In such children, toxic fatty-acid compounds accumulate in the tissues of the nervous system, producing
nerve-cell degeneration, blindness, paralysis, and early
death (Ampola 1982; Volk 1964).
The Tay-Sachs mutation is significantly more common
among people of Ashkenazic ancestry than it is among
other Central and Eastern Europeans (Aronson 1964). It
might well prove to be the case that, when all the TaySachs homozygotes are counted into the picture, the average genetic potential for various mental abilities is lower
at birth among Ashkenazim than it is among non-Jews
from the same area. I do not know whether this is in fact
true, and I am not particularly anxious to find out, because
the answer would be socially and politically uninteresting.
Even if Ashkenazim have on the average lower intelligence than their neighbors, that would not imply that any
particular member of the groupsay, Albert Einsteinis
mentally defective. Congenital mental deficiency is not
caused by belonging to an ethnic group that has a lot of
congenitally stupid people in it. It is caused by carrying
certain combinations of genes in certain environments.
Again, that last phrase needs to be stressed. Because no
gene acts independently of its environment, genetic variants that affect mental abilities in one setting may have no
effect in other environments. The metabolic disease called
phenylketonuria illustrates this rule. Like Tay-Sachs disease, this disorder is caused by a recessive allele. People
who are homozygous for this allele cannot properly metabolize the amino acid phenylalanine. If they ingest it, toxic
compounds accumulate in their tissues and cause neurological damage, resulting in epilepsy and mental retardation. However, such people can avoid this fate simply by
avoiding phenylalanine. If they adhere to a suitably restricted diet from birth onward, they suffer little or no
damage (Ampola 1982; Kaiser and Bickel 1971; Williamson et al. 1971). In environments where phenylketonuriacs are unable or unprepared to avoid phenylalanine, the genetic locus of the phenylketonuria mutation



represents a "gene for intelligence." In other environments, it does not. Our science and technology enable us to
create favorable environments in which phenylketonuria
becomes a relatively harmless genetic variant. Someday,
we may be able to create environments in which TaySachs homozygotes, or other sorts of so-called "congenital mental defectives," are not handicapped. As we alter
our environments in pursuit of such aims, the contribution
of genetic variation to variation in human mental abilities
will decline accordingly.
I suspect that the question of interethnic differences in
average mental abilities attracts more attention than it deserves because some of the people who write about it and
ought to know better are not really thinking about heredity
in terms of particulate Mendelian inheritance. Rather,
they are thinking about it in terms of our folk concepts of
"blood" heredity. Suppose for the sake of argument that,
say, the inhabitants of Ireland make lower average scores
on IQ tests than the inhabitants of Scotland. Average genetic differences between the two populations might well
be contributing to that difference in test results. B ut even if
all this were true, it would not imply that Irish "blood"represents some sort of a hereditary taint that dooms all those
of Irish descent to some degree of congenital thick-headedness. It might simply mean that, say, the phenylketonuria mutation occurs in higher frequencies in Ireland
than in Scotland. Any genetically conditioned difference
in average mental abilities between two human groups
will fit this general description.
As the theory of natural selection would lead us to expect, all genetic variants known to yield gross mental deficiencies in the present range of human environments occur in quite small percentages in every ethnic group, local
population, or "race." No doubt there are undiscovered
variants that produce smaller and subtler deficiencies in
various environments. Such genetic variants will be less
heavily selected against than the seriously deleterious
variants that produce phenylketonuria or Tay-Sachs disease, and therefore may occur in higher frequencies; but
they are equally unlikely to occur uniformly throughout
any ethnic group. A possible exception may occur in cases
where certain phenotypes are culturally interpreted as
markers of ascribed membership in a stigmatized group.
In cultural settings where (say) left-handed or darkskinned or obese children are held to be congenitally stupid, we might expect such children to be differentially neglected, discriminated against, and taught to think poorly
of themselves. In these environments, but not in others,
genetic variants promoting right-handedness, light skin,
or slenderness may turn out to be "genes for intelligence."
Again, it should be emphasized that there is no such thing
as a "gene for intelligence" outside a particular environmental context, and that culture always affects the interaction between genes and environment in our species.


Summary and Conclusions

Almost every sort of human potential is limited by both
environmental and genetic factors, but it makes no sense
to ask whether a particular capacity is limited chiefly by
heredity or by environment. The environment (including
culture) determines the contribution of genetic factors to
phenotypic variation. Genetic variants that affect aphenotypic trait in one setting may have no effect on it in other
environments. Superior or fitter genetic variants are superior only in a specific environmental context. There is no
such thing as "heritability," "fitness," or "biological superiority" in the abstract.
Hereditary differences between human individuals are
real and important, and there are significant average differences in various respects between some regional populations. Correlations between genetics and geography are
a legitimate subject for scientific investigation. However,
these facts do not oblige us to think of human variation in
racial terms. Regional populations that differ significantly
in one respect usually resemble each other, and contrast
with some third population, in certain other respects.
Many regional populations today (e.g., those of North
America) have been largely formed by centuries of massive immigration from widely separate parts of the world.
The sympatric "racial" groups conventionally recognized
within such populations are neither geographically, phenotypically, nor genetically discrete. The aggregate variation within such populations encompasses the entire
range of variation in all the immigrant groups combined,
and any typological "racial" groups that we attempt to distinguish in the population will contain large numbers of
individuals descended from members of the other groups.
If human races are geographically delimited populations characterized by regionally distinctive phenotypes
that do not occur elsewhere in significant numbers, then
races no longer exist and have probably not existed for
centuries, if ever. And if races are not geographically delimited, then racial classificatory categories are merely labels for polymorphisms that vary in frequency from one
part of the world to another, like redheadedness or Type A
blood. If "Negroid" and "Caucasoid" people occur on
every continent, it makes no more sense to describe these
groupings as geographical subspecies than it would to describe redheads or people with Type A blood as human
subspecies. In particular, it makes no sense to try to study
differences between races by subdividing a sample of
North Americans. Yet a lot of the existing literature on
supposed racial differences offers to do just that. Structuring our samples using these chimerical racial categories
often obscures the nature and causes of past and present
human variation.
Like other social constructs, races are real cultural entities. For many people, membership in a racial group constitutes an important part of their social identity and self-



image. But social facts are not necessarily part of the biological landscape. In multiethnic regional populations,
races are merely ethnic groups linked to vague, inconsistent, and stereotypical ideal phenotypes. Growing awareness of the meaninglessness of racial taxonomy is currently leading increasing numbers of U.S. citizens to
refuse to classify themselves racially, or to allow themselves to be so classified by others (Fish 1995). In the long
run, we would probably be better off if we all followed
their example.

Acknowledgments. I am grateful to George Aimelagos, Kayc
Brown, Jon Marks, Dan Schmitt, Ted Steegman, and three
anonymous referees for their helpful comments on earlier drafts
of this paper.
1. For a quick survey of the polar positions among physical
anthropologists on the subject of race, see the recent issue of
Evolutionary Anthropology in which two distinguished physical anthropologists were asked to provide separate reviews of
the same four books dealing with issues of race and human genetic diversity (Aimelagos 1995; Harpending 1995). Each reviewer praised the same books that the other condemned.

References Cited
AAPA (American Association of Physical Anthropologists)
1996 AAPA Statement on Biological Aspects of Race.
American Journal of Physical Anthropology 101:569
Ampola, M.G.
1982 Metabolic Diseases in Pediatric Practice. Boston: Little, Brown.
Aimelagos, G.
1995 Race, Reason, and Rationale. Evolutionary Anthropology 4:103-109.
1964 Epidemiology. In Tay-Sachs Disease. B. W. Volk.ed.
Pp. 118-153. New York: Grune and Stratton.
Barkan, E.
1992 The Retreat of Scientific Racism: Changing Concepts
of Race in Britain and the United States between the World
Wars. Cambridge: Cambridge University Press.
Blakey, Michael L.
1994 Passing the Buck: Naturalism and Individualism as
Anthropological Expressions of Euro-American Denial. In
Race. S. Gregory and R. Sanjek, eds. Pp. 270-284. New
Brunswick, NJ: Rutgers University Press.
19% Skull Doctors Revisited: Intrinsic Social and Political
Bias in the History of American Physical Anthropology,
with Special Reference to the Work of Ales Hrdlicka. /n
Race and Other Misadventures: Essays in Honor of Ashley
Montagu in His Ninetieth Year. L. T. Reynolds and L. Lieberman,eds. Pp. 64-95. Dix Hills, NY: General Hall.
Brace, C.L.
1964 A Non-Racial Approach toward the Understanding of
Human Diversity. In The Concept of Race. M. F. A. Montagu, ed. Pp. 103-152. New York: Free Press.



VOL. 100, No. 3

1996 A Four-Letter Word Called "Race." In Race and

Other Misadventures: Essays in Honor of Ashley Montagu
in His Ninetieth Year. L. T. Reynolds and L. Lieberman,
eds. Pp. 106-141. Dix Hills, NY: General Hall.
1996 Genes, Peuples et Langues. Paris: Editions Odile
Coon, C. S., S. M. Garn, and J. B. Birdsell
1950 Races: A Study of the Problems of Race Formation in
Man. Springfield, IL: Charles C. Thomas.
Crosby, A. W.
1986 Ecological Imperialism: The B iological Expansion of
Europe, 900-1900. Cambridge: Cambridge University
1995 Mixed Blood. Psychology Today, November: 55-87.
Goodman, A. H., and G. J. Armelagos
1996 The Resurrection of Race: The Concept of Race in
Physical Anthropology in die 1990s. In Race and Other
Misadventures: Essays in Honor of Ashley Montagu in His
Ninetieth Year. L. T. Reynolds and L. Lieberman, eds. Pp.
174-186. Dix Hills, NY: General Hall.
1995 Human Biological Diversity. Evolutionary Anthropology 4:99-103.
Harrison, G. A., J. S. Weiner, J. M. Tanner, and N. A. Barnicot
1977 Human Biology: An Introduction to Human Evolution, Variation, Growth, and Ecology. 2nd edition. Oxford:
Oxford University Press.
Herrnstein, Richard J., and Charles Murray
1994 The Bell Curve: Intelligence and Class Structure in
American Life. New York: Free Press.
Kaiser, S., and H. Bickel
1971 Psychological and Developmental Evaluation of
Phenylketonuric Patients Treated with a PhenylalanineRestricted Diet In Phenylketonuria and Some Other Inborn Errors of Amino Acid Metabolism. H. Bickel, F. P.
Hudson, and L. I. Woolf, eds. Pp. 263-269. Stuttgart:
Georg Thieme.
Keita, S. O. Y., and R. A. Kittles
1997 The Persistence of Racial Thinking and the Myth of
Racial Divergence. American Anthropologist 99:534544.
Landau, M.
1997 A People's History of Human Biodiversity. American Anthropologist 99:392-394.
Lewontin, R. C.
1972 The Apportionment of Human Diversity. Evolutionary Biology 6:381-398.
Lieberman, L., andF. L. C. Jackson
1995 Race and Three Model s of Human Ori gin. American
Anthropologist 97:231 -242.
Lieberman, L., and L. T. Reynolds
1978 The Debate over Race Revisited: An Empirical Investigation. Phylon 39:333-343.
1996 Race: The Deconstruction of a Scientific Concept In
Race and Other Misadventures: Essays in Honor of Ashley
Montagu in His Ninetieth Year. L. T. Reynolds and L. Lieberman, eds. Pp. 142-173. Dix Hills, NY: General Hall.


Littlefield, A., L. Lieberman, and L. T. Reynolds

1982 Redefining Race: The Potential Demise of a Concept
in Physical Anthropology. Current Anthropology 23:
Livingstone, F.B.
1962 On the Non-Existence of Human Races. Current Anthropology 3:279-281.
Marks, J.
1995 Human Biodiversity: Genes, Race, and History. New
York: Aldine de Gruyter.
Molnar, S.
1992 Human Variation: Races, Types, and Ethnic Groups.
Englewood Cliffs, NJ: Prentice-Hall.
Montagu, M. F. A.
1942a The Genetic Theory of Race and Andiropological
Method. American Anthropologist 44: 369-375.
1942b Man's Most Dangerous Myth: The Fallacy of Race.
New York: Columbia University Press.
Rushton, J. P., and A. F. Bogaert
1989 Population Differences in Susceptibility to AIDS: An
Evolutionary Analysis. Social Science and Medicine
Sanjek, R.
1994 The Enduring Inequalities of Race. In Race. S. Gregory and R. Sanjek, eds. Pp. 1-17. New Brunswick, NJ:
Rutgers University Press.
1997 Culture as a Stressor: A Revised Model of Biocultural
Interaction. American Journal of Physical Anthropology
Shipman, P.
1994 Facing Racial DifferencesTogether. Chronicle of
Higher Education, August 3: B1-B3.
1992 Proceedings of the Sixty-First Meeting of the American Association of Physical Anthropologists. American
Journal of Physical Anthropology 89:505-515.
1993 Proceedings of the Sixty-Second Meeting of the
American Association of Physical Anthropologists.
American Journal of Physical Anthropology 92:549-560.
1964 Pathologic Anatomy. In Tay-Sachs' Disease. B. W.
Volk, ed. Pp. 36-67. New York: Grune and Stratum.
Weizmann, F., N. I. Wiener, D. L. Wiesenthal, and M. Ziegler
1996 Inventing Racial Psychologies: The (Mis)Uses of
Evolutionary Theory. In Race and Other Misadventures:
Essays in Honor of Ashley Montagu in His Ninetieth Year.
L. T. Reynolds and L. Lieberman, eds. Pp. 187-205. Dix
Hills, NY: General Hall.
Williamson, M., R. Koch, T. Clair, J. Dobson, and Y. Lee
1971 Treatment Effects on Growth and Intell igence among
Phenylketonuric Children. In Phenylketonuria and Some
Other Inborn Errors of Amino Acid Metabolism. H.
Bickel, F. P. Hudson, and L. I. Woolf, eds. Pp. 199-207.
Stuttgart: Georg Thieme.
Wolpoff, M., and R. Caspari
1997 Race and Human Evolution. New York: Simon and


FromTypesto Populations:A Centuryof Race,

PhysicalAnthropology,and the American
ABSTRACT Inthe1960s,U.S.physical
a periodofintrospection
a changefrom
thatwas largely
racebasedto thenewphysical
a decade,
and socialinfluences
on physical
and biologiessentialism,
as cladespersists
as a legacyofthe


CarletonCoon's TheOriginofRacesin 1962 reflected a major change in U.S. physicalanthropology.Coon

suggestedthatfivemajorracesof humansevolvedin parat fivedifferent
allel fromHomoerectus
timesand at different rates. He furthersuggestedthat each racial lineage
crossedthe sapiens"threshold"at different
timesin preand
had been
history implied
in the sapiensstatewas correlatedwith the level of "culturalachievement"of different
racial groups.Coon contended that Causcasoids and Mongoloids crossed this
a claimthatclearly
Capoids) and Australians(Australoids),
had socialimplications.
Race had held immenseimportancewithinthe field
of physicalanthropologyduringthe time leading up to
the publicationof Coon's work.At the emergenceof the
subdiscipline,race was the major theoreticalfoundation
of anthropology;physicalanthropology
was virtuallysynonymouswiththe studyof race. In 1902, at the inception
of the AmericanAnthropological
anthropologistsconsidered"race" to representthe way
the humanspecieswas internally
was implicitin thisidea; a racewas thoughtto represent
naturalcategorywithunique featuresthatdefinedthe essenceof thatcategory.'It seemedobviousto manyanthroAMERICAN ANTHROPOLOGIST 105(1):65-76.

pologiststhat thesebiological subdivisionscorresponded

to the social meaningsof race,a notionthatlinkedphysical and behavioralcharacteristics.
This link betweenthe
componentsof an essenceprovidedthe basis forthe biologicaldeterminism
prevalentin the racialthinkingof the
time.Throughoutthe 20th century,race also had an evolutionarycomponent.Raceswereeffectively
clades. Different
essenceswere explainedas a productof
poorlyunderstoodevolutionaryprocesses,as exemplified
byCoon's notionofindependently
The discourseCoon's book spawned contributedto
currentswithinthe fieldthatultimatelyforcedan end to
the old physical anthropologycenteredmainly on the
raceconceptand helpedusherin thenew physicalanthropology,espoused by SherryWashburn,which had been
developingthroughoutthe 1950s. The new anthropology
was eclectic(incorporating
to genetics)and was an evolutionary
science,whose populationalapproacheswereincompatiblewiththe essentialismcentralto theraceconcept.TheOriginofRacesbrought
to a head the riftswithinphysicalanthropology
as a discipline, the tensionsbetweenthe subdisciplinesof anthropology,and discussionsabout the roleof anthropologyin
the publicarena.
The AAA'sreactionto thebookwas decisive.Washburn,
then presidentof the association, delivereda scathing




AmericanAnthropologist * Vol. 105, No. 1 * March 2003

addressdenouncingthebook aroundthetimeof itspublicationat theAAAAnnualMeetingin Chicago on November 16, 1962. The publishedversion(Washburn1963) is
much less harsh,focusingon the limiteduse of race as a
valid object of studyand the lack of scientificsupportfor
Public denunciationof
any claims of racial inferiority.
Coon's ideas seemed necessary;segregationists
were alto
variety responses
community.Statementson racewereissuedbyboththeAAAand theAmerican Associationof PhysicalAnthropologists
(AAPA).Several edited volumes appeared throughoutthe 1960s
critiquingthe race concept.In 1966, MargaretMead and
Theodosius Dobzhanskyorganizedan AmericanAssociation forthe Advancementof Science (AAAS)symposium
meantto deliverthe scientificvoice againsta popularracism based on "misinformation"
and "evil myths"about
race.As embodiedby itsorganizers,
the symposiumrepresentedan alliancebetweenBoasian culturalanthropology
and evolutionarybiology,includingdiverseperspectives
genetics,ethnology,psychology, and sociology.With few exceptions,most anthropologists had become opposed to hereditarianclaims
about race and intelligence,
and manywerenow skeptical
of the race conceptitself.What became clearby the mid1960s was thatrace was no longera unifyingconceptin
justas it had ceased to
be a unifyingconcept foranthropologyas a whole since
Boas's workon race a halfcenturyearlier.In physicalanthropology,race was now a divisiveconcept. Although
Washburnhad publishedhis ideas about the new anthropologyearlier,this periodmarkeda turningpoint in the
on the
race concept.Some have even arguedthat it markedthe
demiseof theraceconcept.
Severalfactorsinfluencedchangingviews about race
withinphysicalanthropologyduringthis time. First,social factorspromptedscientiststo challengeassumptions
and intellectualinferiority
associatedwiththe race concept.The Holocaustin the 1930s
and 1940s and the controversy
school desegsurrounding
regationin the early1960s may have been the most importantexamples.Anothercomponentof social pressure
resultedfromthe relationshipbetweenanthropologyand
interestin raceand racialinequality,an ingovernmental
terestthat had promotedthe "racialization"of U.S. anthropologyin the firstplace. Second,the race conceptitselfwas challengedby thepopulationalprinciplesespoused
in the modernsynthesis;evolutionaryideas were incompatiblewith the essentialistfoundationsof the race conviewsofpopulationand clines,based
cept,and alternative
largelyon understandings
manyscholarsto considerrace an invalidtool forunderstandingbiologicalvariation.Finally,theevolutionofU.S.
fromitsemergenceas a subfieldto
the presentday, has been influencedby its relationship
withthe restof anthropology-specifically,

thropologyas embodiedby the AAA.It is interesting
as earlyas 1894, a quartercenturypriorto the emergence
of physicalanthropologyas a truesubdiscipline,Boas began to challengethe race concept. By the time physical
anthropologyclearlyemergedin the 1920s,Boas's followers held some of the mostpowerfulpositionswithinU.S.
anthropologyand were a dominant voice in the AAA.
Therefore,the racial physicalanthropologythat was rejectedin the 1960s developedwithina broaderanthropologicalcontextthathad been grapplingwiththe raceconcept foryears;partsofthatcommunityalreadyquestioned
race,and the AAAhad been involvedin struggles
issue of race betweenanthropologyand governmentpolicies and funding,as well as strugglesbetweenanthropologyand othersciences.The rejectionofrace in the 1960s
was not so new;itwas a partoftheheritageof physicalanthropologywithinU.S. anthropology.
This historysuggeststhatthe raceconcepthas no remaininglegacyin physicalanthropology.What actually
changed?Is the race conceptreallydead? What elements
of the race conceptstillpersistand influencephysicalanthropology
today?In thisarticle,I addressthesequestions,
themwithinthecontextofthe scientific
social influenceson mainstreamphysicalanthropology
thatwere a major forcein the evolutionof the race concept. I arguethatsome elementsof the raceconceptwere
in factrejected,but thatothersremain,subtlyinfluencing
ourviewsofwhatwe todayterm"populations."
The race concept that was examined and rejectedby so
manyin the 1960s includesassumptionsabout the cause
and natureofgeographicand otherkindsofvariation.The
historybehind these assumptionshas helped createthe
conceptthatwe grapplewithtoday.Althoughforthe last
100 years the race concept has been thoughtabout in
terms,its most fundamentalelements
clades,and biologicaldeterminism.
attributesare clearlyrelated,and all of them have informedthe theoriesabout human variationin physical
The raceconcepthas changed,yettheseatanthropology.
tributesof racehave not all changedtogether.While bioand its social implicationshave been
questioned since the inceptionof the field,essentialism
and the concomitantrenderingof races as clades have
been less amenableto change.
The racesdefinedby the Westernrace conceptwerecodifiedby Linnaeus(1758) and by the definitive10thedition
Naturae,in whichhe describedfivesubspecies
of humans,listingforeach typeboth the morphological
and behavioralcharacteristics
thatwereconsidereda part
of the essenceof the category.These wereimplicitly(and
explicitly)understoodto be partof theintrinsicbiologyof
the race. European prejudiceswere clearlyincorporated

Caspari * Race:FromTypesto Populations 67

into Linnaean typologiesand taxonomiesintegralto the
naturalhistorytradition.Fromitsveryinception,the race
conceptembodiedboth essentialismand biologicaldeterminism.
In manycases thisessentialism(and the naturalhistorycontextto whichit applied) renderedthinkingabout
raceverysimilarto thinkingabout biologicalspecies.This
is exemplifiedin the polygenismso prevalentin the U.S.
and Frenchschools of thoughtthat dominatedmuch of
anthropologyforthe firsthalfof the 19th century(Brace
1982; Stanton1960; Stocking1968; Wolpoffand Caspari
Even afterthe widespreadacceptance of evolution
and manyelementsof Darwiniantheory,a formof polyscientists
genismcontinuedto thrivebecauseevolutionary
essentialist(and racial) perspective.Taxonomiccategories,includingsubspecificones,continuedto
be conceptualizedas discretegroups,whiletheessencesof
the categorieswereexplainedas productsof separateevolutionaryhistories.Races,like speciescategories,weredepictedas brancheson an evolutionarytree,whose differences could now be explainedthroughtheirindependent
evolution,at different
Clades: EvolutionaryEssentialism
Conformingto the Darwiniannotionof the commondemodels
scentof all species,treemodelsbecameappropriate
fordiagrammingthe relationshipsbetweenspecies.After
splittingfroma common ancestor,two daughterspecies
difisolatedby definitionand represent
are reproductively
intuitiveessenferentbranchesof a phylogeny.Therefore,
tialismand older treemetaphorsdid not impede understandingof evolutionaryprocessesat the species level,
because the categoriesare discrete.However,the storyis
below the specieslevel,because branching
quite different
cannot accuratelyreflectrelationshipsbetween groups
The essentialistlink betweendepictingvariationbetweenspeciesand variationwithinthe humanspecieswas
nowherecleareror moreinfluentialthan in the worksof
ErnstHaeckel.Haeckelused evolutionarytreesbothto describetheplace ofhumansin thenaturalworldand therelationshipsof human races within the human species.
This had unfortunateimplicationsforunderstandings
human variation.As Linnaeantaxonomywas "evolutionized" and relationshipsamong taxa expressedin termsof
evolutionarytrees, human races, like species, became
branches(or twigs)on the tree,each with its own definable essence. This approach providedscientificexplanahuman groupswere effections forhuman differences;
tivelyspecieson a smallerscale, whose differences
be accountedforthroughindependentevolution.
Throughoutthe 19th and 20th centuries,the use of
phylogeniesto characterizehuman relationshipsin sociopolitical spheres provided the conceptual underpinTheyprovidedjusningsof Westernracialclassifications.

tificationfor"interracialcompetition"(Keith 1936), the

basisforclaimsof thebiologicalinferiority
of socialclasses,
and supported
to various eugenic policies and the applied biology of
Nazism (Gasman 1971; Stein 1988; Wolpoffand Caspari
The assumptionof monophylyimplicitin treemetaphorswas made explicitwiththe generalconcensusthat
treebranchesare clades,definedas monophyleticgroups.
A monophyleticgroupincludesan ancestraltaxon and all
its descendents;clearly,races are not monophyleticand
Yet treebranches
theirdepictionas cladesis inappropriate.
are theunderlying
as a naturalcaterepresentation
This construction
underliesmuch of the thinkingpresent
the historyof U.S. physicalanthropology.
Biological Determinism
is implicitto the raceconcept,and
it is this component that has been most ardentlyaddressedby the fieldbecause of its obvious social implications. In the 1960s, biological determinismwas a focal
school depointofimportantcurrentissues(in particular,
was at thecenterofpoliticaldiscussionsthatmanyin the
anthropologicalcommunityof the 1960s foundit importantto address.
anthropologyembodied both a
racial thinkingand evolutionismthat explainedcultural
variation.At the Turnof the Century,virtuallyall social
scientistswereevolutionists,holdingthe idea that primitive races and theirculturesrepresentedstagesin evolulengthson an evotionaryhistoryor branchesof different
biologyand culture
than others,some
biology influencingcultural
change,otherslike Lewis HenryMorgan(1877, reprinted
in 1964), forexample,thoughtcultureaffectedbiological
changein the brain,in a Lamarckiansense. However,biolofculturaldifogywas usuallyconsideredthe determinant
as it was practicedthroughout
much of the 19th century,was a singlebioculturaldiscipline,withrace linkingthe components.FranzBoas sevand while not
eredthisconnectionforU.S. anthropology,
all anthropologists
agreedwithhim,he and his followers
forceda kindof introspection-biological(racial)determicould no longerbe acceptedas
a blanketassumptionin U.S. anthropology.
is nota necessarypartofracial
rejectedwithoutthe rejectionofthe
raceconceptas a whole.Throughout
validityof racial
many anthropologistsquestioned
of culturalcapacitieswithoutcompletelyredetermination
jectingthe race concept and its underlyingessentialism.
in general,thebiologicaldeterminism
race conceptwas deeply
anthropologyas a


AmericanAnthropologist * Vol. 105, No. 1 * March2003

major assumptionof racial studies,and in Europe,racial

was a majorcomponentof anthropological
In the UnitedStates,ironically,giventhe influenceof the
earlier"AmericanSchool," race had become somewhat
less importantthan in Europeananthropology.Becauseof
its emergencewithin(or in some cases beside) the broader
anthropologicaltraditionembodied after 1902 by the
moreof a
strugglebetweenracial (i.e., those who focusedon race)
and nonracialelements--orshould I say "less-racial"elements,because the race conceptreallyunderlayall thinking about human variation.To some extent,the new
espousedby Washburnrepresented
a realliancewith the Boasian partsof anthropologythat
had questionedthe assumptionsof the race conceptsince
the 1890s.
The storyof racein U.S. anthropology(includingphysical
anthropology)cannot be discussedwithoutreviewingthe
roleof Boas and the AAA.This has been treatedby a large
numberof Boas scholars(to name a few:Cole 1999; Stocking 1968; Williams1996), and I onlybrieflyoutlinethese
relationshipshere to underscorethe professionaland politicaltensionsaffecting
as itemerged
as a subdiscipline.
Until the 1920s, there were reallyno U.S. degrees
awardedin what would be specifically
considered"physical anthropology."Nevertheless,
race and racial assumptionsstillplayedan important,
rolein anthropology.The fieldfocusedon NativeAmericanethnology
and archaeologyand was descriptive;
even while
race may have been considereda cause of culturalvariation,itwas not emphasized.The pre-Darwinian
AmericanSchoolofSamuelMortonhad no students,
and althe
Agassiz produced
Ward Putnam,who had a fundamentalinfluence
on the developmentof anthropology
in the UnitedStates,
Putnam'sinterestwas archaeology,notrace.
As anthropologyemergedas a professionat the Turn
ofthe Century,a commitmentto NativeAmericanstudies
and theidea of professionalization
(i.e.,trainingin anthropologyratherthanrelateddisciplines,orworse,none at all)
was what held the earlyassociationtogether,in spiteof
earlydivisionsbetweenthe "Washingtonians"and "Boasians" along this veryline (Stocking1968). Boas was responsibleforthe four-field
trainingof manyearlyPh.D.s
in U.S. anthropology,even those fromoutside his home
institution(Columbia). Unlike the European model for
anthropology,Boas thought trainingin anthropology
should includeall subdisciplines,as did his own research.
At Harvard,Putnamconcentratedon archeology,the fo-

cus of most of the 15 Ph.D.s Harvardproducedbetween

1894-1919. Yet,manyofthesestudentsalso trainedunder
Boas in ethnology,linguistics,
and physicalanthropology
in the trainingof Har(Cole 1999). Boas was instrumental
vard studentssuch as RolandDixon (who was laterto become an influenceon EarnestHooton's racialthinking)as
well as his own famousdescendentsfromColumbia. By
would head every
1926, Boas's students(or sympathizers)
in thecountry(Stocking1968).
It is sometimesforgotten
that Boas was a practicing
physicalanthropologistearlyin his career,probablythe
onlyone trainingstudentsin theUnitedStatesat theTurn
of the Century.In 1894, Paul Topinard(the preeminent
Frenchanthropologistof his time,and studentof Broca)
wrote that Boas was "the man, the anthropologist,I
wishedforin the UnitedStates"(Stocking1968:166).
Boas receivedhis Ph.D. in physicsin 1881 butby that
time had become a geographer.Previouslyuntrainedin
he soughtguidancefromAdolphBastianin
Rudolph Virchowin physicalanthropolbefore
leavingBerlinforBaffinIsland. Boas muchadmiredVirchow,who trainedhim in anthropometrics
1999; Stocking1968). LikeVirchow,Boas was interested
physiologicalprocessesand never became a Darwinian
(i.e., selectionist),althoughhe did recognizecommondescentand humanevolutionary
to thenatural
world.Likemanyothersof his time,Boas had Lamarckian
ideas (see Wolpoffand Caspari 1997: ch. 8) and neverunderstoodselection.He acceptedtheviewof manyGerman
scientiststhat selectioncould only effectsmall changes
as Franz Weidenreichcalled them), not
Moreover,and, perhaps,more importantly,
Boas consideredVirchow'smost significantlegacyto be
the organizationof the fieldin Germany,and, later,Boas
consciouslysoughtto be a similarfigurein U.S. anthropology(Stocking1968).
In the UnitedStates,Boas continuedto make contributionsto physicalanthropology,
whichhe recognized,as
did everyoneat the time,as racial studies.However,insteadof acceptingthe assumptionsofthe raceconcept,he
treatedthemas objectsofinquiry.He wound up rejecting
biological determinismratherearly in the game, and,
later,his workquestioned the validityof human types,
thus challengingthe essentialismat the core of the race
concept.However,he neverreallyrelinquishedessentialist
notions of major races-broad geographicentities-even
as he questionedthe validityof human typesforsmaller
racial categories,such as variousnationalities(e.g., "Nordics" or "Alpines").
His strongestcontributions
to physicalanthropology
whichhe appliedto studiesofmetrichuman
variation.He was veryinterestedin the new biometrics
being advancedby FrancisGalton and KarlPearson,with
whom he corresponded,
and he developedhis own methods of analysisas well. A major outcomeof these studies
was his appreciationoftheimportanceofvariation,which
he used laterto critiquethe idea of racialtypes.This can

Caspari * Race: FromTypesto Populations 69

be juxtaposedwith Hooton's use of Pearson'sbiometrics
yearslater,whichhe used less criticallyto delineateracial
types.Hooton influencedthe developmentofa race-based
physicalanthropologyin the UnitedStates;Boas and his
to itsdemise.
By the time Boas came to Columbia in 1896, he was
deeply concerned with questions of race and had researchedproblemsof variationand change. He was alreadydevelopinghis ideas of relativism,sparkedby his
1884 expeditionto BaffinIsland,and by then had largely
rejectedthe idea that race determinedculturalachievement.As earlyas 1894, he explicitlyrejectedracialdeterminismof culture:"Historicaleventsappearto have been
much more potent in leading races to civilizationthan
theirfaculty,and it followsthatachievementsof racesdo
not warrantus to assume that one race is more highly
giftedthananother"(Boas 1894:303).
He thoughtphysicalanthropologywas importantin
even acceptingraces as "real,"he recognizedthe importanceof environment
and historyas influenceson human
Boas was interestedin growthand
partof physicalanthropology,
(environmentaland hereditary
affectson growth)
influencedthe modificationofinheritedform.Priorto movingto Columbiain 1896, he initiateda studyof Worcesterschoolchildrenin which he
statisticallydemonstratedthe problems with inferring
longitudinal informationfrom cross-sectionalstudies
(and, thus,advocatedforlongitudinalstudiesin growth)
and emphasizedforthe firsttimethe importanceof variation in temposof growth.Thus, beforethe Turnof the
Century,he was lookingat human variationin nonracial
ways,more interestedin the impactof the environment
(includingculture)on biologythan the affectof biology
(race)on culture.
Boas investigated
BritishAssociationfor the Advancementof Science. He
looked for relationshipsbetween heredityand environbetweenreservation
Indians of the NorthPacific
and nonreservation-dwelling
Coast. He also looked at problemsof racialadmixturein
in racialhybrids.In these"mixed"popureducedfertility
variationin cranialproportions,
he noted the distribufacial
tionwas bimodaland not normal.In theseand otherstudies, averagesdid not representthe "type."He laterundermined the conceptof "type,"questioningits meaning:If
averagesdid not represent
oftraits,not theconformawas thedistribution
of interest
tion to typesor the creationof new,intermediate
thatwas criticalto the race conthe case of interbreeding
cept. His mostfamousworkregardingracewas performed
between 1908-10 on head shape in U.S. immigrants,
funded (somewhat ironically)by the U.S. Immigration
Commission,which was seekinga scientificbasis to re-

In a studyof over 18,000 immigrants,
he found changes in head formthat underminedthe
dogmaof thestabilityof racialtypesand the Europeanfocus on head shape as a major indicatorof race. He could
not explainthe causes of change,althoughhe considered
themin some way "environmental"
and, as an empiricist,
importantwas the documentation
of thechangeitself.Throughhis workon racialquestions,
Boas challengedboth biologicaldeterminism
and the nature of racial categories,two criticalcomponentsof the
race concept.These challengeswere a centralpartof anthropologicalthinkingin the U.S. beforeHooton started
producingPh.D.sin physicalanthropology.
"Racializing" PhysicalAnthropology
Both governmentaland privateforcespromotedthe renaissanceofthe "scienceof race"in U.S. anthropology
develI, running
opingin muchof U.S. anthropology.
also tensionsbetween Boasians and other,at the time,
smallerfactionsof the AAAwho weresympathetic
to the
anthropologistsassociated with Washingtoninstitutions
(Stocking1968). It mustbe remembered
startedproducingPh.D.s at Harvardin 1925,therewas no
specifictrainingin physicalanthropologyin the United
States.Only six U.S. Ph.D.s in physicalanthropologyhad
been awardedpriorto 1925-five of thesefromHarvard,
trainedby specialistsin otherdisciplines.Ales HrdliEka,
the founderofAmerican
1918, and AAPAin 1928, had no studentsin his position
at theNationalMuseumofNaturalHistory.
Duringthe second decade of the 20th century,many
scholarswho claimedto representphysicalanthropology
were actually eugenicistsfrom other disciplines (that
claimed scientificsuperiority
to anthropology),
and some
wereverypowerfulin theU.S. scientific
Theseincludedmembersofthe GaltonSociety,whichwas
dedicatedto the studyof racialanthropology,
such as the
the AmericanMuseumof NaturalHistory),and the biologistsRaymondPearland JohnC. Merriman(head of the
NationalResearchCouncil [NRC]).Manyin the anthropologicalcommunitysaw themas a threat;theywereracial
deterministswith a political agenda, and the BoasiandominatedAAAdid not acceptthemas anthropologists.
Therewas clearlya need forphysicalanthropologists
This dearthbecame veryapwhen
Councilsoughtto form
an anthropology
whichwas to deal withphysicommittee,
cal anthropologyand eugenics.Aside fromHrdliekaand
Boas, therewerefewphysicalanthropologists
by the AAAto serve(Stocking1968). Madison Grantand
Charles Davenport,ardentracistsand eugenicists,founders of the Galton Societywith strongpoliticalagendas
and tiesto Washington,servedon the originalcommittee.
Whilethe AAArefusedto recognizeGrantor Davenportas


AmericanAnthropologist * Vol. 105, No. 1 * March 2003

thereremainedenormouspressureon ananthropologists,
thropology "racialize,"both from the government,
which had become increasinglyinterestedin restricting
immigrationon racialgrounds,and fromthe eugenicinterestscontrollingothermajor fundingsources.Members
of the Galton Societyincluded the heads of institutions
thathad been (or potentiallycould be) importantsources
of anthropologicalfunding.By the 1920s, fundingincreasedforstudiesof race and racialpsychology.U.S. anthropologistsrespondedto this fundingincreaseand to
criticismsthat they neglected biology and the racial
makeupofthe U.S. byexpressingmoreinterestin physical
anthropology.Ironically,severalof Boas's students(e.g.,
Mead, Herskovitz,
Klineberg)werefundedby NRC fellowships in the biological sciences forworkthat supported
the culturalbasis forracial differences,
and Boas himself
on race. Other
studentsof Boas, such as AlfredE. Kroeber(and Roland
Dixon), as well as more conservativenon-Boasian elements of the anthropologicalcommunity,also became
in physicalanthropology,
placingthe race
concept and eugenicsat the focus of the emergingnew

EarnestHooton was one of the mostinfluentialfiguresin
physicalanthropology(Giles 1999). He was a professorat
Harvardfrom1913 until his death in 1954 and was responsible fortrainingvirtuallyan entireacademic field,
spawningseveralgenerationsof studentswhen fewother
offeredphysicalanthropologyas partof their
curricula.Hooton's Ph.D. (in 1911 fromWisconsin)was in
classics.He had littlebackgroundin anthropology,
and it
took some time to get his programoffthe ground,but
startingin 1926, a floodof Ph.D.s in physicalanthropology emergedfromHarvard.Withina fewyears,physical
anthropologywas a majorpartof U.S. anthropology,
this was reflectedin AAAmembershipand the developmentoftheAAPA.
Race studiescame to be the focusof Hooton's career,
but he formedhis ideas about race and physicalanthropologyin generalafterhe came to Harvard.His workwas
typologicaland manifested,like Haeckel's,as an "evolutionarypolygenism"(Wolpoffand Caspari1997). Hooton's
Harvardcolleague, Dixon, influencedhis views on race;
Hooton's 1931 classification
is verysimilarto Dixon's 1923
The polygenismof Hooton and Dixon was complicated,groundedin the beliefin once-pureracesthathad
with fundamentallypolygenistideas, they understood
thatpresenthumanvariationcould not be accommodated
withina fewracial types.Hooton thoughtthat the complexity of human variation could be accounted for
recentlyunderwenta secondaryrace

formationstage and then a tertiary

stage--ineach stage,
ones. Thus,they
hybridracesformedfromthe preexisting
argued,pure racesexistedand persistedinto the present,
but secondaryand tertiary
human evolutionthroughhybridization.
Hooton's views
were stillessentialist;he believedin "pure" races,but he
realizedthata fewracialtypescould not accountforreal
Hooton'sthinkingabout racehad all the attributes
the race concept;he was an essentialist,
he explainedthe
branches,and he was a biological determinist
as is clearlyshown in his eugenicwriting
(Hooton 1937, 1939; Wolpoffand Caspari 1997). Given
this, Hooton's views on racismcould appear paradoxical
(Wolpoffand Caspari 1997). While he believed in races,
and even in "racialcharacter,"he was more active than
most membersof the academic communityin antiracist
activities,enteringinto a relationshipwith Boas that
Barkantermed"the frustrated
antiracistcampaignof an
odd anthropologicalcouple" (1988:182). Aftermany attemptsto mobilizethe academic communityagainstracism,Boas turnedto Hooton,who senta statementhe had
authoredto seven leading U.S. physicalanthropologists
outlininghis view on the stateof scientificknowledgeof
race differences.
Amongotherthings,he concludedthata
correlationbetweenphysicalfeaturesand mentalability
had not been demonstratedand that therewas insufficient scientificevidence to assign evolutionaryranksto
races.Only Hrdli6kawould sign it. In 1936, Hooton then
published his own "Plain Statementabout Race" in Science,speakingagainstthe racismunderlyingNazism. In
of attemptsto organ1940, as Hooton realizedthe futility
ize even the AAPAagainstracism,his studentWilliamW.
Howellsasked him what could be done. He replied:"Not
only has the horsebeen stolen,but thebarnhas also been
It is hardto overestimate
U.S. physicalanthropology.
Hooton'sthinkingon racewas
adoptedby some ofhis students,rejectedby others,but in
eithercase, it stronglyinfluencedsubsequentgenerations
of scholarsbecauseit limitedtheirunderstanding
ofdifferent ways of interpreting
studentssuch as Howells,who largelyrejectedHooton's
views(Caspariand Wolpoff
1996; Wolpoffand Caspari 1997). For instance,Howells
was so conditionedby polygenicmodels that he did not
interpretFranz Weidenreich'spolycentricmodel of human evolutionas a network,
as it was originally
(Weidenreich1946). Followingan initialexchangein the
(AA) (Howells 1942; Weidenreich
1940), Howells describedWeidenreich'sideas as a polygenic tree (the "candelabra").Even afterdiagramsof the
trellisappeared (Weidenreich1946, 1947), Howells continuedto depictit as a candelabrain numeroussecondary
sourcesand textbooksthroughout
his career(e.g., Howells
1959,1993).Itmaybe thatHowellswasinpartreacting
polygenismof R. RugglesGates, a racistplant geneticist

Caspari * Race:FromTypesto Populations 71

who derivedsupportfrombothHootonand Coon, thinking
Weidenreichsharedtheseviews(Gates 1944; Wolpoffand
(1946) specifically
and the candelabraHowellsdescribed
was actuallymore like Hooton's model,and not Weidenreich's."Tree-thinking"
saw the"candelabra"as a reasonablesimplification
an oversimplification)
of a trellis,but one thatrepresented
of racesciencedid not permeatephysicalanthropologyin
the UnitedStatesforlong. It was nevertheoverwhelrhing
traditionthat it had been in Europe,and his students,
only a single generationlater,were responsiblefor the
new physical anthropologythat disavowed the importance of race. Washburn,the most well-knownamong
these,activelyrejectedthe racialthinkingofhis mentor;it
thatWashburn'sfirstjob was at Columis not surprising
he joined those predominantly
the Boasians,includingAshleyMontagu.
While few of his studentsshared Hooton's eugenic
manyof themcontinuedhis focuson race and human variation,at least fora while. Some, such as Stanley
Garn and Coon, focusedon problemsof race definition,
races,and problemsofraceformation(e.g., Brues1972; Coon et al. 1950; Garn1957, 1962).
and manyotherscontributedto discusThese researchers
sions about the numberof races-some recognizedhundreds,some onlya few.Authorssuch as Coon et al. (1950)
suggestedit was just a matterof resolution:Microraces
could be definedby a largernumberof traitsand representedsubdivisionsof broad major raceswhose constituents uniquelyshareda smallernumberof traits.Some of
natureof racialclassifithesestudiesimpliedthe arbitrary
cation.Coon et al. (1950) wroteon the potentialadaptive
significanceof racialtraits.HarryShapiro(1939) and Fred
Hulse (1962), also studentsof Hooton and interestedin
draquestioningthe stabilityof racialtraits,demonstrated
maticmorphologicalchangesin first-generation
in Hawaii,similarto Boas's conclusionsearlier
in the 20th century.Some students,such as WilliamSheldon in his famoussomatotypestudies,retainedHooton's
biological determinism(and in Sheldon's case expanded
on it); others,such as Coon, inheritedHooton's polygenism;stillothers,such as Washburn,rejectedHooton's
emphasis on race, turninginstead to the evolutionary
ideas underlyingthe modernsynthesisas the foundation
ofthenew physicalanthropology.
With the modernsynthesisof the 1940s, Hooton's
studentsalso facedthe need to bringevolutionarytheory
into theirstudies.They did this in different
ways. Coon
was a typologistwho never incorporatedpopulational
thinkinginto his perspective;however,he considered
himselfan evolutionist,
largelythroughhis interestin adof
aptation.He did not extendthis to an understanding

populationalprocesses,a focuson variationwithinpopulations,or on the fluidityof populations(Wolpoffand

came to be
Caspari1997). The new physicalanthropology
viewedas the studyof human evolution,not the description of human types. Some, like FrederickHulse, addressedthisby lookingat racesas evolutionaryepisodes,
viewingracesas largelyephemeral,causedbyevolutionary
processes.Washburnsoughtto developa new physicalanthropologywithoutrace,groundedin evolutionarybiologyand the populationalthinkingof the synthesis.Only
threeyears afterthe Princetonsymposiumthat marked
the "official"birthof the modernsynthesis,Washburn
and Dobzhanskyorganizedthe famous15thAnnualCold
SpringHarborSymposiumthatclarifiedthe evolutionary
In additionto
programof the new physicalanthropology.
a focus on human evolution and prehistory,the new
physicalanthropologyespoused ways that biology was
relevantto studiesof the human condition-thatbiology
and culturecould be interrelated
of intions"insteadof "races,"or studiedthe distribution
dividualtraitsin clinalstudies(Brace1964).
Ironically,Hooton himselfcontributedto the changthroughhis skillsas an
educatorand his respectforhis students(Giles 1999). As
HarryShapiropointsout in Hooton's obituaryin the AA,
Hooton encourageddiversityof thoughtin his students.
He did not want to establisha "school" and "neverattemptedto establishintellectualascendancyover his students" (1954:1082). He encourageddissentingopinions,
tellingShapiro: "You know, none of my studentshave
been yesmen. ... ThankGod!" (Shapiro1954:1082). Hooton's studentsremaineddiverse,as theyestablishedphysiand museumsaroundthe
in universities
cal anthropology
and racialapmaintained
least in part,forwhat has been consideredthe demiseof
Public Science
The verypublicrejectionof raceby manyanthropologists
in the 1960s was one of a numberofresponses,beginning
in the 1930s,by the scientific
communityto racismin the
thinkingabout the race conlargersociety.
developmentof the modcept
ernsynthesisin biology,and the applicationof its principles to human variation and evolution (not only by
but also by the architectsof the synthesis
of its architects,especially Ernst
Mayr (1982, 1991) and Dobzhansky(1944, 1962, 1963)
saw thepopulationalthinkingofthe synthesisand emergof populationgeneticsto be influential
ing understandings
weapons in a warwaged by scienceagainstpublic racism.


AmericanAnthropologist * Vol. 105, No. 1 * March2003

However,the reactionsparkedby Coon's publicationof

TheOriginofRaceswas also a responseto Coon's tacitalliance withracistsseekingto influencepublic policy(Jackson 2001).
as well as otherscientists,had
Some anthropologists,
been activein antiracismcampaignssincethe early1930s.
The abuses of biologyand anthropologythatwereat the
root of the eugenicsmovementand Nazi biopolicyproand evolupelledat leasta fewbiologists,anthropologists,
to presentscientific
tionistswitha sense of responsibility
argumentsthatwould underminethis"scientific"racism.
coaliThis is when Boas and Hooton formedtheirfruitless
tionto generatesupport
U.S. academia. The BritishevolutionarybiologistJ. B. S.
Haldane spoke out against racismat the 1934 London
Meetingof the InternationalCongressof Anthropological
and EthnologicalSciences(ICAES),warninghis audience
againstthe abuse of sciencein supportof race theories.In
1935, respondingto risingracismin Europe,JulianHuxley
an important
and AlfredHaddon publishedWe Europeans,
antiracisttract.In additionto underminingbiologicaldeterminismand assumptionsof racial inferiority,
questionedthe veryexistenceof race and suggestedthat
ethnicgroupreplacethe termrace,a harbingerof Ashley
Montagu's1942 Man's MostDangerousMythand his 1950
UNESCO statementon race.
Montagu,who receivedhis Ph.D. withBoas at Columbia afterstudyinganatomywith G. Elliot Smithin Lonat the forefront
don, was theU.S. physicalanthropologist
of the public antiracismcampaign after the war. He
authorednumerouspopulararticlesand books,as well as
the firstUNESCO statementon race,which was verycontroversialbecause of his claim thatraces were a "myth,"
not because of his denunciationof notionsof differences
in racialcapacitiesforachievement.
In the 1960s,an even largergroupof scientistssought
to underminethe scientificracismused to supportopponentsof the civil rightsmovement.This reactionwas especiallystrongin the anthropologicalcommunity.Once
again, as in the days of Madison Grantand othertimes
the AAAfounditselfpittedagainst
groups seekingto influencepublic racial policy in the
name of science.CarletonPutnamand othersdirectlyattackedthe AAAas a left-wing
concealed the "truth"about race. Coon was squarelyin
the middle of all this (Jackson2001), contrathe mostly
(Coon 1981) depictionsof him as a purely
whose work was misused by others
withouthis approval(Shipman 1994). As the civil rights
movement became strongerand the Supreme Court
laws (activelyresistedin the South),
passed desegregation
Coon subtlyparticipatedin movementsmeantto undermine Boasian interpretations
of race. Coon was sympatheticto thesegregationist
Pamphletsand books such as Race and Reason(1962)
by CarletonPutnam(Coon's cousin) used Coon as scientificauthority.These publicationsconsciouslypittedthe

subdisciplinesof anthropologyagainsteach other,claiming that "scientific"anthropologists(like Coon) rejected

the dismissalof race and that theyhad evidenceof racial
inequalitythat made blacks undeservingof full citizenship. These writingshad wide circulation;theywerepublishedin newspapersthroughouttheSouth,and therewas
even a "PutnamLettersCommittee"dedicatedto raising
funds to publish the lettersin Northernnewspapers,
and were
where they appeared as paid advertisements
used as mass mailingsof segregationist
requiredreadingin the
Louisiana public schools (Jackson2001)--evidence of its
prominencein the South.
AttacksfromPutnamand otherracistslikeHenryGarrett(1961) and Wesley George (1962) promptedresolutions on race fromboth the AAAin November1961 and
the AAPAin 1962. Froma pressreleaseon the 60th Annual Meetingof the AAA,GordonWilley,then president
ofthe AAA,called fora resolutionin responseto "publicaas a basisforsocial and
tionson raceand racialdifferences
believe to be false
of our professionby personswho
and misrepresentative
are not recognizedby theAmericanAnthropological
Association as professionalanthropologists"(Jackson2001:
263). The resolutionpassedunanimously.
A fewmonthslater,the AAPApassed a resolutionintroducedby Stanley Garn that specificallycondemned
Race and Reasonand the misuse of science withinit. Followingtheresolution,Coon resignedfromthepresidency
of the AAPA,claimingthe resolutionwas inappropriate
and thatscientistsshouldkeep out ofthe integration
the scenesof the segregationist
cause throughhis associationwithPutnamand others(Jackson2001).
Some of the authoritysegregationists
citedalso came
fromeugenicistswhose work (by the 1960s withoutthe
eugenicslabel) continuedto be fundedby Wycliffe
founder of the Pioneer fund, and other like-minded
sources,whichtodaycontinuesto fundresearchmeantto
demonstratehuman inequality.This line of research,and
its financial foundation,representsa thread running
throughoutthe historyof U.S. anthropology(Lieberman
have embraced,but that
2001) thata fewanthropologists
the communityat largehas consistently
in spite
oftheirinfluenceat thetime,boththe AAAand the AAPA
have continuedto denyanthropologicalidentityto their
thesedescendentsalso claim they are estrangedfromthe field
because of the left-wing,
politicalcorrectnessof anthrouse
politicalclaims to deflectcriticismoftheirwork(Relethford
2001). Nevertheless,
produced and fundedby incarnationsof the same foundationsthatsupportedsimilarworkthroughout

Caspari * Race:FromTypesto Populations 73

Public attentiontherefore
discussionabout the validityof race withinphysicalanthropology;questionsabout the intellectualcapacitiesof
racialgroupswere addressed,as were questions
of the "reality"of race. Some individualsrejectedat least
some elementsof theraceconcept.However,thisrepudiation of the typeconceptwas more directlyinfluencedby
evolutionarybiology;because of evolutionaryand genetic
influences,the newergenerationof physicalanthropologists grew up thinkingabout human variationin ways
thatwerenot (at leastexplicitly)racial.

Genetics,Populations,and Clines
The need to confrontpublicmisrepresentations
thatwereactivelyused in the 1960s fosteredalliancesbetween various elements that had foughtracism before
with some that had not-the architectsof the modern
synthesis,"mainstream"anthropologistsas represented
as represented
by the AAA,and physicalanthropologists
Mead-Dobzhanskysymposium represented
thisalliance,as did a numberofvolumes
on the studyofraceproducedat the time(e.g.,Mead et al.
1968; Montagu 1964) thatbroughttogetherworkfroma
varietyof disciplines.One way of attackingracismwas
(and is) througha focus on the inadequacy of the race
conceptforexplaininghumanvariation.Studiesof clines,
the geographicdistributionof individualmorphological
and genetic traits,were introducedand population replaced race as a focus of study.This was by no means
purelypolitical;it was a consequenceof the evolutionary
approachofthenew physicalanthropology.
As C. LoringBrace pointed out in 1964, races,and
even populations,are inadequateforthe studyof human
variation.Instead,he advocatedforthe studyof individual traits-thestudyoftheirdistribution
and the selection
thatcauses theirvariation.The studyof clinescame to replace race as a focus of analysis for many researchers.
in his 1962 articleon the nonexistence
of human races,eloquentlylays out whyraceor any subspecifictaxonomyis misleading:
is notonly
to anypartofsuchvariation
buttendsto obscure
He was a strongproponentof nonracialclinal studies,arguing,"thereare no races,onlyclines"(Livingstone1962:
Others,however(e.g., Brues 1972), acceptedthe importanceof clines but arguedthat the biologyof groups
themselveswas also a valid targetof inquiry.With the
populational thinkingof the modern synthesis,which
formedthe basis of Washburn'snew physicalanthropology,populationsreplacedrace as the unit of study.What
did this mean? How did the studyof populationsdiffer
fromrace? Mayr himselfsuggeststhat the populational

thinkinghe developedhelped bringabout the demiseof

the race concept,as essentialismis the antithesisto Darwinian approachesto variation.By emphasizingintraspecificevolutionary
on variationand the fluidity
betweenpopulations-on all
the processesthatreduceor increasevariationwithinand
betweenpopulations.An emphasison gene flowand its
the disrelationshipto otherevolutionaryforcesaffecting
tributionof differenttraitsacross populations is what
populationalthinkingis all about, and it underminedthe
race concept. The approach is verydifferent
used to understandphyleticevolution,the focusof many
and when populationsare studied
withtheoriesand methodsappropriateforphyleticanalyses, the workis no longerpopulational,but essentialist.
When thesetwo verydifferent
confusedwithone another,populationsare treatedmuch
as racesonce were,and theworkdoes not represent
Therefore,despite the shiftin focus fromrace to
populationas a unit of study,populationalthinkingdoes
not necessarily
go hand-in-handwiththe studyofpopulations.Many 20th-century
conceivedofpopuing genes (Boyd 1950) or morphology,
lation as just another term for race. They thoughtof
populationsas breedingpopulations,isolatedfromother
groups.Some recognizedthis implicitly,some explicitly.
Garn wrote,"the contemporaryapproach to race stems
frompopulation genetics,where a race is viewed as a
breedingpopulation,neithermore nor less" (1962:6). He
identifiedsmall"local races"likethe "Bushmen"ofSouth
Africaas moreor less isolatedbreedingpopulations(Garn
1962). In spite of Washburn's(1963) admonitionthat
racesor populationswereopen systems,populationswere
theexistconceptualizedas closed.Therefore,
ence of typeswas implicit,even ifthe scientific
on theiradaptations.AsArmelagos,Carlson,and Van Gerven (1982) pointedout, manystudiesin skeletalbiology
and geneticscontinuedto employtypologicalmethodsto
typologicalends:the recognitionand delineationofpopulations.Theirconclusionin 1982 was thatwhetherusing
skeletalor genetictraits,many studiesof populationsare
justas typologicalas studiesofrace.
In spiteof the typologicalapproachof some genetic
studies,geneticshad a stronginfluenceon the changing
race concept, especiallythe population geneticsof the
modern synthesis.Population geneticiststhroughthe
yearshave providedcompellingevidenceforhumanunity.
In 1972, whenRichardC. Lewontinmade famoustheestimates of much more variationwithinthan betweenhuman groups,he was showingwhat populationgeneticists
like Dobzhanskyhad suspectedand said all along. While
Dobzhanskyargued that races were not a "myth,"and
that there were biological differencesbetween populations,he arguedfortheirfluidityand forthe conceptof
isolation by distance.More recently,Alan Templeton,a
geneticistwith anthropologicaltrainingwritingin the


AmericanAnthropologist * Vol. 105, No. 1 * March 2003

pages of AA, looked at race froma geneticsperspective,

showingthat subspeciesdo not existin humansand emphasizingthattreemodelsdo not adequatelydescribehuman populationrelationships
However,tree models have continued to thrive.As
discussedearlier,the polygenictreewas such a powerful
metaphorin the thinkingof the 1940s and 1950s that
as a tree.
Weidenreich'snetworkwas originallyinterpreted
evoluThis historicbackgroundcontinues
tionarythinkingabout humanpopulationstoday.
Relationshipsbetweenpopulationsare still oftendepicted as branches on a tree,thereforeimplyinginterisolation
are due to reproductive
fromother groups.The magnitudeof that difference
the extentof the relationshipis sometimessaid to reflect
the length of time since the populations diverged.Althoughthisis appropriateforspecies(whichcannotinterbreed),and may even somewhataccuratelydepict intergroup relationships in species with marked genetic
betweengroups,branchingmodelsdo not dedistinctions
scribehuman relationships.Ironically,the geneticliteraTreesarecommonheutureis fullof suchrepresentations.
risticdevices used to depictvariationof geneticsystems.
at this level,
While treescan be valid forrepresentation
if the
theyfailto accuratelypredictpopulation
gene divergencesare assumedto reflectthe relationships
betweenpopulations.Worldwideanalysisof different
genetic systemsshows that theyhave different
variation-that is, their treeshave different
likemtDNAhave shallowcoalescencetimes,while others
likebeta-globinand HLA have much deeperones (Hawks
et al. 2000). Because of recombination,the historiesof
geneseven withinthe same individualare different.
because gene historiesare not digene tree is different
rectlylinked togetherin population histories(Harding
2000; Hawkset al. 2000; Relethford
1998). Yet,in muchof
a treederivedfroma singlegenetic
the geneticliterature,
the historyofpopulations.
systemis assumedto represent
How did the raceconceptchangein the 1960s?Whatelementswerealtered?Can we reallycelebrateitsdemise?Of
the threeattributesof race discussed,biologicaldetermiof such traitsas intelligence,
nism,or racial determinism
has been most activelyaddressedsince the beginningsof
the AAA,and despitemisgivingson the partof some culturalanthropologists,
the physicalanthropologycommunitylargelyrejectsit today.Withthe growthof evolutionary psychologyand behavioralecology,theremay be a
resurgenceof emphasison the biological basis of behavioraltraits,but,forthe mostpart,these studiesrecognize
the difference
betweenevolutionaryfoundationsand biologicaldeterminism.
The link betweenbiological determinismand racial
determinismdepends on races being naturalcategories,

and physicalanthropologistsno longersupportthe notion that races are subspecies.The importanceof gene
flowand the fluidityof the speciesis recognizedeven by
whose continueduse of typological and raciallycharged termsmakes them appear less
populationalthan theyoftenreallyare. However,in spite
of the rejectionof racesas subspecies,and a reluctanceto
use the term race,populations are oftenthoughtof in
much the same way that raceswere in the earlierliterature. Essentialismcontinues to influence conceptions
about humangroups,and thisis exemplified
by the use of
treesas metaphorsforhuman populationrelationshipsin
studiesof morphologicaland genetichumanvariation.Inclades are an enduringlegacyof racial anthrotraspecific
pologyand continueto informour thinkingabout populations.The raceconceptmaybe rejectedby anthropology,
but itsunderlyingracialthinking
pologists longerstudy
Populationsare now studied, butnot all approachesto the studyofpopulationsare

MI 48109-1382

I wouldliketothank
me to participate
in thissymposium
and inspiring
me to think
in thecontext
and an anonymous
I especially
1. Ultimately,
raceis a taxonomy
basedon socialfactors,
Becauseall taxonomies
is a critical
is a product
mayhavea psychological
worldin similar
social,and physical
(beyondthe information
given)about constituent
tobe part
to particular
domains(basedon different
have been termed"livingkinds."Peoplelearn"livingkinds"
thanthoseused to learn
aboutinanimatethingsor the processesthatrelateto them.
a cognitive
domainthatallowsthemto easilylearn"humankinds"
ofa particular
to a culture;
turetobe intrinsic
toa person's
Justas biological
abouttheessenceof a species(or a dog
theyaresupposedto looklike,thinklike,
ofa category
do notconformto thestereotype
does notdispelthestereotype.

Caspari * Race: FromTypesto Populations

of "humankinds"areconmentalessence.Becauseclassifications
theymayhave greater
social meaningthan othercategoriesthat do not reflectthe essenceofa person.In U.S. society,presumedgeographicoriginand
phenotypicfeatures,widelyconsideredthe constituentcompoto identity
nentsof race,are consideredto be moreintrinsic
in Western
othercategorieslikeoccupationor religion.Therefore,
society,and globallyto some extentbecauseof culturalinterconnection,Westerndominance,and the legacies of colonialism,
"race"is a "humankind"and, therefore,
has a psychologicaldimensionsinceitis basedon thesamecognitivedomain.According
to this reasoning,we may be psychologically
disposedto racial
(1996) has said, thatracial
thinking.This suggests,as Hirschfeld
and the raceconceptare not one and the same;the race
conceptmay be a productof "mentation,"but racialthinkingis
culturaland psychological.

GeorgeJ.,DavidS. Carlson,and DennisP. Van Gerven
1982 TheTheoretical
1990 Cognitive
1994 CoreDomainsvs.Scientific
1988 Mobilizing
Essayson Biological
W. StockingJr.,
1894 HumanFaculty
as Determined
1974 TheShapingofAmerican
1950 GeneticsandtheRacesofMan.Boston:Little,
Brace,C. Loring
ofHu1964 ANonracialApproach
InTheConceptofRace.M. F.AshleyMontagu,
1982 TheRootsoftheRaceConceptinPhysical
1972 ModelsofClinesandRaces.American
Caspari,Rachel,and MilfordH. Wolpoff
1996 Coon andWeidenreich.
Cole, Douglas
1999 FranzBoas:TheEarlyYears,1858-1906.Seattle:University
1939 TheRacesofEurope.NewYork:MacMillan.
1950 HumanRacesinRelation
on Quantitative
1962 TheOriginofRaces.NewYork:Knopf.
1965 TheLivingRacesofMan.NewYork:Knopf.
1981 Adventures
S. Coon. Englewood
Coon,C. S., S. M. Garn,andJ.B. Birdsell
1950 Races:AStudyoftheProblems
IL:C. C. Thomas.
1923 TheRacialHistory


1944 On Speciesand RacesofLivingandFossilMan.American
1962 MankindEvolving:
1963 Possibility
1957 RaceandEvolution.
1962 HumanRaces.Rev.edition.Springfield,
IL:C. C. Thomas.
1961 TheEqualitarian
1971 TheScientific
1944 Phylogeny
1962 TheBiologyoftheRaceProblem.
DC: National
1999 Hooton,Earnest
vol. 11.Pp.147-149.John
A.GaratyandMarkC. Carnes,eds.
1905 TheWondersofLife.NewYork:Harper.
2000 Diversity,
Facultede MedicineLariboisiereSaint-Louis,
Hawks,John,KeithHunley,Sang-HeeLee,and MilfordH. Wolpoff
2000 Bottlenecks
1996 RaceintheMaking:Cognition,
1998 Natural
LawrenceA.,and SusanA. Gelman,eds.
1994 MappingtheMind:DomainSpecificity
1931 Upfrom
1936 PlainStatements
1937 Apes,Men,andMorons.NewYork:G. P.Putnam.
1939 Twilight
ofMan.NewYork:G. P.Putnam.
1942 FossilManandtheOriginofRaces.American
1959 MankindintheMaking.GardenCity,NJ:Doubleday.
1993 Getting
DC: CompassPress.
1962 Raceas an Evolutionary
C. Haddon
Huxley,JulianS.,and Alfred
1935 WeEuropeans:
2001 "InWaysUnacademical":
1936 History
1972 TheApportionment


AmericanAnthropologist * Vol. 105, No. 1 * March 2003

2001 How"Caucasoids"GotSuchBigHeadsandWhyThey
1758 Systemae
F. B.
1962 On theNon-Existence
1982 TheGrowth
1991 One LongArgument:
RobertE. Light,eds.
1968 ScienceandtheConceptofRace.NewYork:ColumbiaUniPress.
1942 Man'sMostDangerousMyth:TheFallacyofRace.NewYork:
1964 TheConceptofRace.NewYork:FreePress.
1964[1877] Ancient
1962 RaceandReason:AYankeeView.Washington,
DC: Public
1998 GeneticsofModemHumanOrigins

2001 Commenton Lieberman,

Shapiro,H. L.
1939 Migration
1954 Earnest
1994 TheEvolution
1960 TheLeopard'sSpots:Scientific
1988 BiologicalScienceandtheRootsofNazism.American
1968 Race,Culture,
ofAnEssayson theHistory
1998 HumanRaces:AGeneticandEvolutionary
1963 TheStudyofRace.American
An1940 SomeProblems
1946 Apes,Giants,andMan.Chicago:University
1947 FactsandSpeculations
1996 Rethinking
M. H.,and R.Caspari
1997 RaceandHumanEvolution.

1 of 4

About Human Races and the Confusion

By R.C. Lewontin

Published on: Jun 07, 2006

R.C. Lewontin, Alexander Agassiz Professor Emeritus of Zoology at Harvard University, has written a number of books and
articles on evolution and human variation, including Biology as Ideology: The Doctrine of DNA and The Triple Helix: Gene,
Organism, and Environment

Over the last thirty five years a major change has taken place in our biological understanding of the concept
of human race, largely as a consequence of an immense increase in our knowledge of human genetics. As
a biological rather than a social construct, race has ceased to be seen as a fundamental reality
characterizing the human species. Nevertheless, there appear from time to time claims that racial

categories represent not arbitrary socially and historically defined groups but objective biological divisions
based on genetic differences. The most recent widely noticed rebirth of such claims is an essay by Armand

Marie Leroi on the Op-Ed page of The New York Times (March 14, 2005), an essay that illustrates both the
classical confusions about the reality of racial categories and the more recent erroneous conclusions about
the relevance of such racial identifications for medical practice.

There are four facts about human variation upon which there is universal agreement. First, the human

species as a whole has immense genetic variation from individual to individual. Any two unrelated human
beings differ by about 3 million distinct DNA variants.

Second, by far the largest amount of that variation, about 85%, is among individuals within local national
or linguistic populations, within the French, within the Kikuyu, within the Japanese. There is diversity

from population to population in how much genetic variation each contains, depending upon how much
immigration into the population has occurred from a variety of other groups and also on the size of the

population. The United States, with a very large population whose ancestors came from all over the earth

including the original inhabitants of the New World, is genetically very variable whereas small populations
of local Amazonian tribes are less genetically variable, although they are by no means genetically uniform.
Despite the differences in amount of genetic variation within local populations, the finding that on the

average 85% of all human genetic variation is within local populations has been a remarkably consistent

result of independent studies carried out over twenty-five years using data from both proteins and DNA.
Of the remaining 15% of human variation, between a quarter and a half is between local populations within
classically defined human races, between the French and the Ukrainians, between the Kikuyu and the

Ewe, between the Japanese and the Koreans. The remaining variation, about 6% to 10% of the total human
variation is between the classically defined geographical races that we think of in an everyday sense as
identified by skin color, hair form, and nose shape. This imprecision in assigning the proportion of

variation assigned to differences among population within races as compared to variation among races,

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arises precisely because there is no objective way to assign the various human populations to clear-cut

races. Into which race do the Hindi and Urdu speakers of the Indian sub-continent fall? Should they be
grouped with Europeans or with Asians or should a separate race be assigned to them? Are the Lapps of
Finland and the Hazari of Afghanistan really Europeans or Asians? What about Indonesians and

Melanesians? Different biologists have made different assignments and the number of races assigned by
anthropologists and geneticists has varied from 3 to 30.

Third, a small number of genetic traits, such as skin color, hair form, nose shape (traits for which the genes
have not actually been identified) and a relatively few proteins like the Rh blood type, vary together so that
many populations with very dark skin color will also have dark tightly curled hair, broad noses and a high

frequency of the Rh blood type R0. Those who, like Leroi, argue for the objective reality of racial divisions

claim that when such covariation is taken into account, clear-cut racial divisions will appear and that these
divisions will correspond largely to the classical division of the world into Whites, Blacks, Yellows, Reds

and Browns. It is indeed possible to combine the information from covarying traits into weighted averages
that take account of the traits' covariation (technically known as "principal components" of variation).
When this has been done, however, the results have not borne out the claims for racial divisions. The
geographical maps of principal component values constructed by Cavalli, Menozzi and Piazza in their

famous The History and Geography of Human Genes show continuous variation over the whole world
with no sharp boundaries and with no greater similarity occurring between Western and Eastern

Europeans than between Europeans and Africans! Thus, the classically defined races do not appear from

an unprejudiced description of human variation. Only the Australian Aborigines appear as a unique group.
A clustering of populations that does correspond to classical continental "races" can be acheived by using a
special class of non-functional DNA, microsatellites. By selecting among microsatellites, it is possible to

find a set that will cluster together African populations, European populations, and Asian populations, etc.
These selected microsatellite DNA markers are not typical of genes, however, but have been chosen

precisely because they are "maximally informative" about group differences. Thus, they tell us what we

already knew about the differences between populations of the classical "races" from skin color, face shape,
and hair form. They have the added advantage of allowing us to make good estimates of the amount of
intermixture that has occurred between populations as a result of migrations and conquests.

The every-day socially defined geographical races do identify groups of populations that are somewhat
more closely similar to each other genetically. Most important from the standpoint of the biological
meaning of these racial categories, however, most human genetic variation does not show such "race"

clustering. For the vast majority of human genetic variations, classical racial categories as defined by a
combination of geography, skin color, nose and hair shape, an occasional blood type or selected

microsatellites make no useful prediction of genetic differences. This failure of the clustering of local

populations into biologically meaningful "races" based on a few clear genetic differences is not confined to
the human species. Zoologists long ago gave up the category of "race" for dividing up groups of animal

populations within a species, because so many of these races turned out to be based on only one or two
genes so that two animals born in the same litter could belong to different "races."

In his article, Leroi is inconsistent and shifting in his notion of race. Sometimes it corresponds to the

classical social definitions of major races, but elsewhere he makes race coincident with a small local

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group such as the Negritos or Inuit. In this shifting concept of race he goes back to the varying use of the
term in the 19th century. Then people spoke of the Scots race, the Irish race and the race of

Englishmen. Indeed race could stand for a family group defined by male inheritance, as in the
description of the last male in a family line as the last of his race. This inconsistent usage arises from the
fact that there is no clear criterion of how much difference between groups of genetically related

individuals should correspond to the category race. If it had turned out that groups of related populations

were clearly different in the great majority of their genes from other groups, then racial categories would be
clear and unambiguous and they would have great predictive power for as yet unstudied characters. But
that is not the way it has turned out, at least for the human species.

The fourth and last fact about genetic differences between groups is that these differences are in the

process of breaking down because of the very large amount of migration and intergroup mating that was
always true episodically in the history of the human species but is now more widespread than ever. The

result is that individuals identified by themselves or others as belonging to one race, based on the small

number of visible characters used in classical race definitions, are likely to have ancestry that is a mixture
of these groups, a fact that has considerable significance for the medical uses of race identification.
A common claim, repeated by Leroi, is that racial categories are of considerable medical use, especially in
diagnostic testing because some genetic disorders are very common in ancestral racial populations. For

example sickle cell anemia is common among West Africans, who were brought as slaves to the New
World, and Tay-Sachs disease is common among Ashkenazi Jews. So, it is argued, racial information can

be a useful diagnostic indicator. Certainly classical race contains some medically relevant information in

some cases, as for example white as opposed to African American if the contrast is between Finland and
West Africa, but not if it is a contrast between a white Mediterranean and an Asian Indian. There is a

confusion here between race and ancestry. Sickle cell anemia is in high frequency not only in West Africans

but also in some white Middle Eastern and Indian populations. Moreover, a person with, say, one African
great-grandparent, but who is identified by herself and others as white has a one in eight chance of

inheriting a sickle-cell mutation carried by that ancestor. There are, in addition, a number of other simply
inherited hemoglobin abnormalities, the thalassemias, that are in high frequency in some places in the
Mediterranean (Sardinia), Arabia and southeast Asia. The highest frequency known for a thalassemia
(80%) is in Nepal, but it is rare in most of Asia. The categorization of individuals simply as white or

Afro-American or Asian will result in a failure to test for such abnormal hemoglobins because these

abnormalities do not characterize the identified race of the patient. Even group identities below the level
of the conventional races are misleading. Two of my incontrovertibly WASP grandchildren have a single
Ashenazi Jewish great-grandparent and so have a one in eight chance of inheriting a Tay-Sachs

abnormality carried by that ancestor. For purposes of medical testing we do not want to know whether a
person is Hispanic but rather whether that persons family came from a Caribbean country such as Cuba,
that had a large influx of West African slaves, or one in which there was a great deal of intermixture with
native American tribes as in Chile and Mexico, or one in which there was only a negligible population of
non-Europeans. Racial identification simply does not do the work needed. What we ought to ask on

medical questionnaires is not racial identification, but ancestry. Do you know of any ancestors who were
(Ashkenazi Jews, or from West Africa, from certain regions of the Mediterranean, from Japan)? Once
again, racial categorization is a bad predictor of biology.

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There has been an interesting dialectic between the notion of human races and the use of race as a general

biological category. Historically, the concept of race was imported into biology, and not only the biology of
the human species, from social practice. The consciousness that human beings come in distinct varieties
led, in the history of biology, to the construction of race as a subgrouping within species. For a long time
the category race was a standard taxonomic level. But the use of race in a general biological context

then reinforced its application to humans. After all, lots of animal and plant species are divided into races,
so why not Homo sapiens? Yet the classification of animal and plant species into named races was at all

times an ill-defined and idiosyncratic practice. There was no clear criterion of what constituted a race of

animals or plants that could be applied over species in general. The growing realization in the middle of the
twentieth century that most species had some genetic differentiation from local population to local

population led finally to the abandonment in biology of any hope that a uniform criterion of race could be
constructed. Yet biologists were loathe to abandon the idea of race entirely. In an attempt to hold on to the

concept while make it objective and generalizable, Th. Dobzhansky, the leading biologist in the study of the
genetics of natural populations, introduced the geographical race, which he defined as any population

that differed genetically in any way from any other population of the species. But as genetics developed and
it became possible to characterize the genetic differences between individuals and populations it became

apparent, that every population of every species in fact differs genetically to some degree from every other
population. Thus, every population is a separate geographic race and it was realized that nothing was
added by the racial category. The consequence of this realization was the abandonment of race as a
biological category during the last quarter of the twentieth century, an abandonment that spread into
anthropology and human biology. However, that abandonment was never complete in the case of the

human species. There has been a constant pressure from social and political practice and the coincidence
of racial, cultural and social class divisions reinforcing the social reality of race, to maintain race as a

human classification. If it were admitted that the category of race is a purely social construct, however, it
would have a weakened legitimacy. Thus, there have been repeated attempts to reassert the objective
biological reality of human racial categories despite the evidence to the contrary.

Genetic Similarities Within and Between Human Populations

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Genetics. 2007 May; 176(1): 351359.

doi: 10.1534/genetics.106.067355

PMCID: PMC1893020

Genetic Similarities Within and Between Human Populations

D. J. Witherspoon,* S. Wooding, A. R. Rogers, E. E. Marchani,* W. S. Watkins,* M. A. Batzer, and L. B. Jorde*,1

Department of Human Genetics, University of Utah Health Sciences Center, Salt Lake City, Utah 84112, Department of Anthropology, University of

Utah, Salt Lake City, Utah 84112, McDermott Center for Human Growth and Development, University of Texas Southwestern Medical Center, Dallas,
Texas 75390 and Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana 70803

1Corresponding author: Department of Human Genetics, Eccles Institute of Human Genetics, University of Utah, 15 N. 2030 E., Room 7225, Salt Lake
City, UT 84112-5330. E-mail:
Communicating editor: L. E

Received 2006 Oct 25; Accepted 2007 Feb 5.

Copyright 2007 by the Genetics Society of America
This article has been cited by other articles in PMC.


Go to:

The proportion of human genetic variation due to differences between populations is modest, and individuals
from different populations can be genetically more similar than individuals from the same population. Yet
sufficient genetic data can permit accurate classification of individuals into populations. Both findings can be
obtained from the same data set, using the same number of polymorphic loci. This article explains why. Our
analysis focuses on the frequency, , with which a pair of random individuals from two different populations is
genetically more similar than a pair of individuals randomly selected from any single population. We compare
to the error rates of several classification methods, using data sets that vary in number of loci, average allele
frequency, populations sampled, and polymorphism ascertainment strategy. We demonstrate that classification
methods achieve higher discriminatory power than because of their use of aggregate properties of populations.
The number of loci analyzed is the most critical variable: with 100 polymorphisms, accurate classification is
possible, but remains sizable, even when using populations as distinct as sub-Saharan Africans and Europeans.
Phenotypes controlled by a dozen or fewer loci can therefore be expected to show substantial overlap between
human populations. This provides empirical justification for caution when using population labels in biomedical
settings, with broad implications for personalized medicine, pharmacogenetics, and the meaning of race.
DISCUSSIONS of genetic differences between major human populations have long been dominated by two facts:
(a) Such differences account for only a small fraction of variance in allele frequencies, but nonetheless (b)
multilocus statistics assign most individuals to the correct population. This is widely understood to reflect the
increased discriminatory power of multilocus statistics. Yet B
et al. (2004) showed, using multilocus
statistics and nearly 400 polymorphic loci, that (c) pairs of individuals from different populations are often more
similar than pairs from the same population. If multilocus statistics are so powerful, then how are we to
understand this finding?

All three of the claims listed above appear in disputes over the significance of human population variation and
race. In particular, the A
(1997, p. 1) stated that data also show
that any two individuals within a particular population are as different genetically as any two people selected from
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Genetic Similarities Within and Between Human Populations

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any two populations in the world (subsequently amended to about as different). Similarly, educational material
distributed by the H
(2001, p. 812) states that two random individuals from any one
group are almost as different [genetically] as any two random individuals from the entire world. Previously, one
might have judged these statements to be essentially correct for single-locus characters, but not for multilocus
ones. However, the finding of B
et al. (2004) suggests that an empirical investigation of these claims is
In what follows, we use several collections of loci genotyped in various human populations to examine the
relationship between claims a, b, and c above. These data sets vary in the numbers of polymorphic loci
genotyped, population sampling strategies, polymorphism ascertainment methods, and average allele frequencies.
To assess claim c, we define as the frequency with which a pair of individuals from different populations is
genetically more similar than a pair from the same population. We show that claim c, the observation of high ,
holds with small collections of loci. It holds even with hundreds of loci, especially if the populations sampled
have not been isolated from each other for long. It breaks down, however, with data sets comprising thousands of
loci genotyped in geographically distinct populations: In such cases, becomes zero. Classification methods
similarly yield high error rates with few loci and almost no errors with thousands of loci. Unlike , however,
classification statistics make use of aggregate properties of populations, so they can approach 100% accuracy
with as few as 100 loci.

Data sets: Three

Go to:

data sets were used. Loci or individuals with >10% missing data were not included in any data
set (loci were pruned first and then individuals). The first data set (insertions) consists of 175 polymorphic
transposable element insertion loci (100 Alu and 75 L1) previously genotyped in 259 individuals. The population
sample consists of 104 individuals from sub-Saharan Africa, 54 East Asians, 61 individuals of northern European
ancestry, and 40 individuals from Andhra Pradesh, India (W
et al. 2005; W
et al. 2006). The
second data set (microarray) consists of 9922 biallelic single-nucleotide polymorphism (SNP) loci genotyped in
278 individuals (55 Africans, 42 African Americans, 40 Native Americans, 22 Indians, 20 East Asians, 62
Europeans, 18 HispanoLatinos from Puerto Rico, and 19 individuals from New Guinea). This data set is derived
from that of S
et al. (2005). The third data set (resequenced) is derived from the 10 ENCODE regions of
the HapMap project, release 16c.1 of phase I, June 2005 (I
2005). These
regions were resequenced in 48 individuals to identify SNPs without ascertainment bias in favor of loci with
common polymorphisms. These SNPs were then genotyped in 209 unrelated individuals: 60 Yoruba in Ibadan,
Nigeria (YRI); 60 Utah residents with ancestry from northern and western Europe (CEU, from the CEPH
diversity panel); and 89 Japanese in Tokyo, Japan, plus Han Chinese in Beijing, China (CHB + JPT). Our subset
consists of 14,258 SNPs. All markers in all three data sets are biallelic. The proportions of missing genotypes are
2.4, 2.1, and 0.36%, respectively.

Data subsampling: To

examine the effect of population sampling (i.e., the effects of comparing relatively isolated
populations vs. more closely related or admixed ones), two subsets were constructed from each of the insertions
and microarray main data sets: one consisting of the entire data set, with all its labeled populations, and another
consisting of East Asian, European, and sub-Saharan African population groups only. The resequenced data set
consists only of the latter three population groups.
To investigate the effect of allele frequency, these five data subsets were subdivided according to three further
treatments: loci with common polymorphisms (with minor allele frequency, MAF, > 0.1); loci with rare
polymorphisms (MAF < 0.1); and all polymorphic loci, regardless of frequency. Henceforth we refer to these
classes of loci as rare polymorphisms, common polymorphisms, or all polymorphisms. For this classification,
allele frequencies were computed across the entire sample in the parent data set. To investigate the effect of
incrementally increasing the number of loci used, loci from each of these 15 data subsets were sampled (without

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Genetic Similarities Within and Between Human Populations

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replacement) to produce 200 independent data sets with numbers of loci varying in 21 steps on a logarithmic scale
from 10 to the maximum.

Pairwise genetic distance: We

use the shared alleles genetic distance (C

and J 1993; B
et al. 1994; M
and C
1997), which defines the distance between two individuals at a locus
as one minus half the number of alleles they share. The genetic distance between individuals is the average of
their per-locus distances. Pairs of individuals are classified as within population or between population
according to whether the individuals were sampled from the same or different groups of populations as defined
be the probability that a pair of individuals randomly chosen from different
populations is genetically more similar than an independent pair chosen from any single population. We compute
all possible pairwise genetic distances, classify them as within- or between-population distances (the sets dW or
dB, respectively), and then calculate the frequency with which dW > dB (that is, a within-population pair is more
dissimilar than a between-population pair). This fraction,
is an estimator of . The expected value of ranges
from 0 to 0.5 (regardless of the number of populations). At = 0, individuals are always more similar to
members of their own population than to members of other populations; at = 0.5, individuals are as likely to be
more similar to members of other populations as to members of their own. The distributions of pairwise genetic
distances implied here resemble the common ancestry profiles proposed by M
and R
(2005), who use a different measure of genetic distance. The shared-alleles distance used here generally yields
slightly lower values of

Dissimilarity fraction

: Let

Centroid misclassification rate CC: The

centroid classification method is also based on pairwise genetic distances,

with one critical difference: Every individual is compared to the centroid of each population, rather than to every
other individual. The centroid is the genetic average of a population, an individual whose pseudogenotypes at
each locus are the frequencies of the genotypes in that population (not including the individual being compared to
the centroid). This genetic distance is equivalent to the average of the genetic distances from an individual to all
other individuals in the target population. Each individual is then assigned to the population with the closest
centroid, as in C
et al. (1999). These assignments are compared to the known populations of origin, and
the proportion of individuals misclassified is reported as CC. The expected classification error for random
assignment of individuals to populations is 1 1/n, where n is the number of populations.

Population trait value misclassification rate CT: Our

definition of CT is implicit in the theoretical illustrations of

et al. (2002) and E
(2003). These authors used simplified models to show how modest differences
between populations can nonetheless enable accurate classification. In both cases, population membership is
treated as an additive quantitative genetic trait controlled by many loci of equal effect, and individuals are divided
into populations on the basis of their trait values.

This method is inherently limited to dividing individuals into just two clusters using only biallelic loci, so we
limit our definitions to that situation. Consider individuals sampled from two populations, A and B, and
genotyped at many biallelic loci. At each locus, we identify the allele whose frequency is higher in population A
and assign it a value of 0. The other allele (more frequent in B than in A) is assigned a value of 1. Let qij represent
the genotype of individual i at locus j, defined as the average of the assigned values of the two alleles carried by
that individual at that locus. Now define qi as the average of qij over all loci j (so qi is a polygenic quantitative
genetic trait). Given these definitions, if populations A and B are typified by even slightly different allele
frequencies at many loci, then qi will usually be smaller for a member of population A than for a member of
population B. Thus the value of the trait qi indicates membership in one population or the other, so we call qi the
population trait value of individual i.
Individuals are assigned to population A or B depending on whether their population trait value qi falls below or
above some dividing criterion qC, respectively. In the case of just two populations, these assignments are

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compared to the known origins of the individuals, and the proportion misclassified is reported as CT. The
classification criterion qC is chosen as follows. Let
be the mean of qi taken over all individuals in population
A, and define
similarly for population B. If the distributions of qi for individuals from the two populations are
symmetric with equal variance, then letting qC = ( + )/2 minimizes misclassification (cf. R
et al. 2002;
2003). To better account for unequal variances, we generalize slightly and solve for a criterion qC such
that r(qC) = s(qC) and
< qC <
where r and s are normal probability density functions with means and
variances estimated from the distributions of qi for populations A and B, respectively.
To extend this inherently pairwise approach to more than two populations, assignments for each individual are
initially computed with reference to each possible pair of populations. The values (0 or 1) assigned to particular
alleles, the criterion qC, and all qi are calculated anew for each pair of populations. Individuals are finally
assigned to a population only if they were assigned to it in all pairwise comparisons involving that population.
The proportion of individuals misclassified (or not classified, since this method can fail to classify individuals) is
reported as CT. For comparison, a single-locus classification error rate is computed by using this method to
classify individuals using each locus singly and then averaging the results over all loci.

Go to:

The statistics
CC, and CT are closely related by design. To illustrate the relationships
between them, the distributions of the genetic measures that underlie them are shown in Figure 1. For simplicity,
only two populations (Europeans and sub-Saharan Africans) and 50 typical loci randomly chosen from the
insertions data set are used. The distributions of pairwise genetic distances for within- and between-population
pairs of individuals (Figure 1A) overlap considerably even for these geographically isolated populations. The
dissimilarity fraction,
is 20%, indicating that between-population pairs are more similar than within-population
pairs one-fifth of the time. In contrast, the distributions of individuals' distances to the centroids of their own or
different populations (Figure 1B) show much less overlap, resulting in CC = 4.2%. The population trait value
distributions for Africans and Europeans overlap for just three individuals, yielding CT = 1.8%. Classifications
using model-based methods such as Structure (P
et al. 2000) achieve 90% accuracy or better using the
same data (B
et al. 2003; W
et al. 2006).

Distributions of distances:

Frequency distributions of the underlying genetic measures used to
CC, and CT, for a subset of 50 loci genotyped in 104
sub-Saharan African and 61 European individuals of the insertions data set.
The measures shown are (A) 13,530 pairwise genetic ...

The variances of the distributions are much greater for the individual-to-individual comparisons (Figure 1A) than
for the centroid-to-individual comparisons (Figure 1B). The distribution means are nearly identical, however, so
the distributions overlap more in Figure 1A than in 1B, and thus > CC. The difference in variances is due to the
fact that each genetic distance to a centroid (each datum in Figure 1B) is equivalent to the average of a sizable
subset of pairwise genetic distances represented in Figure 1A (see
). That averaging step
eliminates considerable variation and produces the narrower distributions of Figure 1B.

The simplifications introduced by R

et al. (2002) and E
(2003) allow an alternative view, represented
in Figure 1C. Here, each individual i is assigned a unidimensional genetic location qi (the individual's population
trait value; see
). The trait distance between any two individuals x and y is now just the
horizontal distance between them, |qxqy|. This simplification is possible only in the two-population case and
requires a population-specific coding of allele states, so the trait distance is not equivalent to the genetic distances
represented in Figure 1, A and B. Nonetheless, it is instructive to consider the analogy using Figure 1C as a guide.

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For example, an African individual x with qx = 0.52 will be more similar to a European y with qy = 0.60 than to
another African z with qz = 0.4. Yet that individual x will still be closer to the population mean trait value for
Africans (qA 0.48, the African centroid) than to the mean value of Europeans (qB 0.68). It follows that many
individuals like this one will be correctly classified (yielding low CC and CT) even though they are often more
similar to individuals of the other population than to members of their own population (yielding high ).

To empirically and quantitatively understand the relationships and contrasts between and the misclassification
rates CC and CT, we examine three primary factors that influence them: the number of polymorphic loci used, the
allele frequencies at those loci, and the degree of differentiation between the populations examined.

Data subset statistics: Three

data sets, labeled insertions, microarray, and resequenced, were used, and 15 subsets
were constructed from these to examine the effects of different data collection strategies (see
). Table 1 lists the 15 data subsets and reports
CC, and CT (each computed over all loci in each data
subset) as well as the expected value of CT when only a single locus is used. Table 1 also gives values of five
descriptive statistics for each data subset: the proportion of genetic variance explained by interpopulation
differences (FST); the observed proportions of heterozygotes (% het); the absolute differences in allele
frequencies between population pairs (averaged), ; the fraction of polymorphisms that are rare (MAF < 0.1) in at
least one population and at the same time common (MAF > 0.1) in another population (% rare and common); and
the fraction of loci that are monomorphic in at least one population and common polymorphisms in another (%
fixed and common). The values observed are typical of human population genetic data sets (N 1973; D
al. 1994; I
2005; S
et al. 2005; W
et al. 2006).
Data set descriptions and summary results
Figure 2 shows the dependency of
CC, and CT on the number of loci for
each of the 15 data subsets listed in Table 1. As the number of included loci increases,
CC and CT decrease.
This is the expected behavior for CC (S
and C
1986; M
et al. 2002; C
et al.
2003) and CT (R
et al. 2002; E
2003). However, does not decrease nearly as rapidly. Figure 2A
shows the results for a diverse sample of individuals genotyped at 175 insertion loci, a number that is typical of
many studies of human genetic diversity published during the last decade. The downward trend in is apparent,
but even with the full data set it remains at 15% (with all four population groups; Table 1). Across all data sets
and using <100 polymorphisms, generally exceeds 10% (Figure 2). With <100 loci, then, it will often be the
case that two individuals from different populations are more similar to one another than are two individuals from
the same population.

Dependency of

on number of loci:

Behavior of the dissimilarity fraction ( ) and error rates of the centroid
(CC) and population trait (CT) classification methods (red, blue, and
green lines, respectively) for each of 15 data subsets (see Table 1 and ...
The power of large numbers of common polymorphisms is most apparent in the microarray data set, comparing
the European, East Asian, and sub-Saharan African population groups (Figure 2C). approaches zero (median
0.12%) with 1000 polymorphisms. This implies that, when enough loci are considered, individuals from these
population groups will always be genetically most similar to members of their own group. In general, CC and CT
decrease more rapidly and to lower values than

Allele frequency effects: The

rare polymorphism subsets defy this trend by converging toward high values of

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as loci are added. This is largely because the frequencies of rare polymorphisms are necessarily quite similar
across populations, whereas higher-frequency polymorphisms have the potential to differ more. For example, the
frequency of an allele with an overall MAF of 5% can differ by at most = 10% between two populations (absent
in one, at 10% frequency in another). This situation yields > 0 and very poor classification accuracy, since most
between-population pairs are identical but some within-population pairs differ. In contrast, an allele with an
overall frequency of 50% across two populations could be fixed in one and absent in the other, resulting in = 0
and allowing perfect classification. It is these frequency differences that allow populations to be distinguished, so
the data sets with lower (and thus generally lower FST) have lower classification power.
The sensitivity of these statistics to allele frequencies explains some differences between the data sets. The
microarray data set exhibits strong ascertainment bias for common polymorphisms, and it is with this data set that
drops most rapidly and to its lowest values (Figure 2, C and D). The insertions data set exhibits a weaker
ascertainment bias and includes more rare polymorphisms, so remains higher (Figure 2, A and B). Similarly,
CC and CT drop more rapidly for the microarray data set than for the insertion data set. The resequenced data set
polymorphisms were ascertained by resequencing a sizable panel of individuals from the genotyped populations
and thus include many rare polymorphisms, but this is partially offset by the equally large number of common
polymorphisms (Figure 2E). The classification methods are less affected by the inclusion of rare polymorphisms.

Population sampling effects: We

contrast two choices: sets of populations that have been relatively isolated from
each other by geographic distance and barriers since the earliest migrations of modern humans out of Africa and
sets that include populations that were founded more recently, are geographically closer to one another and
therefore more likely to exchange migrants, or have recently experienced a large genetic influx from another
population in the set. Sampling only from the more distinct populations yields lower -values, as expected.
Figure 2, A, C, and E, shows the results of using only the three most distinct population groups (Europeans, East
Asians, and sub-Saharan Africans). Figure 2, B and D, expands the samples used in Figure 2, A and C, to include
recently founded and/or geographically intermediate populations (Indians in the insertions data set and New
Guineans, South Asians, and Native Americans in the microarray data set) and admixed populations (i.e., those
that have recently received many migrants from different populations, such as the African American and
HispanoLatino groups in the microarray data set). With just 175 loci, choosing to sample distinct populations vs.
more closely related ones makes only a modest difference (insertions data set, compare Figure 2A to 2B; Table 1).
The effect of population sampling becomes more pronounced when 1000 loci are available. In the microarray
data set, drops to zero at 1000 loci if only distinct populations are sampled. With geographically intermediate
and admixed populations added, however, reaches an asymptotic value of 3.1%, CC remains well above zero,
and even CT does not reach zero (microarray data, Figure 2, C and D; Table 1).
also appears to reach a nonzero asymptotic value in the resequenced data set, instead of continuing to trend
downward as would be expected given the distinct populations used. This may be due to the fact that many of the
polymorphisms in that data set are physically linked and therefore nonindependent. Overall, the responses of the
two classification methods to data set composition variables are qualitatively similar to the behavior of (
Figure 2). The most apparent difference is that the misclassification rates (CC and CT) decrease much more
rapidly, and to lower values, than does as the number of loci considered increases.

Go to:

It has long been appreciated that differences between human populations account for only a small fraction of the
total variance in allele frequencies (typically presented as FST values of 1015%; L
1972; N and
1972; L
1980; B
et al. 1997; J
et al. 2000; W
et al. 2003;
2005; R
et al. 2005). Such observations triggered controversy
from the outset. Some geneticists concluded the differences were negligible (L
1972); others disagreed
1978). Despite the limited data, it soon became apparent that even a modest number of loci should allow
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accurate assignment of individuals to populations (M

1978; S

et al. 1982).

More recently, the H

(2001) (HGP) highlighted the basic genetic similarity of all humans,
yet subsequent analyses demonstrated that genetic data can be used to accurately classify humans into populations
et al. 2002, 2005; B
et al. 2003; T
and E
2003; T
et al. 2005; L et
al. 2006). R
et al. (2002) and E
(2003) used theoretical illustrations to show why accurate
classification is possible despite the slight differences in allele frequencies between populations. These
illustrations suggest that, if enough loci are considered, two individuals from the same population may be
genetically more similar (i.e., more closely related) to each other than to any individual from another population
(as foreshadowed by P
and T
1978). Accordingly, R
et al. (2002, p. 2007.5) state that two
Caucasians are more similar to each other genetically than a Caucasian and an Asian. However, in a reanalysis of
data from 377 microsatellite loci typed in 1056 individuals, Europeans proved to be more similar to Asians than
to other Europeans 38% of the time (B
et al. 2004; population definitions and data from R
et al.
With the large and diverse data sets now available, we have been able to evaluate these contrasts quantitatively.
Even the pairwise relatedness measure,
can show clear distinctions between populations if enough
polymorphic loci are used. Observations of high and low classification errors are the norm with intermediate
numbers of loci (up to several hundred). These results bear out the observations of B
et al. (2004). The
high observed there was due primarily to the slow rate of decrease of with increasing numbers of loci.
Although R
et al. (2002) achieved a very low misclassification rate with the same data, far more loci
would be needed to reduce to similarly small values (assuming such values could be reached at all for those

Thus the answer to the question How often is a pair of individuals from one population genetically more
dissimilar than two individuals chosen from two different populations? depends on the number of
polymorphisms used to define that dissimilarity and the populations being compared. The answer, can be read
from Figure 2. Given 10 loci, three distinct populations, and the full spectrum of polymorphisms (Figure 2E), the
answer is
0.3, or nearly one-third of the time. With 100 loci, the answer is 20% of the time and even using
1000 loci,
10%. However, if genetic similarity is measured over many thousands of loci, the answer becomes
never when individuals are sampled from geographically separated populations.

On the other hand, if the entire world population were analyzed, the inclusion of many closely related and
admixed populations would increase
This is illustrated by the fact that and the classification error rates, CC
and CT, all remain greater than zero when such populations are analyzed, despite the use of >10,000
polymorphisms (Table 1, microarray data set; Figure 2D). In a similar vein, R
et al. (2002) and S
and P
(2004) have suggested that highly accurate classification of individuals from continuously sampled
(and therefore closely related) populations may be impossible. However, those studies lacked the statistical power
required to answer that question (see R
et al. 2005).

How can the observations of accurate classifiability be reconciled with high between-population similarities
among individuals? Classification methods typically make use of aggregate properties of populations, not just
properties of individuals or even of pairs of individuals. For instance, the centroid classification method computes
the distances between individuals and population centroids and then clusters individuals around the nearest
centroid. The population trait method relies on information about the frequencies of each allele in each population
to compute individual trait values and on the means and variances of the trait distributions to classify individuals.
The Structure classification algorithm (P
et al. 2000) also relies on aggregate properties of populations,
such as HardyWeinberg and linkage equilibrium. In contrast, the pairwise distances used to compute make no
use of population-level information and are strongly affected by the high level of within-groups variation typical
of human populations. This accounts for the difference in behavior between and the classification results.

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Since an individual's geographic ancestry can often be inferred from his or her genetic makeup, knowledge of
one's population of origin should allow some inferences about individual genotypes. To the extent that
phenotypically important genetic variation resembles the variation studied here, we may extrapolate from
genotypic to phenotypic patterns. Resequencing studies of gene-coding regions show patterns similar to those
seen here (e.g., S
et al. 2001), and many common disease-associated alleles are not unusually
differentiated across populations (L
et al. 2006). Thus it may be possible to infer something about an
individual's phenotype from knowledge of his or her ancestry.

However, consider a hypothetical phenotype of biomedical interest that is determined primarily by a dozen
additive loci of equal effect whose worldwide distributions resemble those in the insertion data set (e.g., with =
0.15; Table 1). Given these assumptions, the genetic distance used in computing and CC is equivalent to a
phenotypic distance, so Figure 2 can be used to analyze this hypothetical trait. Figure 2A shows that a trait
determined by 12 such loci will typically yield = 0.31 (0.200.41) and CC = 0.14 (0.0540.29; medians and
90% ranges). About one-third of the time ( = 0.31) an individual will be phenotypically more similar to
someone from another population than to another member of the same population. Similarly, individuals will be
more similar to the average or typical phenotype of another population than to the average phenotype in their
own population with a probability of 14% (CC = 0.14). It follows that variation in such a trait will often be
discordant with population labels.

The population groups in this example are quite distinct from one another: Europeans, sub-Saharan Africans, and
East Asians. Many factors will further weaken the correlation between an individual's phenotype and their
geographic ancestry. These include considering more closely related or admixed populations, studying
phenotypes influenced by fewer loci, unevenly distributed effects across loci, nonadditive effects, developmental
and environmental effects, and uncertainties about individuals' ancestry and actual populations of origin. The
typical frequencies of alleles that influence a phenotype are also relevant, as our results show that rare
polymorphisms yield high values of
CC, and CT, even when many such polymorphisms are studied. This
implies that complex phenotypes influenced primarily by rare alleles may correspond poorly with population
labels and other population-typical traits (in contrast to some Mendelian diseases). However, the typical
frequencies of alleles responsible for common complex diseases remain unknown. A final complication arises
when racial classifications are used as proxies for geographic ancestry. Although many concepts of race are
correlated with geographic ancestry, the two are not interchangeable, and relying on racial classifications will
reduce predictive power still further.
The fact that, given enough genetic data, individuals can be correctly assigned to their populations of origin is
compatible with the observation that most human genetic variation is found within populations, not between
them. It is also compatible with our finding that, even when the most distinct populations are considered and
hundreds of loci are used, individuals are frequently more similar to members of other populations than to
members of their own population. Thus, caution should be used when using geographic or genetic ancestry to
make inferences about individual phenotypes.

Go to:


Go to:

We thank Jinchuan Xing, Michael Bamshad, Dennis O'Rourke, and Thomas Doak for thoughtful comments. This
work was supported by National Science Foundation grants BCS-0218338 (M.A.B.), BCS-0218370 (L.B.J.), and
EPS-0346411 (M.A.B.); by National Institutes of Health grant GM-59290 (L.B.J. and M.A.B.); by the Louisiana
Board of Regents Millennium Trust Health Excellence Fund HEF (2000-05)-05 (M.A.B.), (2000-05)-01
(M.A.B.), and (2001-06)-02 (M.A.B.); and by the Intramural Research Program of the National Institute of
Diabetes and Digestive and Kidney Diseases, National Institutes of Health.

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American Anthropological Association, 1997. Response to OMB directive 15: race and ethnic standards for
federal statistics and administrative reporting (original statement at
/web/19990507115624/; amended 2000 statement at
Bamshad, M., S. Wooding, B. A. Salisbury and J. C. Stephens, 2004. Deconstructing the relationship between
genetics and race. Nat. Rev. Genet. 5: 598609. [PubMed]
Bamshad, M. J., S. Wooding, W. S. Watkins, C. T. Ostler, M. A. Batzer et al., 2003. Human population genetic
structure and inference of group membership. Am. J. Hum. Genet. 72: 578589. [PMC free article] [PubMed]
Barbujani, G., A. Magagni, E. Minch and L. L. Cavalli-Sforza, 1997. An apportionment of human DNA diversity.
Proc. Natl. Acad. Sci. USA 94: 45164519. [PMC free article] [PubMed]
Bowcock, A. M., A. Ruiz-Linares, J. Tomfohrde, E. Minch, J. R. Kidd et al., 1994. High resolution of human
evolutionary trees with polymorphic microsatellites. Nature 368: 455457. [PubMed]
Campbell, D., P. Duchesne and L. Bernatchez, 2003. AFLP utility for population assignment studies: analytical
investigation and empirical comparison with microsatellites. Mol. Ecol. 12: 19791991. [PubMed]
Chakraborty, R., and L. Jin, 1993. A unified approach to study hypervariable polymorphisms: statistical
considerations of determining relatedness and population distances. Exper. Suppl. 67: 153175. [PubMed]
Cornuet, J. M., S. Piry, G. Luikart, A. Estoup and M. Solignac, 1999. New methods employing multilocus
genotypes to select or exclude populations as origins of individuals. Genetics 153: 19892000.
[PMC free article] [PubMed]
Dean, M., J. C. Stephens, C. Winkler, D. A. Lomb, M. Ramsburg et al., 1994. Polymorphic admixture typing in
human ethnic populations. Am. J. Hum. Genet. 55: 788808. [PMC free article] [PubMed]
Edwards, A. W., 2003. Human genetic diversity: Lewontin's fallacy. BioEssays 25: 798801. [PubMed]
Human Genome Project, 2001. The human genome. Nature 409: 812.
International HapMap Consortium, 2005. A haplotype map of the human genome. Nature 437: 12991320.
[PMC free article] [PubMed]
Jorde, L. B., W. S. Watkins, M. J. Bamshad, M. E. Dixon, C. E. Ricker et al., 2000. The distribution of human
genetic diversity: a comparison of mitochondrial, autosomal, and Y-chromosome data. Am. J. Hum. Genet.
66: 979988. [PMC free article] [PubMed]
Lao, O., K. van Duijn, P. Kersbergen, P. de Knijff and M. Kayser, 2006. Proportioning whole-genome singlenucleotide-polymorphism diversity for the identification of geographic population structure and genetic
ancestry. Am. J. Hum. Genet. 78: 680690. [PMC free article] [PubMed]
Latter, B., 1980. Genetic differences within and between populations of the major human subgroups. Am. Nat.
116: 220237.
Lewontin, R. C., 1972. The apportionment of human diversity. Evol. Biol. 6: 381398.
Lohmueller, K. E., M. M. Mauney, D. Reich and J. M. Braverman, 2006. Variants associated with common
disease are not unusually differentiated in frequency across populations. Am. J. Hum. Genet. 78: 130136.
[PMC free article] [PubMed]
Manel, S., P. Berthier and G. Luikart, 2002. Detecting wildlife poaching: identifying the origin of individuals with
Bayesian assignment tests and multilocus genotypes. Conserv. Biol. 16: 650659.
Mitton, J. B., 1977. Genetic differentiation of races of man as judged by single-locus and multilocus analyses.
Am. Nat. 111: 203212.
Mitton, J. B., 1978. Measurement of differentiation: reply to Lewontin, Powell, and Taylor. Am. Nat. 112:
Mountain, J. L., and L. L. Cavalli-Sforza, 1997. Multilocus genotypes, a tree of individuals, and human
evolutionary history. Am. J. Hum. Genet. 61: 705718. [PMC free article] [PubMed]
Mountain, J. L., and U. Ramakrishnan, 2005. Impact of human population history on distributions of
individual-level genetic distance. Hum. Genomics 2: 419. [PMC free article] [PubMed]
Nei, M., 1973. Analysis of gene diversity in subdivided populations. Proc. Natl. Acad. Sci. USA 70: 33213323.
5/14/2016 11:46 PM

Genetic Similarities Within and Between Human Populations

10 of 10

[PMC free article] [PubMed]

Nei, M., and A. K. Roychoudhury, 1972. Gene differences between Caucasian, Negro, and Japanese populations.
Science 177: 434436. [PubMed]
Powell, J. R., and C. E. Taylor, 1978. Are human races substantially different genetically? Am. Nat. 112:
Pritchard, J. K., M. Stephens and P. Donnelly, 2000. Inference of population structure using multilocus genotype
data. Genetics 155: 945959. [PMC free article] [PubMed]
Risch, N., E. Burchard, E. Ziv and H. Tang, 2002. Categorization of humans in biomedical research: genes, race
and disease. Genome Biol. 3: 2007.12007.12. [PMC free article] [PubMed]
Romualdi, C., D. Balding, I. S. Nasidze, G. Risch, M. Robichaux et al., 2002. Patterns of human diversity, within
and among continents, inferred from biallelic DNA polymorphisms. Genome Res. 12: 602612.
[PMC free article] [PubMed]
Rosenberg, N. A., J. K. Pritchard, J. L. Weber, H. M. Cann, K. K. Kidd et al., 2002. Genetic structure of human
populations. Science 298: 23812385. [PubMed]
Rosenberg, N. A., S. Mahajan, S. Ramachandran, C. Zhao, J. K. Pritchard et al., 2005. Clines, clusters, and the
effect of study design on the inference of human population structure. PLoS Genet. 1: e70. [PMC free article]
Serre, D., and S. Pbo, 2004. Evidence for gradients of human genetic diversity within and among continents.
Genome Res. 14: 16791685. [PMC free article] [PubMed]
Shriver, M. D., R. Mei, E. J. Parra, V. Sonpar, I. Halder et al., 2005. Large-scale SNP analysis reveals clustered
and continuous patterns of human genetic variation. Hum. Genomics 2: 8189. [PMC free article] [PubMed]
Smouse, P. E., and R. Chakraborty, 1986. The use of restriction fragment length polymorphisms in paternity
analysis. Am. J. Hum. Genet. 38: 918939. [PMC free article] [PubMed]
Smouse, P. E., R. S. Spielman and M. H. Park, 1982. Multiple-locus allocation of individuals to groups as a
function of the genetic variation within and differences among human populations. Am. Nat. 119: 445.
Stephens, J. C., J. A. Schneider, D. A. Tanguay, J. Choi, T. Acharya et al., 2001. Haplotype variation and linkage
disequilibrium in 313 human genes. Science 293: 489493. [PubMed]
Tang, H., T. Quertermous, B. Rodriguez, S. L. Kardia, X. Zhu et al., 2005. Genetic structure, self-identified
race/ethnicity, and confounding in case-control association studies. Am. J. Hum. Genet. 76: 268275.
[PMC free article] [PubMed]
Turakulov, R., and S. Easteal, 2003. Number of SNPs loci needed to detect population structure. Hum. Hered. 55:
3745. [PubMed]
Watkins, W. S., A. R. Rogers, C. T. Ostler, S. Wooding, M. J. Bamshad et al., 2003. Genetic variation among
world populations: inferences from 100 Alu insertion polymorphisms. Genome Res. 13: 16071618.
[PMC free article] [PubMed]
Watkins, W. S., B. V. Prasad, J. M. Naidu, B. B. Rao, B. A. Bhanu et al., 2005. Diversity and divergence among
the tribal populations of India. Ann. Hum. Genet. 69: 680692. [PubMed]
Witherspoon, D. J., E. E. Marchani, W. S. Watkins, C. T. Ostler, S. P. Wooding et al., 2006. Human population
genetic structure and diversity inferred from polymorphic L1(LINE-1) and Alu insertions. Hum. Hered. 62:
3046. [PubMed]
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