A defense of the race concept

Below is a defense of the biological race concept.

1 The core element of “Race” is ancestry
2 The ordinary concept of race means “people related by region, ancestry, and appearance”
3. The ordinary concept of race is logically coherent
4. The scientific concept of race is subspecies
5. The ordinary concept of race has biological reality
6. The biological basis of race is independent from the existence of subspecies
7. There are scientific natural kinds that roughly correspond to the ordinary concept of race
8. The issue of socially significant heritable differences is independent from that of scientific natural kinds and subspecies
9. The genetic diversity between populations is sufficient for substantial between population heritable differences
10. The humans species is polytypic, given an often used subspecies concept

1. Race means ancestry

It’s often asserted that the term “race” has many meanings and concluded that “race” therefore is “a social construct, with no biological meaning.” This argument is an obvious Non Sequitur. Nonetheless, it’s worth pointing out that race is a polyseme with a core element of “ancestry.”

While there have been many concepts of race (taxonomic lineage, continental ancestry, breeding population, genealogy, nation, Ur myth, etc,), the concepts are related in that they deal with populations that are genetically (whether in the classical Greek sense of “origin“ — genetikos — or the modern sense of DNA) related. This should not be surprising as etymological analysis shows that the root of the term “race” is “origin.”

2. The ordinary concept of race

The ordinary concept of race (sometimes called the “folk concept”) is the concept that ordinary people mean when they use the term race.

The ordinary concept of race (OCR) is “people related by region, ancestry, and appearance” [1]. Sometimes it’s contended that the ordinary concept of race refers to “people of similar appearances” [1]; it’s difficult, however, to reconcile this interpretation with the historic practice of distinguishing people by racial admixture (e.g. the one drop rule) or the practice of identifying race with region (e.g. Asian = Asian, Black = African American). For example, by the US government’s definition [ 2], race, in part, refers to the regional ancestry of individuals:

In Directive No. 15, the “White” category includes persons having origins in any of the original peoples of Europe, North Africa, or the Middle East. The public comment included suggestions for subcategories and related changes in terminology to collect more detailed information on White ethnic groups according to the geographic region of their ancestors.”

[1] Hardimon, 2009. Wallis Simpson was Wrong
[2] For a discussion of this refer to: Hardimon, 2003. The ordinary concept of race
[3] Office of Management and Budget. Standards for the Classification of Federal Data on Race and Ethnicity. http://www.whitehouse.gov/omb/fedreg_race-ethnicity

3. The ordinary concept of race is logically coherent

It’s often argued that the ordinary concept of race is logically incoherent; it clearly isn’t. With regards to the conceptual coherence of race, it should be first noted that “race” (R1, R2, R3..) is a set of classes (i.e. a system of classification). Whether classifications really exist is an interesting question, but not one germane to the topic. By semantic theory for a system of classification to be coherent, there must be a criterion for distinguishing classes R1/R2/…, the classes must differ in some way in addition to the criterion used to distinguish them, and some of the classes must contain more that 0 members.

With regards to the ORC, the criterion is simple: continental regional ancestry, where ancestry means: the region where most of your ancestors lived X to Y years ago. ** (e.g., Black/Negro/ =African-American). To clarify, the criterion is not “different essences,” “unique genes,” “unique alleles,” or “discrete groups.” These latter are criterion used to set up and knock down straw man definitions of the ordinary concept of “race.” When Cavalli-Sforza, for example, claims that there are no human populations marked by unique genes and concludes that there are no races (or more properly, that no human racial classifications have more than 0 members), he obviously is not talking about the OCR.

Now, not everyone fits into a race classification delineated accordingly — when so, they are called “multiracial.” As for the requirement of additional differences, they would be, at very least, patterns of skin color, hair color and form, and facial traits. It could be objected that 0 members belong to the class R1/R2.. and share a common additional difference. This objection fails since it’s sufficient that there are patters of differences between R1/R2/R3.

To clarify, the criterion is regional ancestry; the additional differences are phenotypic patterns. Now it’s trivially true that races, as we ordinarily qualify “region”/“most”/and “ancestry” in this context, have some members; as such, we can conclude that the ordinary concept of race is coherent.

* This is an adaptation of the argument made by Levin (2002).
**”Region” and “Most” is open to qualification — as is the time frame. The point of this exercise is to just establish the conceptual coherence of the ordinary concept of “race” as we use the term; others define race in terms of region. See for example, Risch et al. (2002) “Categorization of humans in biomedical research: genes, race and disease.” Quote: “For the purpose of this article, we define racial group on the basis of primary continent of origin.”

4. The dominant scientific concept

The dominant scientific concept of race (DSCR) is “subspecies.” There is no consensus on the definition of subspecies and the criteria for subspecies classifications. As it is, there is no consensus on the meaning of species [1, 2]. The definition used in the US for the purpose conservation is [3]:

The subspecies category has been defined as “a geographically defined aggregate of local populations which differ taxonomically from other subdivisions of the species.” A valuable recent modification urged that the evidence for BCS subspecies designation should come from the concordant distribution of multiple, independent, genetically based traits. In an attempt to provide formal criteria for subspecies classifications we offer the following guidelines: Members of a subspecies share a unique geographical range or habitat, a group of phylogenetically concordant phenotypic characters, and a unique natural history relative to other subdivisions of the species. Because they are below the species level, different subspecies are reproductively compatible, They will normally be allotropic and they will exhibit recognizable phylogentic partitioning, because of time dependent accumulations of genetic difference in absence of gene flow. Most subspecies will be monophyletic, however they will also derive from ancestral subspecies hybridization. ….Occasional introgression or inbreeding should not be viewed as inconsistent with subspecies status, they simply change the phylogenetic description. (O’Brien and Mayr, 1991. Bureaucratic Mischief: Recognizing Endangered Species and Subspecies.)

[1] Queiroz, 2007. Species concepts and species delimitation
[2] Wilkins, 2002. Summary of 26 species concept
[3] O’Brien and Mayr, 1991. Bureaucratic Mischief: Recognizing Endangered Species and Subspecies

5. The biological reality of the ordinary concept of race

The ordinary concept of race is biologically real. In the philosophy of science, to say that races are biologically real, to use the common parlance, is to say:

a) There is a logically coherent definition of race, such that race X can be distinguished from race Y
b) Members of race X (or Y) can be distinguished from members of race Y (or X) and can be accurately identified as a member of race X (or Y)
c) Race X differs from race Y in patterns of practically significant heritable characteristics

There is a logically coherent definition of race (above). Members of a given race are genetically more similar due to common ancestry [1] and so can be distinguished from members of other races; individuals can be correctly classified with a high degree of probability based both on phenotype [2, 4, 5] and geneotype [6]. Races differ in patterns of heritable phenotypic characteristics [3], some of which have practical significance [7]. (Some would argue that for human populations to be “biologically real” every members would have to differ by some common observable characteristic, yet even the common subspecies criteria doesn’t require this. Classically, populations qualified as subspecies if 75% of the members could be correctly assigned to the population [8, 9].)

[1] Witherspoon et al., 2007. Genetic similarities within and between human population.

A good measure of the robustness of racial genetic differentiation is the answer to the following question: ‘‘How often does it happen that a pair of individuals from one population is genetically more dissimilar than two individuals chosen from two different populations?’’ In fact, if many thousands of loci are used as a basis for judging genetic similarity and when individuals are sampled from geographically separated populations, the correct answer, which many will probably find surprising, is: ‘‘Never.’’

[2] Brues, 1990. The once and future diagnosis of race
[3] Gill. Craniofacial criteria in the skeletal attribution of race
[4] Konigsberg, et al. 2009. Estimation and evidence in forensic anthropology: sex and race
[5] Ousely, et al., 2009. Understanding Race and Human Variation: Why Forensic
Anthropologists are Good at Identifying Race

Further, our results show that humans can be accurately classified into geographic origin using craniometrics even though there is overlap among groups

[6] Mountain and Risch, 2004. Assessing genetic contributions to phenotypic differences among ‘racial’ and ‘ethnic’ groups
[7] Burchard et al., 2003. The Importance of Race and Ethnic Background in Biomedical Research and Clinical Practice
[8] Amadon, D. 1949. The seventy-five percent rule for subspecies
[9] Smith et al., 1997. Subspecies and Classification

6. The biological basis of race is independent from the existence of subspecies

It is often argued that races are not biological since there are no human subspecies.

P1 zoologically, race = subspecies
P2 what we call races aren’t subspecies
P3 therefore, races don’t exists as subspecies
P4 therefore races are not scientific natural kinds
P4 therefore races are “social constructs”
P5 social constructs have no biological content
Conclusion: therefore races have no biological content

This is a Non Sequitur. 1) Races can have biological reality without being scientific natural kinds. In such a case, the boundaries of the concept are defined socially (e.g. hail from continental Europe) while the content is biological (e.g. are ancestrally Caucasian). Moreover, 2) Races can refer to scientific natural kinds (e.g. clades) without referring to subspecies.

7. There are scientific natural kinds that roughly correspond to the ordinary concept of race

There are several major human ancestral population lineages [1] or clades (SS. African (Black), West + Central Eurasian (White), (South Asians), East Eurasian, Oceanian, Amerindian). A human clade is defined as a reasonably reproductively isolated population related ancestrally-descendantly [2, 3]. These clades overlap with the [8] several major phylogenetically delineated population clusters [4] which resulted from historic human divergence. The cladistically and phylogenetically (ancestrally and genetically) defined populations complement each other [5]. These are scientific natural kinds which roughly correspond with the OCR.

[1] Wilson and Cann, 1992. The recent African genesis of humans
[2] Andreasen, 2003. The Cladistic Race Concept: A Defense
[3] Andreasen, 2005. The Meaning of ‘Race’: Folk Conceptions and the New
[4] See, for example: McEvoy, et al., 2010. Whole-Genome Genetic Diversity in a Sample of Australians with Deep Aboriginal Ancestry
[5] See for example: Zhang, 2008. Tree-guided Bayesian inference of population structures

As shown in Figure 4, TIPS identified 23 population clusters from this dataset. The six main population clusters of the globe were detected by our method and they formed distinct clades in the estimated tree. We also identified many subpopulations within each of the six major population clusters, and they corresponded well to their known origins, such as Yakut and Mongola in East Asia, Kalash in Central South Asia. We observed near perfect separation of populations in Africa, Oceania and America, but only weak separation in Europe and Middle East. In addition, the reconstructed population tree appeared to be roughly consistent with the geographic distributions of the known populations and their migration histories. For example, the African populations formed the outmost clade in the tree in support of the out of Africa model (Stringer and McKie, 1996). Overall, our result was concordant with the previous analysis.

8. The issue of socially significant heritable differences is independent from that of scientific natural kinds and subspecies

Race has a biological basis (ancestry). The substantiate issue concerns the boundaries of the concept. As such, much of the debate about the “reality of race” revolves around the ability to distinguish this race (ancestral population) form that. But this is an artificial debate. One can define the boundaries in non-biological terms (e.g religion, culture, geography), giving the concepts of relgiorace, ethnorace, and regional race. Assuming some coherent boundary, those races will qualify as being “biologically real.”

9. The genetic diversity between populations is sufficient for substantial between population heritable differences.

Genetic variability is often measured by fixation index (Fst). Fst isn’t a particularly good measure of genetic-phenotypic mediation. As a case example, Long and Kittles (2003) found a between human population Fst of 11% based on their sample; when they added chimpanzees, the between population Fst increased only to 18% [1]. Mountain and Risch (2004), citing this example, note that ‘‘a low FST estimate implies little about the degree to which genes contribute to between-group differences.” [2]

Regardless, based on 650,000 single-nucleotide polymorphisms, Li et al., (2008) found a between-region variance of ~10% [3]; when averaged across studies and indexes (e.g RFLP loci, enzymes, SNPs), 10% is the median between Continental race variance found [4, 5]. Sewall Wright, who helped develop the index, noted that this signified moderate genetic diversity.

“0 to 0.05 indicates little genetic differentiation.
“0.05 to 0.15 indicates moderate genetic differentiation.
0.15 to 0.25 indicates great genetic differentiation.
0.25 indicate very great genetic differentiation.
(Wright, S. 1978. Evolution and the genetics of populations)”

It’s not unreasonable to suppose that a moderate amount of phenotypic diversity is mediated by genetics. If we naively translate the Fst and other variance estimates into phenotypic differences we do in fact see that there would be a substantial amount of differences.

If we have a 90% within race variance, we have the equivalent of a 45% between individual, within race variance. (Roughly half of the within group variance is within individuals). This gives of a 22% between individual, between race variance (10/45). http://occidentalascent.wordpress.com/2011/04/14/did-sarich-get-it-right/ Imagine some trait for which there is a 1:1 phenotypic/geneotypic relation and for which the within population SDs are 100, assuming equally numerous populations, if the genes for that trait were randomly dispersed throughout the genetic variance, a 22% between group variance would be equivalent to 1.1 SD difference.

[1] Li, et al., 2008. Worldwide human relationships inferred from genome-wide patterns of variation
[2] Long and Kittles, 2003. Human Genetic Diversity and the Nonexistence of Biological Races
[3] Mountain and Risch, 2004. Assessing genetic contributions to phenotypic differences among ‘racial’and ‘ethnic’groups
[4] Campbell and Trishkoff, 2008. AFRICAN GENETIC DIVERSITY: Implications for Human Demographic History, Modern Human Origins, and Complex Disease Mapping (Estimate 10-16%)
[5] Holsinger and Weir, 2009. Genetics in geographically structured populations: defining, estimating and interpreting FST (Estimate 5-10% between populations)

10. The subspecies question

First, what is the question? To rephrase it: Were we not to conceptually gerrymander, would human population clusters be divided into subspecies, given how other vertebrates are classified? This last point is essential if the use or disuse of the word race in humanity is to have any taxonomic validity.

Following . O’Brien and Mayr’s (1991) criteria for human races to be subspecies, the members

1] share a unique geographical range or habitat
2] share a group of phylogenetically concordant phenotypic characters
3] share a unique natural history relative to other subdivisions of the species.

When it comes to geographic ancestral populations, criteria 3 is clearly fulfilled. The continental populations are the result of historic migrations and adaptations [1, 2]. As is 1, given how the criterion is used in practice (i.e. a member of a subspecies does not lose its status due to migrating or being sent around the world to a zoo). For example, the subspecies of the American Puma have an expansive range [2], and American Puma that are relocated to a Zoo in Asia do not lose there subspecies membership. The ambiguity results from 2. What does it mean for members to “share a group of phylogenetically concordant phenotypic characters”? Does this mean that members must share in a gene pool which manifests heritable characteristics (i.e. share patterns of traits resulting from phylogenetic history)? Or does this mean that members must have a set of characteristic traits. If the former, is the issue quantitative (frequency) or qualitative (uniqueness)? Since there are patterns of differences and individuals can be correctly assigned to racial groups with a high degree of accuracy [6], it’s evident that members of races do share common distinguishable phenotypic patterns in the sense of frequency [4, 5]. And some of these traits are more or less uniquely characteristic (e.g. EDAR codes for thick hair in most East Asians; SLC24A5 codes for skin color in most Caucasians [7, 8]).

But how do we decide whether we mean patterns or sets. Since our question was “….given how other vertebrates are classified?” we have to refer to how other subspecies are classified. Based on the usages [9, 10], if we were conceptually consistent we would probably conclude that the major population clusters/clades do represent subspecies. This is worth clarifying:

a) There are (some) genetic clusters/clades.
b) Members share (or did share ) a unique geographical range or habitat
c) Members share a unique natural history relative to other subdivisions of the species.
d) Members share patterns of heritable phenotypic differences (frequency).

If humans were turtles, the subspecies label would be used in the classification of populations. As such, Ernst Mayr, who co-developed this subspecies concept stated:

Let me begin with race. There is a widespread feeling that the word “race” indicates something undesirable and that it should be left out of all discussions. This leads to such statements as “there are no human races.” Those who subscribe to this opinion are obviously ignorant of modern biology….

…No matter what the cause of the racial difference might be, the fact that species of organisms may have geographic races has been demonstrated so frequently that it can no longer be denied. And the geographic races of the human races – established before the voyages of European discovery and subsequent rise of a global economy – agree in most characteristics with the geographic races of animals. Recognizing races is only recognizing a biological fact. (Mayr, 2002. The biology of race and the concept of equality

[1] Forster, 2004. Ice Ages and the mitochondrial DNA chronology of human dispersals: a review
[2] Burchard et al., 2003. The

Importance of Race and Ethnic Background in Biomedical Research and Clinical Practice
From the perspective of genetics, structure in the human population is determined by patterns of mating and reproduction. Historically, the greatest force influencing genetic differentiation among humans has been geography. Great physical distances and geographic barriers (e.g., high mountains, large deserts, and large bodies of water) have imposed impediments to human communication and interaction and have led to geographically determined endogamous (i.e., within-group) mating patterns resulting in a genetic substructure that largely follows geographic lines. The past two decades of research in population genetics has also shown that the greatest genetic differentiation in the human population occurs between continentally separated groups.

[3] Culver, et al., 2000. Genomic Ancestry of the American Puma (Puma concolor) Figure 1.
[4] Relethford, 2009. Race and Global Patterns of Phenotypic Variation
[5] Gill. Craniofacial criteria in the skeletal attribution of race
[6] Ousely, et al., 2009. Understanding Race and Human Variation: Why Forensic
Anthropologists are Good at Identifying Race
[7] Fujimoto, 2008. A scan for genetic determinants of human hair morphology: EDAR is associated with Asian hair thickness
[8] Sabeti, 2007. Genome-wide detection and characterization of positive selection in human populations
[9] Bickham, 2007. Turtle Taxonomy: Methodology, Recommendations, and Guideline

We propose that subspecies classification, if used, should describe the major patterns of variation found within a species. A precise definition of “major” is elusive, but the formal subspecific description of small, isolated populations, particularly in low vagility species, should be avoided unless there is strong reason to do otherwise. This could avoid the proliferation of named forms of small, isolated populations such as occurred with pocket gophers in western North America (Smith and Patton, 1988).

[10] Phillips, 2009. Systematics of Steller sea lions (Eumetopias jubatus): subspecies recognition based on concordance of genetics and morphometrics

For males, three variables were used in stock assignment to correctly identify 88.13% and 86.59% of individuals from the eastern and western stocks, respectively. Through the same method the correct identification in stock assignment using five selected variables for female eastern and western stock individuals was 86.27% and 88.1%, respectively

………….
Common Critiques and responses:

1) Nested African differences — Africans don’t represent one phylogenetically defined race; given the cladistic relation, since Africans aren’t a race, non-Africans don’t represent cladistic races. (Hunely et al., 2009. “The Global Pattern of Gene Identity Variation Reveals a History of Long-Range Migrations, Bottlenecks, and Local Mate Exchange: Implications for Biological Race)

We are either using a cladistic criteria or a phylogenetic one. If we are using the later, the nestedness — which only applies to African populations — suggests that there are multiple African races defined in terms of relative genetic difference. If we are using the former, the genetic dissimilarity between Africans is irrelevant; there would be multiple African cladistic races; since Africans represent cladistic races, their non-race status can not be ground for dismissing the race status of other populations.

2) Clinal not discrete phenotypic differences. (Relethford, 2009. Race and Global Patterns of Phenotypic Variation)

If populations can genotypically discriminated then it doesn’t matter if the phenotypic differences are continuous. Moreover some phenotypic differences are not continuous. Asian skin tone and hair texture aren’t.

3) Not enough genetic differences. The Fst is too low. (Barbujani and Colonn, 2010. Human genome diversity: frequently asked questions)

It’s not infrequently argued that human subspecies do not exist because the between population Fst ~.10 is deemed to be too low for subspecies classifications (even though many other subspecies have lower values). For example, Graves (2010) states:

In 1978, Sewall Wright published a four volume treatise entitled: The evolution and genetics of populations. Volume four is devoted to variability within and among populations…. Chapters 9 & 10 of this volume focus on variability within human populations and what he describes as racial differentiation in mankind. …However, on careful examination we see that Wright based on this own criteria for the existence of race, contradicted himself. The mean Fst did not exceed, nor did it come close to his pre-established value for the existence of subspecies, which he equated with geographical race, Fst>0.25 (Graves, 2010. Biological V. Social Definitions of Race: Implications for Modern Biomedical Research)

Here’s what Wright actually said:

There is no question that all mankind constitutes a single species in view of the absence of any physiological bar to hybridization between the most diverse races or of any recognizable loss of vigor in the first or later generations.

There is also no question, however, that populations that have long inhabited separated parts of the world should, in general, be considered to be of different subspecies by the usual criterion that most individuals of such populations can be allocated correctly by inspection. It does not require a trained anthropologist to classify an array of Englishmen, West Africans, and Chinese with 100% accuracy by features, skin color, and type of hair in spite of so much variability within each of these groups that every individual can easily be distinguished from every other.

It is, however, customary to use the term race rather than subspecies for the major subdivisions of the human species as well as for minor ones. The occurrence of a few conspicuous differences, probably due to selection for adaptation to widely different environmental conditions, does not necessarily imply much difference in general. Nei and Roychoudhury (1974) have shown that the differences among negroids, caucasoids, and mongoloids in the protein and blood group loci are slight compared with those between individuals within any one of them. There is disagreement on the number of major races that should be recognized. At a minimum, the Australoids are added to the three referred to above.
….

Diversification is much greater on the average among than within the major races (F (DS) = 0.0715, F (ST) = 0.1248) but there is more uniformity within ).0.042 to 0.141) than among (0.026 to 0.402). The great differences among loci indicates that more than pure sampling drift is involved. (Wright, 1978. Evolution and the Genetics of Populations, Vol. 4: Variability Within and Among Natural Populations. Univ. Chicago Press, Chicago, Illinois. p 439-440)

It doesn’t take “careful examination” to see that Wright did not define subspecies in terms of having a Fst above >.25. He defined subspecies in the same manner Smith et al 1997 do below.

Templeton (1998) states:

A standard criterion for a subspecies or race in the nonhuman literature under the traditional definition of a subspecies as a geographically circumscribed, sharply differentiated population is to have F* values of at least 0.25 to 0.30 (Smith et al. 1997). Hence, as judged by the criterion in die nonhuman literature, the human Fn value is too small to have taxonomic significance under the traditional subspecies definition…Although human “races” do not satisfy the standard quantitative criterion for being traditional subspecies (Smith etal. 1997)…(Templeton, 1998. Human races: a genetic and evolutionary perspective

Here’s what Smith et al actually said:

“The non-discrete nature of subspecies is evident from their definition as geographic segments of any given gonochoristic (bisexually reproducing) species differing from each other to a reasonably practical degree (e.g., at least 70-75%), but to less than totality. All subspecies are allopatric (either dichopatric [with non-contiguous ranges] or parapatric [with contiguous ranges], except for cases of circular overlap with sympatry); sympatry is conclusive evidence (except for cases of circular overlap) of allospecificity (separate specific status). Parapatric subspecies interbreed and exhibit intergradation in contact zones, but such taxa maintain the required level of distinction in one or more characters outside of those zones. Dichopatric populations are regarded as subspecies if they fail to exhibit full differentiation (i.e., exhibit overlap in variation of their differentiae up to 25-30%), even in the absence of contact (overlap exceeding 25-30% does not qualify for taxonomic recognition of either dichopatric populations or of parapatric populations ….

…..The use of multivariate statistical procedures can provide approaches that are reasonably objective and not dependent on preconceptions about taxonomic membership. Nonetheless, the discriminatory power of such methods depends critically on the quality of the characters being analyzed and, in addition, the adopted standard for level of differentiation required for taxonomic recognition. Multivariate analyses (Thorpe 1987) are useful techniques for substantiation of subspecific validity, with revival of the now generally neglected 75% (or similar) rule (idem:7) (Smith et al., 1997. Subspecies and Classification)

Not only do they not discuss Fst, but based on their criteria, human populations clearly qualify as subspecies. The 70-75% they refer to is the percent of correct assignments of individuals to a population (see: Amadon, 1949) . With humans, using multivariate analyses, individuals can be assigned to populations with well over 90% accuracy (Konigsberg, 2009). There is no Fst criteria for subspecies.

Lao and Kayser (2009) state:

So far, studies performed at various loci have shown that the proportion of genetic variation obtained when individuals were clustered according to their geographic continent of origin is quite small (ranging from only 5% up to 15%) compared to that seen when all humans were considered as a single group (approximately 80%) (Romualdi et al., 2002). For comparison: a biological criterion (despite subjective) to define the presence of subspecies is finding estimations of genetic dierentiation greater than approximately 25%( Kittles and Weiss, 2003). (Lao and Kayser, 2009. Human Relationships Inferred from Genetic Variation)

Kittles and Weiss (2003) is cited. They state:

“The traditional, though subjective, criterion for biological subspecies is FST > 0.25 (168, 190). The percentage of genetic diversity between spot-samples from the extremes of the Old World range of human populations (sub-Saharan Africa, East Asia, and Europe) is about 5–15%, with the remaining 85–95% within populations as noted earlier (Table 1) (2, 5, 60, 75, 76). From available sequence data, FST would likely be three times higher among different chimpanzee populations compared to levels for different human populations on separate continents”

Who do they cite? Tempelton and, of all people, Sewall Wright!

References not cited above

Konigsberg, 2009. Estimation and evidence in forensic anthropology: sex and race
Amadon, D. 1949. The seventy-five percent rule for subspecies