Tuesday, September 26, 2017

Tales from old bones

Around three thousand years ago Bantu began to spread east and south from the Nigeria/Cameroun border, eventually replacing the original inhabitants of eastern and southern Africa. Those people no longer exist. Only the DNA in their skeletal remains are left to speak for them.

When scientists began to retrieve ancient DNA from human remains, they succeeded only at sites in the temperate and arctic zones. It seemed impossible to retrieve any at tropical sites, apparently because warm year-round temperatures soon reduce DNA to a meaningless molecular jumble.

This problem seems to be solved. Two years ago, DNA was successfully retrieved from 4,500 year old remains in Ethiopia. Now, we have ancient DNA from several sites across eastern and southern Africa over a range of dates from 10,000 to 400 years ago (Skoglund et al. 2017).

Vanished peoples

This new study shows that eastern and southern Africans have changed a lot since the time of the ancient Greeks. As far north as Tanzania, the continent was once home to peoples related to the Hottentots (now called Khoisans, Khoe-Sans, or simply San)—short in stature, gracile in body build, and light yellowish brown in color. From Zanzibar north, people were of mixed Middle Eastern and Cushitic origin—sort of like present-day Ethiopians but with more Arab ancestry.

What happened to these peoples? They were either replaced or absorbed by Bantus moving in from the west, although it now looks like they were replaced a lot more than they were absorbed. No trace of them remains in Malawi's gene pool:

Population replacement by incoming food producers appears to have been nearly complete in Malawi, where we detect little if any ancestry from the ancient individuals who lived ~8,100-2,500 BP. Instead, present-day Malawian individuals are consistent with deriving all their ancestry from the Bantu expansion of ultimate western African origin. (Skoglund et al. 2017)

The original inhabitants were related to present-day Khoisans but had significantly diverged from them:

Notably, the Khoe-San-related ancestry in ancient individuals from Malawi and Tanzania is symmetrically related to the two previously identified lineages present in the San [...], estimated to have diverged at least 20,000 years ago [...], implying that this was an ancient divergent branch of this group that lived in eastern Africa at least until 1,400 BP. (Skoglund et al. 2017)

This is in line with previous DNA findings from the Fwe (a Bantu group of southwestern Zambia), particularly the presence of Khoisan admixture that resembles nothing in present-day Khoisans:

[It is possible] that the Fwe intermarried with a Khoisan group whose genetic composition differed from that of the populations included in molecular anthropological investigations to date. [...] it is plausible that the Fwe ancestors interacted with a Khoisan community that differed genetically from those still settled in southern Africa today, which was ultimately replaced by the newcomers. (Barbieri et al. 2013)

Aside from these scattered fragments of DNA, we also have the testimony of ancient observers. Two tenth-century Arab geographers state that "in the outer reaches of the land of the Zanj there are cool highlands in which live white Zanj" (Lewis 1990, p. 121, n. 3). The Zanj are the dark-skinned peoples of east Africa and the term 'white' is better translated by 'lighter-skinned.' (The words 'black' and 'white' are often used in a relative sense in Arabic). The highlands might be the Drakensberg Escarpment of South Africa. 

Encounters with the archaic Other

Modern humans arose some 80,000 years ago in eastern Africa through a series of population expansions that culminated twenty thousand years later in a big bang that spread outward in Africa and then into the Middle East, Europe, and Asia (Watson et al. 1997). There, they encountered more archaic hominins: Neanderthals and, farther east, Denisovans. There was some intermixture. How much? Some have argued that modern Europeans and Asians are 3.4 to 7.9 percent admixed (Lohse and Frantz 2013). Most still opt for a lower figure of 1.5 to 2.1 percent (Prüfer et al. 2014).

But it wasn't only in Eurasia that modern humans encountered Neanderthal-like groups. Archaic hominins were present in Africa itself, some being relatively close to modern humans, and some more distantly related.

The latest DNA study has confirmed that modern humans intermixed with at least one archaic group as they expanded into western Africa:

The possible basal western African population lineage would represent the earliest known divergence of a modern human lineage that contributed a major proportion of ancestry to present-day humans. Such a lineage must have separated before the divergence of San ancestors, which is estimated to have begun on the order of 200-300 thousand years ago. (Skoglund et al. 2017)

This archaic ancestry is visible in human remains found at the Iwo Eleru rock shelter, in southwestern Nigeria, and dated to approximately 16,300 BP:

Our analysis indicates that Iwo Eleru possesses neurocranial morphology intermediate in shape between archaic hominins (Neanderthals and Homo erectus) and modern humans. This morphology is outside the range of modern human variability in the PCA and CVA analyses, and is most similar to that shown by LPA individuals from Africa and the early anatomically modern specimens from Skhul and Qafzeh. (Harvati et al., 2011)

Archaic ancestry is also visible in present-day West Africans, particularly in their teeth: 

[...] compared to other world populations, Africans south of the Sahara Desert are distinct dentally — especially in their expression of nine high- and two low-frequency morphological features. [...] the same nine high-frequency traits are also ubiquitous in the dentitions of extinct hominids and many extinct and extant non-human primates.  
[...] The presence and, indeed, prevalence (see next section), of high-frequency Sub-Saharan dental traits in fossil and recent hominoids—some of which are probably direct ancestors of modern humans, suggests they have been around for a long time.  
[...] A final ancestral feature found with some regularity in Sub-Saharan Africans, relative to other modern groups, is polydontia. Numerous cases of extra incisors, third premolars, and fourth molars have been noted [...] In one study (Watters, 1962) the incidence reached 2.5-3% in several hundred west Africans; many of the extra teeth were fully formed and erupted. "Typical" mammals exhibit three incisors and four premolars (Jordan et al., 1992). Polydontia is also found in living non-human primates. (Irish, 1998)

How much archaic ancestry do sub-Saharan Africans have today? The latest DNA study is silent on this point. Any answer can only be approximate, there being no reconstructed genome of this Neanderthal-like population. Moreover, there was probably more than one such population within Africa. Watson et al. (1997) attribute 13% of the sub-Saharan gene pool to a population that expanded some 111,000 years ago—when Skhul-Qafzeh hominins entered the Middle East from Africa. Those hominins were anatomically modern, or almost so, but culturally Neanderthal. Hammer et al. (2011) estimate that about 2% of the sub-Saharan African genome comes from a much more divergent population that split off from the ancestors of modern humans some 700,000 years ago. That admixture entered the sub-Saharan gene pool about 35,000 years ago, perhaps in Central Africa, since pygmy groups from that region have the most.

It looks like the proportion of archaic ancestry is higher in sub-Saharan Africans than in other modern humans. This is to be expected because of the broader range of archaic populations in Africa, including some that were almost modern anatomically and behaviorally. Admixture with them would have been likelier.

Admixture: good, bad, or neither?

Some alleles have successfully introgressed from archaic hominins, thus helping our ancestors adapt to new climates and new diets (Racimo et al. 2015). In general, however, we should not expect such alleles to perform as well in the body of a modern human as they did in the body of an archaic hominin. It's like taking a part from a Chevy and installing it on a Subaru. It might work, but I wouldn't count on it. 

If we look at Neanderthal admixture in the Eurasian genome, we see that natural selection has tended to remove functional genes, while leaving the non-functioning ones alone. 

Neanderthal ancestry decreases in proximity to functional elements in all populations [...] as does Denisovan ancestry in Oceanians [...] most likely reflecting greater selection against Neanderthal ancestry in low B statistic regions. Power to detect archaic ancestry is elevated close to regions of linked selection due to a reduction in the rates of incomplete lineage sorting caused by the lower effective population size in these regions, so these results are not artifacts of reduced power. Thus, similar processes appear to have worked to remove Neanderthal and Denisovan ancestry near genes. (Sankararaman et al. 2016)

Archaic admixture is also associated with reduced male fertility:

Our study provides new evidence in support of the hypothesis that reduced male fertility may be a common feature of admixture between human populations diverged by at least a half million years, a hypothesis that was previously suggested based on genetic patterns associated with the hybridization between Neanderthals and modern humans.

[...] One line of evidence for reduced fertility in male hybrids is that the proportion of archaic ancestry in modern humans is significantly reduced on chromosome X compared to the autosomes. This is suggestive of reduced male fertility as loci contributing to this phenotype are concentrated on chromosome X in hybrids of other species. We confirm an extreme reduction of Neanderthal ancestry on chromosome X (16%-34% of the autosomes depending on the population) and find a quantitatively similar reduction of Denisovan ancestry (21% of the autosomes in Oceanians).

The second line of evidence in support of the hypothesis of reduced fertility in hybrids is that there is a reduction of archaic ancestry in genes that are disproportionately expressed in testes, a known characteristic of male hybrid fertility (Sankararaman et al. 2016)

In sum, archaic admixture did provide modern humans with some ready-made alleles that have helped them adapt to new climates and new diets, but this advantage hardly applies to Africa. There, modern humans were already adapted to the local climate and diet. Archaic admixture couldn't have done much to help them adapt, since the new environments they faced were cultural ones of their own making.


Barbieri, C., A. Butthof, K. Bostoen, and B. Pakendorf. (2013). Genetic perspectives on the origin of clicks in Bantu languages from southwestern Zambia, European Journal of Human Genetics, 21(4), 430-436.

Hammer, M.F., A.E. Woerner, F.L. Mendez, J.C. Watkins, and J.D. Wall. (2011). Genetic evidence for archaic admixture in Africa, Proceedings of the National Academy of Science (USA), 108(37), 15123-15128, www.pnas.org/cgi/doi/10.1073/pnas.1109300108

Harvati, K., C. Stringer, R. Grün, M. Aubert, P. Allsworth-Jones, C.A. Folorunso. (2011). The Later Stone Age Calvaria from Iwo Eleru, Nigeria: Morphology and Chronology. PLoS ONE 6(9): e24024. doi:10.1371/journal.pone.0024024

Irish, J.D. (1998). Ancestral dental traits in recent Sub-Saharan Africans and the origins of modern humans, Journal of Human Evolution, 34, 81-98.

Lohse, K., and L.A.F. Frantz. (2013). Maximum likelihood evidence for Neandertal admixture in Eurasian populations from three genomes, Populations and Evolution, 1307, 8263

Prüfer, K., F. Racimo, N. Patterson, F. Jay; et.al. (2014). The complete genome sequence of a Neandertal from the Altai Mountains, Nature, 505(7481), 43-49.

Racimo, F., S. Sankararaman, R. Nielsen, and E. Huerta-Sanchez. (2015). Evidence for archaic adaptive introgression in humans, Nature Reviews Genetics, 16(6), 359-371.

Sankararaman, S., S. Mallick, N. Patterson, D, Reich; et al. (2016). The combined landscape of Denisovan and Neanderthal ancestry in present-day humans, Current Biology, 26(9), 1241-1247.

Skoglund, P., J.C. Thompson, M.E. Prendergast, A. Mittnik; et al. (2017). Reconstructing prehistoric African population structure, Cell, 171(1), 59-71

Watson, E., P. Forster, M. Richards, and H-J. Bandelt. (1997). Mitochondrial footprints of human expansions in Africa, American Journal of Human Genetics, 61, 691-704.

Saturday, September 16, 2017

An idea abandoned by its father

Italian wall lizard (Podarcis sicula) (Credit: Charles J. Sharp). Five mating pairs were taken from one island to another, and over the next thirty generations the transplanted population became remarkably different from the parent population.

Unlike other animals, we adapt not only to natural environments but also to cultural environments of our making. We thus direct our own evolution. At the same time we are busy redesigning our cultural environment, the latter is just as busy redesigning us. Like the natural environment, it favors the survival and reproduction of those who best fit in.

This concept of gene-culture coevolution began with anthropologist Claude Lévi-Strauss in a 1971 lecture:

... Among early humans, biological evolution may have selected for pre-cultural traits like upright posture, manual dexterity, sociability, symbolic thinking, and ability to vocalize and communicate. It was culture, however, once it came into being, that consolidated these traits and propagated them. When cultures specialize, they consolidate and favor other traits, like resistance to cold or heat in societies that have willingly or unwillingly had to adapt to extreme climates, like dispositions to aggressiveness or contemplation, like technical ingenuity, and so on. In the form these traits appear to us on the cultural level, none can be clearly linked to a genetic basis, but we cannot exclude that they are sometimes linked partially and distantly via intermediate linkages. In this case, it would be true to say that each culture selects for genetic aptitudes, which then, via a feedback loop, influence the culture that had initially helped to strengthen them.

Credit is usually given, however, to geneticist Luigi Luca Cavalli-Sforza. In 1976, he developed the first mathematical models for gene-culture coevolution with another geneticist, Marcus Feldman, and in 1978-1979 he spoke on this subject to a cultural evolution class at Stanford (Feldman & Cavalli-Sforza 1976; Stone & Lurquin 2005, p. 108). Two of his students were Robert Boyd and Peter Richerson, who later wrote a seminal book on gene-culture coevolution (Boyd & Richerson 1985). In the mid-1980s, he decided to test this concept in the field by investigating the cultural and genetic bases of artistic talent among the Inuit:

One of the most remarkable phenomena in the contemporary Canadian Arctic is the presence of highly-acclaimed art forms — carving in stone and ivory, and printing on paper. The question we ask is: how can we account for the wide-spread distribution of such talent in a small dispersed population? (Berry & Cavalli-Sforza 1986, p. 2)

To answer this question, he organized a joint project with psychologist John W. Berry at Queen's University and anthropologist Bernard Saladin d'Anglure at Université Laval. The Inuit were chosen for study because their high rate of adoption made it possible "to distinguish cultural from biological inheritance by studying correlations of adopted children with foster relatives on one hand and biological relatives on the other" (Berry & Cavalli-Sforza 1986, p. 5). Also, until recently, Inuit had lived off the land and, as such, had "abilities [that] are considered to be adaptive to a nomadic and hunting life style" (Berry & Cavalli-Sforza 1986, p. 3). Berry argued that the artistic talent of the Inuit came from certain mental skills that helped them during hunting.

Hunters, by this way of thinking, require good visual acuity, keen disembedding skills and a well-developed sense of spatial orientation. To hunt successfully, the hunter must be able to discern the object of the quest (which is often embedded in a complex visual landscape), then disembed the object, and finally return to home base. In contrast, agriculturalists need not develop these particular skills, but rather they need to invest in other areas of development, such as conservation (in both the economic and the Piagetian senses) and close social interactions. (Berry 2008, p. 3)

The project fell through. Cavalli-Sforza said he had to quit because of illness. Neither of his biographies, however, mention any illness during that time period (Frost 2014). Interestingly, his American biography ascribes his interest in culture at that time to a desire to disprove the existence of mental differences between human populations:

Yet another source of his interest in culture was the idea that the concept of human cultural learning was a valid weapon against racist arguments that differences between people (for example, different IQ scores among ethnic groups) were due to biologically determined "racial" differences. (Stone & Lurquin, 2005, p. 86)

The reality was a bit different. He believed in the importance of culture, but not as an entity separate and distinct from biology. This put him in opposition not only to the racists he denounced in the 1960s but also to the antiracists who increasingly viewed him with suspicion from the late 1980s onward.

With Cavalli-Sforza out of the picture, research on gene-culture coevolution languished over the next quarter-century. This field of research needed a high-profile champion in academia, and all of the possible candidates were either unable or unwilling.  Cavalli-Sforza never was suited for the job, being too timid and, frankly, too easy to blackmail. (Do you really think his wartime research on anthrax involved only mice?)

Lately, there seems to have been a renewal of interest, as seen in this review article on "Human biological and psychological diversity":

Humans migrated out of Africa at least 50,000 years ago and occupied many different ecological and climatological niches. Because of this, they evolved slightly different anatomical and physiological traits. For example, Tibetans evolved various traits that help them cope with the rigors of altitude; similarly, the Inuit evolved various traits that help them cope with the challenges of a very cold environment. It is likely that humans also evolved slightly different psychological traits as a response to different selection pressures in different environments and niches. One possible example is the high intelligence of the Ashkenazi Jewish people. Frank discussions of such differences among human groups have provoked strong ethical concerns in the past. We understand those ethical concerns and believe that it is important to address them. However, we also believe that the benefits of discussing possible human population differences outweigh the costs. (Winegard et al. 2017)

This article is a good read, and I was intrigued by its examples of fast evolution, particularly the Italian wall lizards. Five mating pairs were taken from one island to another, and over the next thirty generations the transplanted population became remarkably different from the parent population. The lizards were now larger, had shorter hind limbs, and could bite with much more force. Even more remarkably, they had a new morphological trait: a cecal valve—a muscle between the large and small intestines that slows down food movement and allows digestion of cellulose. This is an adaptation to the abundance of plant food on that island, but it is surprising that an entirely new trait could evolve so fast.

As far back as Darwin, biologists have described evolutionary change as slow. This is true only when organisms live in slowly changing environments. Transplant them into a very different one, and they will evolve very fast. This has been especially true for modern humans, who over the past 50,000 years have spread into a wide range of natural environments from the tropics to the arctic and into an even wider range of cultural environments:

Humans, like many animals, actively alter their environment, which changes the selection pressures they face (Laland et al. 2001; Laland and Sterelny 2006). In fact, humans may be the paradigmatic example of a niche-creating species, using brains rather than brawn to conquer the world (Baumeister 2005; Pinker 2010). Across the globe, humans devised distinctive cultural systems to cope with their environments, creating vastly different selective regimes from one culture to another. (Winegard et al. 2017)

Humans are indeed niche creators who have speeded up their own diversification. Nonetheless, they aren't alone in diversifying so fast. For example, some animal and plant species have spread into a wide range of new habitats since the last ice age, thereby giving rise to many new populations. Whether these recent populations are "sibling species," "subspecies," or "races"—the distinction is often arbitrary—their example can help us understand our own genetic diversity (Frost 2011).

These populations, like our own, seem to have evolved much more anatomically than they have genetically. Their anatomies are often distinct from each other, with no overlap, yet their genomes overlap considerably—there is far more genetic variation within each population than between them. So they are easier to tell apart by their appearance than by their genes. 

Why this discordance between genes and anatomy? Genes don't lie, do they? To make sense of this puzzle, we need to understand three points:

  • When a gene has different "alleles" or versions of itself, these alleles vary in their degree of similarity, some performing very differently and others identically or almost so—often because the gene itself is little more than "junk DNA.
  • Population boundaries separate not only different populations but also different natural or cultural environments. This is especially true for humans. The cultural environment usually differs, even when the natural environment is the same.
  • If the alleles of a gene perform differently, some of them will be more useful to one population than to another because they do better in one environment than in another. The more differently they perform, the more their frequencies will differ across population boundaries, with some alleles being more common in some populations than in others. Conversely, if the alleles perform identically, they will do equally well in all environments and tend to be equally common in all populations. To the extent that different alleles are present within a single population, the reasons will be more stochastic and less related to the usefulness of any one allele. Genetic variation within a population is therefore disproportionately due to alleles that perform similarly.

So genetic variation between populations differs qualitatively from genetic variation within each population. The first kind matters a lot more than the second kind. There are exceptions to this rule, e.g., balanced polymorphisms, founder effects, genetic drift, but that's the general picture. So when you read that genes vary far more within human populations than between them, you should keep in mind that we see the same pattern with many sibling species that are nonetheless anatomically and behaviorally distinct. This pattern tells us only that the populations in question are very young. It doesn't tell us how different they really are from each other, since real evolutionary change can happen very fast—as we saw with the Italian wall lizards. 


Berry, J.W. (2008). Models of Ecocultural Adaptation and Cultural Transmission: The Example of Inuit Art, paper presented at the conference Adaptation et socialisation des minoritiés culturelles en région, June 3-4, Quebec City.

Berry, J.W., and L.L. Cavalli-Sforza. (1986). Cultural and genetic influences on Inuit art. Report to Social Sciences and Humanities Research Council of Canada, Ottawa.

Boyd, R. and P.J. Richerson. (1985). Culture and the Evolutionary Process, Chicago: Chicago University Press.

Feldman, M.; Cavalli-Sforza, L. (1976). Cultural and biological evolutionary processes, selection for a trait under complex transmission, Theoretical Population Biology, 9: 238-59.

Frost, P. (2014). L.L. Cavalli-Sforza. A bird in a gilded cage, Open Behavioral Genetics, March 28,

Frost, P. (2011). Human nature or human natures? Futures, 43: 740-748.
Lévi-Strauss, C. 1971). Race et culture, conférence de Lévi-Strauss à l'UNESCO le 22 mars 1971 (Audio). Polit'productions

Stone, L. and P.F. Lurquin. (2005). A Genetic and Cultural Odyssey. The Life and Work of L. Luca Cavalli-Sforza, New York: Columbia University Press.

Winegard, B., B. Winegard, and B. Boutwell. (2017). Human biological and psychological diversity, Evolutionary Psychological Science, 3(2): 159-180.

Monday, September 4, 2017

Gene-culture coevolution in northwest Europe

Notre-Dame-de-Compassion, Paris. Women have more affective empathy than do men, an indication that it originated served to bind a mother to her young children. To different degrees, and in different societies, it has become extended to other human relations. 

I've published a new paper: The Hajnal line and gene-culture coevolution in northwest Europe. This is the abstract:

North and west of a line running from Trieste to St. Petersburg, social relations have long conformed to the Western European Marriage Pattern, i.e., men and women marry relatively late; many people never marry; children usually leave the nuclear family to form new households, and households often have non-kin members. This pattern goes back at least to the thirteenth century and perhaps to prehistoric times. I argue that this environment of weaker kinship caused northwest Europeans to create communities based on shared moral rules, rather than shared kinship. Community members enforced these rules by monitoring not only the behavior of other members but also their own behavior and even their own thoughts. Initially, this new mindset did not have a genetic basis. Individuals acquired it within the bounds of phenotypic plasticity. Over time, however, a genetic basis would have developed through the survival and reproduction of individuals who were better at being socially independent, at obeying universal rules, at monitoring other community members, and at self-monitoring, self-judging, and self-punishing. These psychological adaptations-independent social orientation, universal rule adherence, affective empathy, guilt proneness-are moderately to highly heritable. Although they are complex, they required only minor evolutionary changes to evolve out of mechanisms that were already present but limited to specific behavioral contexts. Affective empathy, for instance, is a species-wide trait but usually confined to relations with close kin, particularly between a mother and her young children. An evolutionary scenario is proposed, and two questions discussed. Are these mental traits too complex to have evolved over a span of 30 to 300 generations? Are they too altruistic to be sustainable?


In a series of tweets, hbd* chick has accused me of plagiarizing her work without acknowledgment. This is a serious accusation, so please let me address it at length. 

I first became interested in the Western European Marriage Pattern (WEMP) in the early 1990s. I wanted to know why marriage systems are so different in Europe from elsewhere in the world. The high incidence of polygyny in sub-Saharan Africa can be explained by the mode of subsistence (year-round horticulture with high female participation and low male participation). But why is polygyny stigmatized so much more in European societies than elsewhere in Eurasia? Why the emphasis on monogamy and on postponement of marriage? Most scholars seemed to blame Christianity, but I knew, from my own readings, that polygyny had been stigmatized in Europe before Christianization. I made this point in an exchange with Kevin MacDonald in the pages of Ethology and Sociobiology in 1991 and 1992. I later discovered Wally Seccombe’s book A Millennium of Family Change. Feudalism to Capitalism in Northwestern Europe (1992). It was this book that introduced me to the WEMP, and it was at that point in time that I began to wonder whether the WEMP might explain the development of "guilt culture" (as opposed to "shame culture") in northwest Europe.

I first put forward some of these ideas in a post on my blog, "The Western European Marriage Pattern" (November 12, 2011). At that time, I knew hbd* chick had also written about the WEMP, but her posts hadn't contributed to the development of my thinking. I recognize that she has come up with original ideas on this topic, particularly her belief in the key role of manorialism, but I don't share that belief. Like Seccombe, I believe that the WEMP goes farther back in time. At one point, I tried to incorporate her ideas into my thinking, but I eventually gave up. I may be wrong, and I would like to be proven wrong, but the northwest European mindset seems to be much older than the Middle Ages.

So what did I plagiarize without acknowledgment? Whatever I know about this subject comes from authors who predate hbd chick, especially Wally Seccombe, but also Alan MacFarlane and Ruth Benedict. And I discovered those authors independently of hbd* chick. I'm especially flabbergasted by her claim that she originated the notion of Western European "guilt culture." Is she older than Ruth Benedict? Nor did she (or I) originate the notion that independent social orientation is stronger in northwest Europe. Alan MacFarlane and, more recently, Joan Chiao and Katherine Blizinsky have been the trailblazers in that area. Yes, credit should be given where credit is due, and I have given credit to those people who have the strongest moral claim to those ideas.

I wanted to insert a reference in my article to hbd* chick, but that insertion would have simply described her belief that the spread of manorialism was key to the WEMP (whereas I feel otherwise). Unfortunately it's difficult to cite an anonymous blogger in an academic publication because reviewers and editors have a strong prejudice against such citations. I tried doing that once for a previous article, and I was persuaded to find another source. Citing Wikipedia is usually possible, but citing an anonymous blogger is a bridge too far. The reasons are understandable. When people write under a pseudonym and without peer review, there is no way to prevent publication of poorly formulated views that are written impulsively and in the heat of the moment. I'm not supporting political correctness here. I'm supporting responsible scholarship.

I can understand why many people choose to publish under a pseudonym, especially on topics like this one. I've had to endure personal attacks for publishing under my own name, and my name can be linked to a lot of things: home address, personal photos, place of work, and so on. That's the price I've chosen to pay. In exchange, my publications can be cited in the academic literature. 

Aside from that one point (the spread of manorialism), I feel no moral obligation to cite hbd* chick for a topic that has long been discussed by many other people. My interest in the WEMP is quite different from hers. I wanted to explain why guilt plays such a strong role in the cultures of northwest Europe. The usual explanation is that northwest Europeans feel guilty because they have a lot to feel guilty about: slavery, colonialism, the Holocaust, etc. But how would that explain a guilt-ridden country like Sweden? The Swedes played little or no role in those historical events. In any case, the roots of northwest European guilt culture go much farther back in time.

I hope everyone will think over what I have written here. I don't like to treat people wrongly, and it pains me to be accused of wrongdoing.


I'm back to blogging, perhaps a new column every two weeks. To be honest, I have mixed feelings. On the one hand, blogging forces me to come up with new ideas, which can later be written up as academic articles, like the above. On the other hand, a single column can easily consume five to seven hours of my time. There are also legal ramifications: I can be held responsible not only for what I write but also for what anonymous commenters write.


Benedict, R. (1946 [2005]). The Chrysanthemum and the Sword. Patterns of Japanese Culture. First Mariner Books.

Chiao, J.Y. & Blizinsky, K.D. (2010). Culture-gene coevolution of individualism-collectivism and the serotonin transporter gene. Proceedings of the Royal Society B, 277, 529-537.

Frost, P. (2017). The Hajnal line and gene-culture coevolution in northwest Europe, Advances in Anthropology, 7, 154-174.

Frost, P. (2011). The Western European Marriage Pattern, November 12, 2011 http://evoandproud.blogspot.ca/2011/11/western-european-marriage-pattern.html

Frost, P. (1991). Letter to the Editors, Ethology and Sociobiology, 12(5), 335-336.

Macfarlane, A. (2012). The invention of the modern world. Chapter 8: Family, friendship and population. The Fortnightly Review, Spring-Summer serial

Macfarlane, A. (1992). On individualism. Proceedings of the British Academy, 82, 171-199. 

Macfarlane, A. (1978). The origins of English individualism: Some surprises. Theory and Society, 6, 255-277. 

Seccombe, W. (1992). A Millennium of Family Change. Feudalism to Capitalism in Northwestern Europe, London: Verso.