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Culture rather than genes provides greater scope for the evolution of large-scale human prosociality

Adrian V. Bella,1, Peter J. Richersonb, and Richard McElreathc

Group in Ecology, bDepartment of Environmental Science and Policy, and cDepartment of Anthropology, University of California, Davis, CA 95616

Edited by Richard E. Nisbett, University of Michigan, Ann Arbor, MI, and approved August 31, 2009 (received for review March 25, 2009)

Whether competition among large groups played an important role in human social evolution is dependent on how variation, whether cultural or genetic, is maintained between groups. Comparisons between genetic and cultural differentiation between neighboring groups show how natural selection on large groups is more plausible on cultural rather than genetic variation.
altruism ͉ cultural FST ͉ group selection ͉ prosociality

uman societies are unusual among vertebrates. While people in small-scale societies exhibit much more cooperation and division of labor than other primates, people in even very large societies also show strong tendencies toward altruism. Warfare, food sharing, and taxation are all examples of prosocial patterns of behavior that are common in human societies but nearly completely absent in other vertebrates. Even when plausible analogues can be found in other vertebrates, the scale of costliness of human altruism is extraordinary (1). Explaining the levels of human altruism observed ethnographically and experimentally has proven to be difficult. Much of this altruism is directed at strangers, and so is difficult to explain as simple reciprocity, or it benefits entire tribes or nations of only distant genealogical kin, and so is difficult to explain as altruism among individuals sharing recent common ancestry. Another scenario many researchers, since at least Darwin (2), are concerned with is competition among residential human groups that are too large to comprise close genealogical kin (2–6). If groups differ in the frequency of individuals who are willing to sacrifice their own labor, time, or safety in ways that promote the competitive ability of the residential group, then over time groups with higher frequencies of such ‘‘altruists’’ may tend to replace groups with fewer (7–9). In this paper, we refer to this scenario as ‘‘group-level selection,’’ the evolution of behavior that reduces individual fitness but increases the average fitness within large groups of only distantly related individuals. By ‘‘distantly related,’’ we mean that most individuals within the residential group do not share very recent common ancestors, and so common descent alone does not maintain much genetic variation among residential groups. Nevertheless, given the right population structure and low rates of mixing among groups, individuals within groups may be much more genetically similar to one another than they are to members of other groups, and therefore they may be closely ‘‘related’’ in one important sense of the term (10). If genetic variation among groups is sufficiently large, evolutionary theory predicts that self-sacrifice on behalf of large residential groups can evolve under the same processes that evolve self-sacrifice on behalf of close kin. This is because all hypotheses about the evolution of altruistic behavior—behavior that reduces the absolute fitness of the actor but increases the absolute fitness of recipients—hinge on processes that change and maintain variation among social groups (11–14). Selection for altruism in such large groups, however, remains a controversial topic in part because it is not clear that enough between-group variation existed in human societies to make it an appreciable evolutionary force (15). In very large residential groups, migration can quickly erode between-group genetic͞cgi͞doi͞10.1073͞pnas.0903232106


variation. Nevertheless, recent work argues that sufficient variation did exist by invoking reproductive leveling (7) [see also (16)]. Reproductive leveling reduces the amount of between group variation needed for selection to favor group-beneficial but individually-costly traits. While it is not known how the estimates of genetic differentiation for small forager groups reported in (7) relate to Pleistocene foraging groups (see SI), it is intriguing to note that reproductive leveling itself already has strong hints of prosociality, begging the question of how it could evolve before altruism (17). This illustrates that for genetic selection to favor altruism in large residential groups, theorists need to invoke particularly strong assumptions. An alternative scenario is that human propensities to cooperate arose through selection on cultural rather than genetic variation (15, 18). Humans developed the capacity for complex culture perhaps beginning 250,000 years ago (19). Since that time, culturally transmitted traits have come, along side of genes, to have a large influence on human behavior. Ever since the advanced human capacity for social learning began, groups of individuals likely began rapid divergence in behavior due to cumulative cultural changes. This behavioral variation between groups can persist, given the right kinds of cultural evolutionary forces (20). Even among our closest living relatives, chimpanzees, plausibly socially-learned traits show some between group variation (21). Selection for culturally-prescribed altruists occurs through the same process as for genes: groups of altruists leave more daughter societies (8, 9). However, one advantage that cultural variation has over genetic is that it does not require violent inter-group competition, nor group extinctions (22, 23). If failed groups were incorporated routinely into successful ones, conformist transmission and other forms of resocialization of failed groups can lead to effective cultural selection on groups even though such a pattern will generate rates of migration that keep genetic FST very low between neighbors. Thus selection on culture can be powerful precisely when genetic selection at the group level is weakest. What is the scope for group-level selection on cultural variation and how does this compare to the equivalent for genes? A number of mechanisms may permit cultural variation to be larger than genetic variation between groups (15, 20). If these mechanisms are important, the scope for group-level selection on culture will be much greater than for genes. Here we compute estimates of cultural variation among human groups and compare these to previous estimates of genetic variation among groups. We restrict ourselves to neighboring groups in the main analysis since only neighbors could compete directly. Despite good reasons to believe our estimates of cultural variation are
Author contributions: A.V.B., P.J.R., and R.M. designed research; A.V.B. performed research; A.V.B. analyzed data; and A.V.B., P.J.R., and R.M. wrote the paper. The authors declare no conflict of interest. This article is a PNAS Direct Submission.

whom correspondence should be addressed. E-mail:

This article contains supporting information online at 0903232106/DCSupplemental.

PNAS ͉ October 20, 2009 ͉ vol. 106 ͉ no. 42 ͉ 17671–17674



the similarity of neighbors is much greater than non-neighbors. We used the World Values Survey (WVS) (25) as a source of data to compute cultural FST for a fairly large sample of national neighbors. the condition for the frequency of altruism to increase is: ␤ ͑ w g.0660) between populations is more than order of magnitude larger than their corresponding genetic FST (mean ϭ 0. we can assess the relative ability of either inheritance system to respond to group-level selection. 17672 ͉ www. The one exception known to us is analyzed in the SI and Table S2. we require systematic samples of individual beliefs or behavior. we do not think that the available data from living populations is consistent with neighbors having FST values as high as 0. The sample size within countries is also large and thus favorable to calculating precise estimates of within and between group variation. In most parts of the Old World for most of the history of Anatomically Modern Humans most populations made rather Bell et al. It is evident that the scope for group-level selection. group beneficial behaviors should be hundreds of times greater than the individual cost to be favored by selection. 1. Above: Histogram of 150 cultural FST (gray fill) and 59 genetic FST (black border) for neighboring countries calculated from the World Values Survey and in (24). There is no reason. the baseline figure used in (7) (see SI). The greater the genetic differentiation (FST) between two groups. but rather is an exact description of how selection works. From these estimates we can compute the minimum group benefit over individual cost ratio that would favor altruism. we find much greater scope for multilevel selection on human culture than on human genes. and ␤(wig.0903232106 Fig. Comparison of genetic and cultural differentiation. Cultural FST (mean ϭ 0. Highly variable climates and a disproportionate emphasis on big game hunting in the last glacial compared to the Holocene would probably have made populations more mobile and more prone to long distance movements in the last glacial. The main Western Eurasian Upper Paleolithic cultures.1073͞pnas. By comparing FST for genes and culture. 13). To compute cultural FST. An obstacle to computing cultural FST is that social anthropologists have not traditionally sampled individuals explicitly. The full tables for genetic FST are given in (24). the greater the scope for selection at the group level. In Table S1 in the SI. in principle. the greater the cultural differentiation is between groups. Certainly. we list all of the pairwise cultural FST values. whereas for culture. Discussion Our calculations show much greater scope for cultural rather than genetic group-level selection. for use in describing cultural differentiation between populations. p s͒ 1 Ϫ FST Ͼ ␤͑wig. The culturally innovative Southern African Still Bay and Howieson’s Poort Middle Stone Age traditions appear to have been widespread spatially like the Upper Paleolithic but were more restricted in time than the Aurignacian and Gravettian (28). These results call for attempts to produce better estimates relevant to quantitative models of human cultural evolution. pig) is the fitness decrease of the individual acquiring the altruistic allele. It is difficult to know how last Glacial population structures might have been like compared to Holocene hunter-gatherers. The WVS asks a large battery of questions that are likely to be heavily influenced by culture in a large number of countries. the left-hand side of equation (1). while the respective mean and median for cultural traits is 16 and 14 (range 3–75). FST estimates the proportion of the total variation in a trait or set of traits (or alleles) that is accounted for by between-group differences. pig͒ FST [1] Here ␤(wg. View the left hand side of the inequality as the benefit-cost ratio for the addition of another altruist in a population at the scale of the group and individual.076. In the case of both culture and genes. We then compare these corresponding genetic FST estimates from (24). pg) is the increase in the mean fitness of the group with an increase the frequency of altruists. the Price equation is axiomatic—it does not depend upon simplifying assumptions. the Aurignacian and Gravettian. Calculating Cultural and Genetic Variation. although we should acknowledge how this inference may be limited. Bottom: Plot of the cultural against genetic FST for 59 pairs of neighboring countries. median ϭ 0. The higher this number. grouplevel selection can operate under much less stringent conditions. Human populations densities in most times and places in the Pleistocene were apparently very low.0800. For genes.0032). much smaller population sizes would have generated more drift. respectively. 1 shows the distribution of FST estimates for culture and genes and the bottom panel shows how these estimates relate to equation (1) and the scope for selection among groups. Even in Upper Paleolithic Western Eurasia last glacial populations were apparently only on the order of tens of thousands of people and division into markedly distinct ethnic groups was absent (26).0053. respectively. most ethnography consists of statements about normative behavior that is often observed to vary among groups. The right-hand side of this inequality should be computed for two populations that may compete. For genes the mean and median benefit across all paired countries is 437 and 311 (range from 31–2. The formal condition for altruism to arise can be expressed using the Price equation (11. Instead. a measure of genetic differentiation between populations (24). . On the other hand. Results The top panel of͞cgi͞doi͞10. cultural FST is the proportion of the total variance in allele (or trait) frequencies found between groups. as described in equation (1). not to use the same F-statistic. occurred over the whole of Europe and neighboring West Asia without any strongly marked stylistically marked subdivision (27). is much greater for culture than genes.272). because the derivation of the Price equation makes no assumptions about the nature of the underlying variants.underestimates. As in the case of genetic FST. The low and very low genetic FST values that characterize modern national neighbors might not be typical of ancestral Pleistocene populations. Unlike most evolutionary theory. Put in terms of regression coefficients and the statistic FST.pnas. FST. median ϭ 0.

Ideally.g. even than the ethnographic and genetic samples we have from small-scale foraging populations. For example. Human genes affecting social behavior. That is. Finding that there is greater scope for inter-group competition to select for prosociality on culture rather than genes does not mean that genes are unimportant to the story. 42 ͉ 17673 SOCIAL SCIENCES ANTHROPOLOGY . as behavior geneticists often report appreciable (genetic) heritabilities for seemingly cultural traits like political preferences (33). questions in the World Values Survey (WVS) were regarded as ‘‘loci’’ and responses as ‘‘alleles. In any case. Thus selection acts on competing equilibria as far as the traits upon which group-level selection is acting. One set of data from Africa does provide data sufficient to compute cultural FST (see SI and Table S2). probably arose by gene-culture coevolution (42). national scale data offer some interesting insights on neighbor cultural and genetic differences. complete with fictive descent to give them a place in the Nuer social order (36). Detailed choice of questions and pooling of responses for all questions in the WVS is available upon request from the corresponding author. following (24).’’ then the first two and the last two would be combined as one response. data such as ‘‘no answer’’ or ‘‘not asked’’ and the like were not included in the calculation because they were not considered responses. One important function of our calculation is to call attention to the importance and feasibility of quantitatively estimating the important parameters of evolutionary models in human populations. if the possible responses were ‘‘agree. Also. The way the losing side was integrated demographically. similar responses were pooled as one response if the choice of responses could not be considered as cardinal. Social selecBell et al. A conjecture based on ethnographic accounts [e. selfreported state of health). Human social life is typically regulated by rules of conduct that take the form of self-reinforcing games (40). Cultural FST estimates reported here are likely the lower bound for ethnic groups as questionnaires typically underestimate behavioral variation across groups (31. Some of the variation in the WVS questions may be genetic. (34)]. Cattle domestication and the innovation of dairy farming led to selection pressure on genes to produce the enzymes to break down milk sugars beyond weaning (37). while cultural variation can remain quite large. Differences in marriage institutions led to a deeper reckoning of kinship among the Nuer compared to the Dinka. 106 ͉ no. We are frustrated to have to use estimates of FST derived from late Holocene populations to infer what transpired in Middle and Late Paleolithic contexts. Population densities of Mode 3 toolmakers were probably even lower than Upper Paleolithic West Eurasians (29). For a locus with L alleles and pij is the frequency of allele i in population j. The Nuer-Dinka case illustrates this feature of human intergroup competition. Data from small-scale human societies that better resemble the kinds of social systems important in our evolutionary past would be particularly interesting. Nevertheless. and re-socialized culturally. For example. The WVS is not the best dataset to use for this purpose. Similarly.simple Middle Stone Age or Mode 3 tools (28). The discussions by anthropologists of differences between tribal scale societies that presumably most resemble the late Pleistocene conditions under which our propensities for cooperation arose suggest that cultural differences between tribes were roughly similar to those obtaining between ethnic groups in modern societies [e. as if each population was nearly monomorphic for the same haploid allele. Support for gene-culture coevolution for well-studied physiological traits can be logically extended to the puzzle of human prosociality.’’ and ‘‘strongly disagree. The situation is as if cultural FST was approximately 1 as far as the relevant trait is concerned.’’ ‘‘strongly agree. Further. genetic FST is generally much smaller than cultural FST so a mixture of genetic and cultural effects will lead to an underestimate of cultural FST. the Nuer versus Dinka tribes. innate propensities to cooperate might have evolved by gene-culture coevolution rather than by selection among groups for solely genetic variants (38). (See SI for an analysis of four east-African populations). Social selection mechanisms such as exclusion from the marriage market. Few people will behave contrary to the institution and the non-conformists that do exist will not usually be adhering to the institutions of a neighboring group. immigration into simple societies was often accompanied by cultural assimilation [e. and execution would have exerted strong selection against genes tending toward antisocial behavior. the Nuer and Dinka peoples were genetically similar. We restrict ourselves to neighboring countries in the main analysis since only neighbors could compete directly. the evolution of cultural rules mandating cooperation between group members could exert ordinary selection pressures for genotypes that obey cultural rules. The large differences we find between cultural and genetic FST argue that that the evolutionary processes acting on these two systems of inheritance would have to be very different in the late Pleistocene to affect the qualitative conclusion that the scope for cultural group-level selection was greater than for genetic group-level selection. Thus the genetic variation between groups in such contexts may be quite small.. as neighbors would typically have been before long distance mass transport was available. most nations have multiple ethnic groups that live within their boundaries and different nations often have the same or similar subcultures as neighboring nations. we would like to have neighbor cultural FST estimates for small-scale societies as close as possible in structure to those that characterized our Pleistocene ancestors. For discrete traits. exemplifies contest-based selection acting on institutional cultural variation among groups. tion in favor of genes that predisposed individuals toward prosociality are also easy to imagine.g.... The paleoclimatology and paleoanthropology suggest that Paleolithic populations were structured much differently than contemporary populations. We matched these estimates to available estimates of genetic FST from a database of calculated genetic differentiation between nations and ethnic groups (24).g. one inheritance system likely exerted pressure on the other. banishment. denial of the fruits of cooperative activities. cultural FST was computed as follows. the same differences in evolutionary processes that allow culture more easily than genes to maintain between-group differences should have operated in the Middle and Upper Paleolithic as well as in contemporary populations. then as now. Materials and Methods Using data from four phases of the WVS.’’ ‘‘disagree. into Nuer society further reduced the possibility of maintaining genetic variation between the groups. (39)] is that cultural selection among groups is often driven by differences in institutions. such as our docility compared to chimpanzees (41). For these discrete traits. which led to the Nuer raising larger fighting forces and the expansion of the Nuer at the expense of the Dinka. Questions that explored personal idiosyncrasies rather than cultural beliefs were omitted (e. 32). These fights were not genocidal and many defeated Dinka families were incorporated into Nuer tribes. 2009 ͉ vol. One welldescribed case. different tribes often have different languages or dialects that may function to limit communication between them (35). We conclude that the available evidence suggests that direct group-level selection on genes played a smaller role in human evolution than group-level selection on cultural variation. but no genetic data are available for these groups (30). With our early ancestors inheriting both cultural and genetic variants.g. we computed a pair-wise cultural FST for 154 neighboring pairs of countries sampled therein. As with modern societies. As we see above. The Nuer-Dinka difference was institutional.’’ In the WVS. PNAS ͉ October 20. Thus. (36)].

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Denniston C (Plenum Press.1). 4. Our mean genetic distance is more than an order of magnitude smaller than that reported in Bowles’ study (1) where non-neighbors were included in the analysis. it is apparent that most near neighboring groups are closer genetically that the average populations.0028. 10. Albuquerque). Sheremet’eva VA (1980) The genetics of circumpolar populations of Eurasia related to the problem of human adaptation.. For example. Science 314:1569 –1572. eds Crawford MH. Princeton. mention of sex or not. Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Would you rather own good farming land but no cattle. Harpending H. 7. Bell et al. Twenty-one questions from (7) on four east-African tribes is used to compute a pairwise cultural FST (Table S1). the FST for neighboring Siberian Eskimo and Chukchi is 0.1). From the multidimensional scaling analysis in (2).11. to be 1. Comparison Between Cultural and Genetic FST. reproductive leveling. Table 4.g. p 135 Do the [population] prefer sons or daughters. 3. Groups that tend to have high pairwise FST. what must a [population] own? P 342 appendix Whom can a person trust? p 324 appendix Is there one person you can trust beyond all others? p 325 appendix Why do people kill themselves? p 325 appendix (see Table S2) 5. have probably not been neighbors for many millennia (3).025 although the Chukchi-Alaskan-Eskimo figure is 0. (2002) High levels of Y-chromosome differentiation among native Siberian populations and the genetic signature of a boreal hunter-gatherer way of life. such as Lapps in the European Arctic and Chukchi in Eastern Siberia. Samples from arctic foragers and other small societies that presumably most resemble Pleistocene populations are often very small. Rychkov YG. Workman PL (University of New Mexico Press. 0. rather than an inbreeding coefficient which would be a smaller value. Two Wahlund F estimates are given in (5). Cambridge. p122–124) necessitate that the FST between neighboring groups be far smaller than this average level. www.056 and that for more distant North American populations is even higher. and the evolution of human altruism. New York). Responses to values picture 7: Woman watched by man. 6. Edgerton RB (1971) in The Individual in Cultural Adaptation. In The Human Biology of Circumpolar Populations.Supporting Information Bell et al. Jenkins T (1973) Genetic Distance Among Southern African Populations. It is not always clear in source material whether corrections for sample sizes have been applied in calculations of FST.0264!!). The value for Indigenous circumpolar Eurasian populations of 0.0125. The mean FST for neighboring groups in (3. eds Crow JF. ed Milan FA (Cambridge Univ Press. 0. based on different subdivisions of the !Kung specified by the authors. Some of the populations in (2) that are most different genetically appear to have no close neighbors in the sample. in (3. p 252) report that the average genetic FST for Arctic populations. Cavalli-Sforza LL. Those data refer to Y chromosome variation. and why? p 322 appendix What is the most important thing for parents to teach a toddler? p 322 appendix What is the worst thing that can happen to a man? p 322 Do unmarried females have sexual relations and is it right? p 323 appendix What should a man do if three of his cattle suddenly die? p 323 appendix If a man could have anything he wanted. (see Table S1) Cultural FST for four small-scale societies. p 185 Responses to values picture 8: Man and woman together inside a house. 2. The estimate in (6) was not corrected for biases due to sample size. as is reflected in standard errors of the estimates for many pairs of populations in (3. New York). Menozzi P. when distantly related groups come into contact. One of Bowles’ primary sources (2) does report an autocorrelation analysis suggesting that some of the genetic variation in Central Asia and Siberia is clinal. mostly derived from small foraging populations similar to the populations used in (1).0067 and 0. Mitochondrial DNA and Y chromosome variants are used to trace the recent ancestry of populations because high rates of mutation lead to many new population specific alleles on time scales of a few thousand 1 of ؊2 . Harpending H. Karafet TM. p 186 How does a psychotic person behave. Table 4.11. p 188 What is the most important thing for a young woman to know before marriage? p 60 Do wives obey their husbands and is it right for them to do so? p 143 What makes a man a good friend? p 141 How do people feel about a rich man? p 137 Under what circumstances can a young adult man tell a mzee (‘‘old man’’) that he is wrong. In Methods and Theories of Anthropological Genetics. what would he choose? p 323 appendix To be considered wealthy. Bowles S (2006) Group competition. Hum Biol 74:761–789. New Jersey).0264 Ϯ 0. Similarly. or good cattle but no farming land? p 169 Is it better to have many friends or many kinsmen (who are not clansmen)? p 174 Under what circumstances can a younger brother tell an older brother that he is wrong? P 178. migration often lowers genetic distances. Berkeley). et al. Summary data reported in (23. A Study of Four East African Peoples (University of California Press.076 in (4) appears to be a genetic variance. Jenkins T (1974) !Kung population structure. Table 4.041 (but on p252 the summary figure for all of N Asia is given as 0.11. The variograms presented in (3. The source data available to (1) is not sufficient to recompute his FST values for nearest neighbors only. p 381.1073/pnas. In Genetic Distance.1) is 0. but (apparently) not for their higher rates of mutation. The FST estimates in (1) derived from (2) are corrected for the higher rates of drift for Y (1/4 as many copies in a population as autosomal genes) compared to autosomal loci.pnas. Piazza A (1996) in The History and Geography of Human Genes (Princeton Univ Press. see e.0903232106 SI Text Genetic Evidence of Differentiation.

www.0747 0.0534 0.0458 Bell et al.1165 0.00353 – – – – – – – – – 0.1371 0.0597 0.1320 0.0539 0.0828 0.0052 2 of ؊2 .0015 0.1144 0.004 0.0415 0.0019 Cultural FST 0.0064 – 0.1868 0.0842 0.0437 0.0835 0.0543 0.0212 0.0692 0.0849 0.0043 – – 0.0604 0.02 – 0.0678 0.0206 0.0484 0.0428 0.0578 0.0708 0.1036 0.0469 0.0859 0.0547 0.1548 0.0571 0.0315 0.0439 0.1531 0.0725 0.024 0.0395 0.2545 0.1631 0.0015 0.0391 0.0830 0.0459 0.0305 0.0016 – 0.0590 0.pnas.0036 0.1116 0.0251 0.0329 0.0036 0.0323 0.0518 0.0535 0.Table S1.0032 0.0772 0.0651 0.0923 0.0455 0.0016 0.0012 – – – – – – – – – – – – 0.0666 0.0019 0.0509 0.0329 0.0660 0.0012 – – – – – – – – – 0.0710 0. Genetic and cultural FST estimates reported in this study Genetic FST Albania Albania Albania Albania Algeria Argentina Argentina Argentina Armenia Armenia Australia Austria Austria Austria Austria Austria Austria Austria Azerbaijan Azerbaijan Azerbaijan Bangladesh Belarus Belarus Belarus Belarus Belarus Belgium Belgium Belgium Belgium Brazil Brazil Brazil Brazil Bulgaria Bulgaria Bulgaria Bulgaria Bulgaria Canada Chile China China China China China China China Colombia Colombia Croatia Croatia Croatia Croatia Croatia Czech Republic Czech Republic Czech Republic Denmark Denmark Denmark Denmark Denmark Greece Italy Macedonia. Republic Of Romania Serbia And Montenegro Turkey United States Peru India Japan Pakistan Philippines Republic Of Korea Taiwan Province Of China Viet Nam Peru Venezuela Bosnia And Herzegovina Hungary Italy Serbia And Montenegro Slovenia Germany Poland Slovakia Germany Germany West Great Britain Norway Sweden – – – – – – – – 0.1568 0.0966 0.0323 0.0778 0. Republic Of Serbia And Montenegro Morocco Brazil Chile Uruguay Georgia Turkey New Zealand Czech Republic Germany Hungary Italy Slovakia Slovenia Switzerland Armenia Georgia Russian Federation India Latvia Lithuania Poland Russian Federation Ukraine France Germany Great Britain Netherlands Colombia Peru Uruguay Venezuela Greece Macedonia.00174 – 0.0852 0.0355 0.

1148 0.2211 Bell et al.0592 0.0375 0.00341 0.0746 0.0132 0.0082 0.0022 0.00116 0.0099 – – – 0.0456 0.1149 0.0536 0.00112 – – – – 0.0926 0.0654 0.0136 – – – – 0.1240 0.0997 0.0456 0.00321 0.2498 0. Republic Of Turkey Romania Serbia And Montenegro Slovakia Slovenia Ukraine Great Britain Ireland Northern Ireland Kyrgyzstan Pakistan Philippines Iraq Pakistan Saudi Arabia Turkey Jordan Saudi Arabia Turkey Great Britain Northern Ireland Jordan Bosnia And Herzegovina France Malta Slovenia Switzerland Republic Of Korea Russian Federation Taiwan Province Of China Belarus Lithuania Russian Federation Sweden Poland Sweden Belgium France Germany Serbia And Montenegro Italy United States Spain 0.0954 0.0039 0.0477 0.0023 – – 0.0022 – 0.0153 0.0137 – – – – – – – – – – – – – – – Cultural FST 0.0487 0.1059 0.0077 – – – 0.pnas.1297 0.0034 0.0658 0.0264 0.0603 0.00044 0.1240 0.0871 0.0634 3 of ؊2 .0999 0.0661 0.0756 0.0027 0.1573 0.0670 0.00075 0.00168 – 0.1830 0.1111 0.0653 0.0555 0.1144 0.0229 0.0842 0.0695 0.0024 0.0471 0.0395 0.0579 0.0644 0. Republic Of Malta Mexico Morocco Puerto Rico Finland Latvia Russian Federation Sweden Norway Russian Federation Sweden Germany Great Britain Italy Spain Switzerland Russian Federation Turkey Belgium France Great Britain Germany Northern Ireland Italy Macedonia.0015 0.0406 0.0645 0.0670 0.0698 0.1100 0.0407 0.0027 0.0649 0.0614 0.2083 0.Genetic FST Dominican Republic Estonia Estonia Estonia Estonia Finland Finland Finland France France France France France Georgia Georgia Germany West Germany West Germany West Great Britain Great Britain Greece Greece Greece Hungary Hungary Hungary Hungary Hungary Iceland Iceland Iceland India India Indonesia Iran (Islamic Republic Of) Iran (Islamic Republic Of) Iran (Islamic Republic Of) Iran (Islamic Republic Of) Iraq Iraq Iraq Ireland Ireland Israel Italy Italy Italy Italy Italy Japan Japan Japan Latvia Latvia Latvia Latvia Lithuania Lithuania Luxembourg Luxembourg Luxembourg Macedonia.0064 – – – – – 0.1503 0.00373 0.0034 – – 0.1501 0.0600 0.0268 0. www.1432 0.0509 0.1925 0.0684 0.0732 0.0662 0.0585 0.0579 0.0464 0.

1320 0.0565 0.0777 Bell et al.0012 0.0016 0.1372 0.0591 0.0535 0.0563 0.1607 0. www.0018 0. United Republic Of Great Britain 0.pnas.0693 0.0406 0.0139 0.0047 – 0.001 – – – Cultural FST 0.0794 0.0680 0.0016 0.0600 0.0644 0.0439 0.0850 0.0525 0.0729 0.0322 0.0589 0.0456 0.2116 0.0568 0.1131 0.0659 0.0048 – – – – – – – – – – – – 0.0439 0.1705 0.0039 0.0082 – 0.0688 0.0025 0.1689 0.0544 0.Genetic FST Netherlands Netherlands Netherlands Netherlands Norway Norway Philippines Philippines Poland Poland Poland Poland Portugal Puerto Rico Puerto Rico Republic Of Moldova Republic Of Moldova Romania Romania Russian Federation Russian Federation Serbia And Montenegro Singapore Slovakia South Africa Spain Sweden Sweden Switzerland Taiwan Province Of China Uganda United States Belgium Germany Germany West Great Britain Great Britain Sweden China Viet Nam Germany Slovakia Sweden Ukraine Spain United States Venezuela Romania Ukraine Serbia And Montenegro Ukraine Turkey Ukraine Bosnia And Herzegovina Indonesia Ukraine Zimbabwe France Germany West Norway Germany Philippines Tanzania.0600 0.0039 0.0315 – 0.0018 0.0017 4 of ؊2 .

003 – 0. Estimates for cultural FST derived from (7) on the lower diagonal.105 Sebei 0.003 0.004 – Bell et al. with bootstrapped standard errors in the upper diagonal Hehe Hehe Kamba Pokot Sebei – 0.003 0.099 Pokot 0.004 – 0.087 Kamba 0. 5 of ؊2 .113 0.Table S2.pnas.123 0.004 0.109 0.