How some populations adapt to vitamin D scarcity

 

Vitamin D metabolism varies from one human population to another, as do most heritable traits. This is the subject of a review article I’ve recently published in the journal Nutrients. Here is the abstract:

 

 

Vitamin D metabolism differs among human populations because our species has adapted to different natural and cultural environments. Two environments are particularly difficult for the production of vitamin D by the skin: the Arctic, where the skin receives little solar UVB over the year; and the Tropics, where the skin is highly melanized and blocks UVB. In both cases, natural selection has favored the survival of those individuals who use vitamin D more efficiently or have some kind of workaround that ensures sufficient uptake of calcium and other essential minerals from food passing through the intestines. Vitamin D scarcity has either cultural or genetic solutions. Cultural solutions include consumption of meat in a raw or boiled state and extended breastfeeding of children. Genetic solutions include higher uptake of calcium from the intestines, higher rate of conversion of vitamin D to its most active form, stronger binding of vitamin D to carrier proteins in the bloodstream, and greater use of alternative metabolic pathways for calcium uptake. Because their bodies use vitamin D more sparingly, indigenous Arctic and Tropical peoples can be misdiagnosed with vitamin D deficiency and wrongly prescribed dietary supplements that may push their vitamin D level over the threshold of toxicity.

 

 

Reference

 

Frost P. (2022) The Problem of Vitamin D Scarcity: Cultural and Genetic Solutions by Indigenous Arctic and Tropical Peoples. Nutrients 14(19):4071. https://doi.org/10.3390/nu14194071

 

Evolution of long head hair

 

Swan princess, John Bauer (1882-1918). Human head hair is of relatively recent origin, reaching incredible lengths in some groups but not in others.

 

I've published an article on the evolution of long head hair in humans. The following is the abstract:

In many humans, head hair can grow to a much greater length than hair elsewhere on the body. This is a "derived" form that evolved outside Africa and probably in northern Eurasia. The ancestral form, which is frizzier and much shorter, survives in sub-Saharan Africans and in other groups whose ancestors never left the tropics. This original hair form is nonetheless relatively straight and silky during infancy. Head hair thus seems to have lengthened in two stages: 1) retention of the infant hair form at older ages; and 2) further lengthening to mid-back and even waist lengths. These changes seem to have gone farther in women, whose head hair is thicker and somewhat longer. The most popular evolutionary explanations are: 1) relaxation of selection for short hair; and 2) sexual selection for women with long hair. Neither hypothesis is satisfactory. The first one cannot explain why head hair lengthened so dramatically over so little time. The second hypothesis suffers from the assumption that some populations have remained naturally short-haired because they consider long-haired women undesirable. Almost the opposite is true in traditional African cultures, which have a long history of lengthening and straightening women's hair. It is argued here that sexual selection produced different outcomes in different populations not because standards of beauty differed but because the intensity of sexual selection differed. In the tropical zone, sexual selection acted more on men than on women and was thus too weak to enhance desirable female characteristics. This situation reversed as ancestral humans spread northward into environments that tended to limit polygyny while increasing male mortality. Because fewer men were available for mating, women faced a more competitive mate market and were selected more severely.

Reference

Frost, P. (2015). Evolution of long head hair in humans, Advances in Anthropology, 5, 274-281.

http://www.scirp.org/Journal/PaperInformation.aspx?PaperID=60916

 

Polygyny makes men bigger, tougher ... and meaner

 

Hadza men are smaller, less robust, and less aggressive than the more polygynous Datoga (Wikicommons - Idobi).

 

Humans differ in paternal investment—the degree to which fathers help mothers care for their offspring. They differ in this way between individuals, between populations, and between stages of cultural evolution.

During the earliest stage, when all humans were hunter-gatherers, men invested more in their offspring with increasing distance from the equator. Longer, colder winters made it harder for women to gather food for themselves and their children. They had to rely on meat from their hunting spouses. Conversely, paternal investment was lower in the tropics, where women could gather food year-round and provide for themselves and their children with little male assistance.

This sexual division of labor influenced the transition to farming. In the tropics, women were the main providers for their families as gatherers of fruits, berries, roots, and other wild plant foods. They were the ones who developed farming, thereby biasing it toward domestication of wild plants.

This may be seen in sub-Saharan Africa, where farming arose near the Niger's headwaters and gave rise to the Sudanic food complex—a wide range of native crops now found throughout the continent (sorghum, pearl millet, cow pea, etc.) and only one form of livestock, the guinea fowl (Murdock, 1959, pp. 44, 64-68). Many wild animal species could have been domesticated for meat production, but women were much less familiar with them. Men knew these species as hunters but had little motivation to domesticate them. Why should they? Women were the main providers. 

And so women shouldered even more the burden of providing for themselves and their offspring. Men in turn found it easier to go back on the mate market and get second or third wives. Finally, men had to compete against each other much more for fewer unmated women.

 

There was thus a causal chain: female dominance of farming => female reproductive autonomy => male polygyny => male-male rivalry for access to women. Jack Goody (1973) in his review of the literature says: "The desire of men to attract wives is seen as correlated with the degree of women's participation in the basic productive process." The more women produce, the lower the cost of polygyny.

In sub-Saharan Africa, the cost was often negative. Goody quotes a 17th century traveler on the Gold Coast: the women till the ground "whilst the man only idly spends his time in impertinent tattling (the woman's business in our country) and drinking of palm-wine, which the poor wives are frequently obliged to raise money to pay for, and by their hard labour maintain and satisfie these lazy wretches their greedy thirst after wines."

Goody cites data from southern Africa showing that the polygyny rate fell when the cost of polygyny rose:

In Basutoland one in nine husbands had more than one wife in 1936; in 1912, it was one in 5.5 (Mair 1953: 10). Hunter calculates that in 1911 12 per cent of Pondo men were plurally married and the figure was slightly lower in 1921. In 1946, the Tswana rate was 11 per cent; according to a small sample collected by Livingstone in 1850 it was 43 per cent. The figures appear to have changed drastically over time and the reasons are interesting. 'The large household is now not a source of wealth, but a burden which only the rich can bear' (Mair 1953: 19). Not only is there a specific tax for each additional wife, but a man's wives now no longer give the same help in agriculture that they did before. One reason for this is that the fields are ploughed rather than hoed. Among the Pondo, 'the use of the plough means that the amount of grain cultivated no longer depends on women's labour' (Goody, 1973)

Although polygynous marriage has become less common in southern Africa, polygynous behavior seems as frequent as ever. To a large degree, polygynous marriage has given way to more transient forms of polygyny: prostitution and other informal arrangements.

Goody also notes that women are much less self-reliant in the northern savannah of West Africa:

In savannah regions where water is scarce and trees scattered, their collection may make great demands on a woman’s time. So too does the grinding of hard grain, in the absence of mills. In all these domestic pursuits the savannah is more demanding on a woman’s time than the forest and consequently she can often make less contribution to agriculture. (Goody, 1973)

Yet polygyny rates have remained high. Goody gives the example of Ghana. Polygyny rates are about the same in the north and the south, yet in the north men participate much more in farming.

So what is going on? Goody concludes that "female farming and polygyny are clearly associated in a general way" but ultimately the "reasons behind polygyny are sexual and reproductive rather than economic and productive." It would be more parsimonious to say that the polygyny rate increases when the cost of providing for a woman and her children decreases for men. Over time, low-cost polygyny selects for men who are more motivated to exploit sexual opportunities. This new mindset influences the subsequent course of gene-culture coevolution.

Such gene-culture coevolution has gone through four stages in the evolutionary history of sub-Saharan Africans:

First stage

Tropical hunter-gatherers were already oriented toward low paternal investment. Men had a lesser role in child rearing because year-round food gathering provided women with a high degree of food autonomy. Women were thus selected for self-reliance and men for polygyny. Pair bonding was correspondingly weak in both sexes.

Second stage

This mindset guided tropical hunter-gatherers in their transition to farming. In short, female-dominated food gathering gave way to female-dominated horticulture—hoe farming of various crops with almost no livestock raising. Women became even more autonomous, and men even more polygynous. There was thus further selection for a mindset of female self-reliance, male polygyny, and weak pair bonding.

Third stage

A similar process occurred with the development of trade. Female-dominated horticulture tended to orient women, much more than men, toward the market economy. This has particularly been so in West Africa, where markets are overwhelmingly run by women. Trade has thus become another means by which African women provide for themselves and their children.

Fourth stage

Female-dominated horticulture has given way to male-dominated farming (pastoralism, cereal crops) in some regions, such as the northern savannah regions of West Africa. Despite higher male participation in farming, the pre-existing mindset has tended to maintain high polygyny rates. We see a similar tendency in southern Africa, where polygyny rates have fallen over the past century, and yet polygynous behavior persists in the form of prostitution and less formal sexual arrangements.

The Hadza and the Datoga

Mode of subsistence, mating system, and mindset are thus interrelated. These interrelationships are discussed by Butovskaya et al. (2015) in their study of two peoples in Tanzania: the largely monogamous Hadza (hunter-gatherers) and the highly polygynous Datoga (pastoralists). In their review of previous studies, the authors note:

In hunter-gatherer societies, such as the monogamous Hadza of Tanzania (Africa), men invest more in offspring than in small-scale pastoralist societies, such as the polygynous Datoga of Tanzania [12-14]. Polygyny and between-group aggression redirect men's efforts from childcare toward investment in male-male relationships and the pursuit of additional mates [15]. When men participate in childcare, their testosterone (T) level decreases [15-18]. Muller et al. [19] found that, among the monogamous, high paternally investing Hadza, T levels were lower for fathers than for non-fathers. This effect was not observed among the polygynous, low paternally investing Datoga. (Butovskaya et al., 2015).

Butovskaya et al. (2015) confirmed these previous findings in their own study:

Datoga males reported greater aggression than Hadza men—a finding in line with previous reports [29,30]. It is important to mention several striking differences between these two cultures. There is a negative attitude toward aggression among the Hadza but not among the Datoga. In situations of potential aggression, the Hadza prefer to leave [30]. In contrast, aggression is an instrument of social control—both within the family and in outgroup relations in Datoga society. Datoga men are trained to compete with each other and to act aggressively in particular circumstances [30]

The authors also confirmed differences in reproductive behavior between the two groups: 

Our research indicates a difference in the number of children in Hadza and Datoga men achieved after the age of 50. This may be interpreted as differences attributable to different life trajectories and marriage patterns. Beginning in early childhood, boys in the two societies are subjected to different social and environmental pressures (e.g., it is typical for Datoga parents to punish children for misbehavior, while parental violence is much less typical for Hadza parents). Hadza men start reproducing in the early 20s, but their reproductive success later in life is associated with their hunting skills [15]. In the Datoga, men marry later, typically in their 30s. Male status and, consequently, social and reproductive success in the Datoga are positively correlated with fighting abilities and risk-taking in raiding expeditions among younger men, and with wealth, dominance, and social skills among older men. In the Datoga, as in other patrilineal societies, fathers do not invest directly in child care, but children do benefit from their father's investment in the form of wealth and social protection, as well as various services provided by father's patrilineal male relatives [56]. In polygynous societies, spending resources on attracting additional wives may be more beneficial [40,57,58]. It would be difficult for some men to invest directly in providing for all their children, given that men with multiple wives can father a considerable number of children, and that households with wives may be located at substantial distance from one another.

This behavioral difference seems to be mediated by differing levels of androgens, such as testosterone:

The effect of androgens, such as T, operates through stimulation of androgen receptors [21-23]. The androgen receptor (AR) gene contains a polymorphic and functional locus in exon 1, comprising two triplets (CAG and GGN). This locus supports a regulatory function that responds to T, with fewer CAG repeat clusters being more effective in transmitting the T signal [22]. Moreover, the length of the GGN repeat predicts circulating and free T in men.

At the androgen receptor gene, the authors found fewer CAG repeats in the Datoga than in the Hadza. The number of repeats was also more variable in the Datoga. The Datoga's higher and more variable polygyny rates thus seem to correlate with higher and more variable levels of testosterone.

The authors also wished to see whether these differing levels of testosterone correlate with differing levels of aggressiveness. To this end, they interviewed the Hadza and Datoga participants:

They were asked to provide information including their age, sex, marital status, number of children, ethnicity and aggression history (especially fights with other tribal members). All questions were read aloud in one-to-one dialogues and further explanations were provided, if necessary. Self-reported aggression was assessed with the Buss-Perry Aggression Questionnaire (BPAQ; [48]). The BPAQ includes 29 statements, grouped into four subscales—physical aggression (9 items), verbal aggression (5 items), anger (7 items), and hostility (8 items)—answered on aLikert scale anchored by 1 (extremely uncharacteristic of me) and 5 (extremely characteristic of me).

Total aggression was found to correlate negatively with CAG repeat number. Age group did not predict aggression.

More polygyny = stronger sexual selection of men

Finally, the authors suggest that Datoga men, with their higher polygyny rate and fiercer competition for access to women, have undergone greater sexual selection. They have thus become bigger and more masculine than Hadza men. Although this selection pressure also exists among the Hadza, the driving force of sexual selection has been weaker because Hadza men are more monogamous and less sexually competitive:

Our findings are in concordance with other research, demonstrating that even among the relatively egalitarian Hadza there is selection pressure in favor of more masculine men [59-62]. At the same time, preference for more masculine partners, with greater height and body size, is culturally variable and influenced by the degree of polygyny, local ecology, and other economic and social factors [59-62]. Many Datoga women commented that they would like to avoid taller and larger men as marriage partners, as they may be dangerously violent [44,62]. Only 2% of Hadza women listed large body size as an attractive mate characteristic [63]. Hadza marriages in which the wife is taller than the husband are common, and as frequent as would be expected by chance [64]. (Butovskaya et al., 2015)

This is consistent with what we see in nonhuman polygynous species. Successful males tend to be the ones that are better not only at attracting the opposite sex but also at fighting off rivals. They thus become bigger, tougher, and meaner.

This is also consistent with what we see generally in the highly polygynous farming peoples of sub-Saharan Africa. They and their African-American descendants exceed European-descended subjects in weight, chest size, arm girth, leg girth, muscle fiber properties, and bone density (Ama et al., 1986; Ettinger et al.,1997; Himes, 1988; Hui et al., 2003; Pollitzer and Anderson, 1989; Todd and Lindala, 1928; Wagner and Heyward, 2000; Wolff and Steggerda, 1943; Wright et al., 1995).

References 

Ama, P.F.M., J.A. Simoneau, M.R. Boulay, O. Serresse, G. Thériault, and C. Bouchard. (1986). Skeletal muscle characteristics in sedentary Black and Caucasian males, Journal of Applied Physiology, 61, 1758-1761.

http://www.educadorfisicoadinis.com.br/download/artigos/Skeletal%20muscle%20characteristics%20in%20sedentary%20black%20and%20Caucasian%20males.pdf  

Butovskaya M.L., O.E. Lazebny, V.A. Vasilyev, D.A. Dronova, D.V. Karelin, A.Z.P. Mabulla, et al. (2015). Androgen receptor gene polymorphism, aggression, and reproduction in Tanzanian foragers and pastoralists. PLoS ONE 10(8): e0136208. 

https://www.researchgate.net/publication/281170838_Androgen_Receptor_Gene_Polymorphism_Aggression_and_Reproduction_in_Tanzanian_Foragers_and_Pastoralists  

Ettinger, B., S. Sidney, S.R. Cummings, C. Libanati, D.D. Bikle, I.S. Tekawa, K. Tolan, and P. Steiger. (1997). Racial differences in bone density between young adult black and white subjects persist after adjustment for anthropometric, lifestyle, and biochemical differences, Journal of Clinical Endocrinology & Metabolism, 82, 429-434.

http://press.endocrine.org/doi/abs/10.1210/jcem.82.2.3732  

Goody, J. (1973). Polygyny, economy and the role of women, in J. Goody (ed.) The Character of Kinship, Cambridge: Cambridge University Press.

https://books.google.ca/books?hl=fr&lr=&id=TFjf4mUHqv4C&oi=fnd&pg=PA175&dq=Polygyny,+economy+and+the+role+of+women&ots=Lf9w6wxY9D&sig=3d0RBnoMbGUd2OYqghzhwdsO3BA#v=onepage&q&f=false

Himes, J. H. (1988). Racial variation in physique and body composition, Canadian Journal of Sport Sciences, 13, 117-126.

http://europepmc.org/abstract/med/3293730  

Hui, S.L., L.A. DiMeglio, C. Longcope, M. Peacock, R. McClintock, A.J. Perkins, and C.C. Johnston Jr. (2003). Difference in bone mass between Black and White American children: Attributable to body build, sex hormone levels, or bone turnover? Journal of Clinical Endocrinology & Metabolism, 88, 642-649. 

https://www.researchgate.net/profile/Conrad_Johnston/publication/10911942_Difference_in_bone_mass_between_black_and_white_American_children_attributable_to_body_build_sex_hormone_levels_or_bone_turnover/links/548f36af0cf225bf66a7fdc1.pdf

Murdock, G.P. (1959). Africa. Its Peoples and Their Culture History, New York: McGraw-Hill. 

Pollitzer, W.S. and J.J. Anderson. (1989). Ethnic and genetic differences in bone mass: a review with a hereditary vs environmental perspective, American Journal of Clinical Nutrition, 50, 1244-1259

http://ajcn.nutrition.org/content/50/6/1244.short 

Todd, T.W., and A. Lindala. (1928). Dimensions of the body: Whites and American Negroes of both sexes, American Journal of Physical Anthropology, 12, 35-101.

http://onlinelibrary.wiley.com/doi/10.1002/ajpa.1330120104/abstract  

Wagner, D.R. and V.H. Heyward. (2000). Measures of body composition in blacks and whites: a comparative review, American Journal of Clinical Nutrition, 71, 1392-1402.

http://ajcn.nutrition.org/content/71/6/1392.short

Wolff, G. and M. Steggerda. (1943). Female-male index of body build in Negroes and Whites: An interpretation of anatomical sex differences, Human Biology, 15, 127-152.

Wright, N.M., J. Renault, S. Willi, J.D. Veldhuis, J.P. Pandey, L. Gordon, L.L. Key, and N.H. Bell. (1995). Greater secretion of growth hormone in black than in white men: possible factor in greater bone mineral density-a clinical research center study, Journal of Clinical Endocrinology & Metabolism, 80, 2291-2297.

http://press.endocrine.org/doi/abs/10.1210/jcem.80.8.7543111  

Brain size and latitude: Why the correlation?

Human variation in cranial capacity. Black, 1,450 cc and over; checkerboard, 1,400-49 cc; crosshatching, 1,350-99 cc; horizontal striping, 1,300-49 cc; diagonal striping, 1,250-99 cc; dots, 1,200-49 cc; white areas, under 1,200 cc (Beals et al., 1984)

So Stephen Jay Gould was wrong, and Samuel George Morton was right. Brain size does vary among human populations. But what does this variation mean? Why does it exist?

This question is briefly addressed by Lewis et al. (2011):

[…] cranial capacity variation in human populations appears to be largely a function of climate, so, for example, the full range of average capacities is seen in Native American groups, as they historically occupied the full range of latitudes

For more information, the reader is referred to an earlier study: Beals, Smith, and Dodd (1984). The latter plotted cranial capacity from 122 human populations. The resulting map is shown above. A few caveats: there is a LOT of interpolation and extrapolation in the above map. There are no data points from Cambodia or southern Vietnam, and hence nothing to justify the very low values assigned to that region. Southern India is assigned a very low cranial capacity on the basis of a small sample that includes Veddas—a relic group of hunter-gatherers from Sri Lanka.

All in all, and keeping these caveats in mind, cranial capacity does seem to correlate with latitude. Why? According to the authors, heads are larger at higher latitudes to reduce heat loss. An object will lose less heat if its ratio of volume to surface area is high. There has thus been natural selection to make heads broader and more globular at higher latitudes. The increase in brain size is incidental.

This explanation was challenged in the comments section following the Beals, Smith, and Dodd (1984) article. A Japanese commenter, Iwatoro Morimoto, pointed out that "in recent centuries, brachycranic skulls show a considerable increase in frequency in Eurasian populations, including the Japanese." Since mean temperatures have changed little in recent centuries, there must have been another factor at work. Unfortunately, Morimoto provided no references to back up this counter-argument.

Another commenter, Erik Trinkaus, similarly pointed out that Neanderthal cranial capacity was no bigger during glacial periods than during interglacials. The same was true for early modern humans. For populations already established at northern latitudes, cranial capacity shows no evidence of rising and falling with mean temperature.

A recent analysis has nonetheless found a significant correlation between cranial capacity and latitude among ancestral hominids in general, ranging from A. Afarensis to H. sapiens (Henneberg and Miguel, 2004). The correlation remained even when the authors controlled for each skull’s time period and, thus, was not due to the overall rise in cranial capacity over time and the parallel expansion of ancestral hominids into higher latitudes.

In sum, cranial capacity does correlate with latitude. It is less clear, however, whether this correlation is mediated by mean temperature and the need to reduce heat loss.

Higher cognitive demands at higher latitudes?

Could it be that cognitive demands increased as ancestral humans entered higher latitudes? Not because mean temperatures were lower but because the yearly cycle presented a greater diversity of environments and required much more foresight. Between ‘summer’ and ‘winter,’ the differences are much greater in the temperate and arctic zones than in the tropics.

This point is elaborated upon by Hoffecker (2002, p. 135). Among early modern humans, tools and weapons were more complex at arctic latitudes than at tropical latitudes. “Technological complexity in colder environments seems to reflect the need for greater foraging efficiency in settings where many resources are available only for limited periods of time.” Arctic humans coped with resource fluctuations and high mobility requirements by planning ahead and by developing untended devices (e.g., traps and snares) and means of food storage.

In addition, these increased cognitive demands fell on both men and women. Paternal and maternal investment were much more equal than in the tropics, where women provided for their families year-round with less male assistance (Kelly, 1995, pp. 268-269; Martin, 1974, pp. 16-18). Indeed, because men were the main food providers beyond the tropical zone, women could care for their families by developing a new range of tasks: food processing (e.g., butchery and carcass transport); shelter building; garment making; leather working; transport of material goods; etc. (Waguespack, 2005). This technological revolution would ultimately lead to what we now call ‘civilization’ (Frost, 2008).

Further thoughts

Curiously, Beals, Smith, and Dodd (1984) cite Gould’s 1978 Science article—the one claiming that Morton had unconsciously fudged his data to make brains look bigger among Europeans than among sub-Saharan Africans. Yet these authors declined to mention the inconsistencies between Gould’s findings and their own. Their reference to Gould is studiously neutral: “Critiques of the use of brain size in typology have been offered by Gould.”

There has not been much comment on the Beals, Smith, and Dodd (1984) article. The most substantive one seems to be a blog post by Robert Lindsay (2010) who calls their map a “train wreck” for claims that cranial capacity correlates with IQ:

White racists like to make a big deal about the supposed correlation between head size and intelligence and race. A nice little chart showing the basically dishonest portrayal they attempt based on cherry-picking data is below.

Methinks that Lindsay takes the fine details on that map a bit too seriously. Many of the details are simply creative extrapolation and infilling; otherwise, the map roughly corresponds with world distribution of mean IQ. Furthermore, no one is claiming that cranial capacity is the only determinant of IQ. There are undoubtedly many others: cortical surface area, myelinization of nerve fibers, relative importance of domain-general thinking, etc.

But he does make a good point about the Amerindian data.

As you can see, in the Americas, there is no good evidence whatsoever for head size and IQ. I am not aware that Amerindian IQ varies in the Americas. The average is apparently 87 across the continent. If anyone can show me that it varies by latitude, please do.

Agreed. No one can, for now. But a hypothesis is not false because no one has bothered to test it.

References

Beals, K.L., C.L. Smith, and S.M. Dodd (1984). Brain size, cranial morphology, climate, and time machines, Current Anthropology, 25, 301–330.

Frost, P. (2008). The path to civilization? Evo and Proud, March 10, 2008.
http://evoandproud.blogspot.com/2008/03/path-to-civilization.html

Henneberg, M. and C. de Miguel. (2004). Hominins are a single lineage: brain and body size variability does not reflect postulated taxonomic diversity of hominins, Journal of Comparative Human Biology, 55, 21–37

Hoffecker, J.F. (2002). Desolate Landscapes. Ice-Age Settlement in Eastern Europe. New Brunswick: Rutgers University Press.

Kelly, R.L. (1955). The Foraging Spectrum. Diversity in Hunter-Gatherer Lifeways. Washington: Smithsonian Institution Press.

Lewis, J.E., D. DeGusta, M.R. Meyer, J.M. Monge, A.E. Mann, R.L. Holloway. (2011). The Mismeasure of Science: Stephen Jay Gould versus Samuel George Morton on Skulls and Bias, PLoS Biology, 9(6) e1001071

Lindsay, R. (2010). The Head Size/IQ/Race Trainwreck, March 11
http://robertlindsay.wordpress.com/2010/03/11/the-head-sizeraceiq-trainwreck/

Martin, M.K. (1974). The Foraging Adaptation — Uniformity or Diversity? Addison‑Wesley Module in Anthropology 56.

Waguespack, N.M. (2005). The organization of male and female labor in foraging societies: Implications for early Paleoindian archaeology. American Anthropologist, 107, 666-676.