Human skin variation comes down to how much UV radiation a population is exposed to. Over time, this leads to changes in genetic expression. If that new genotype is advantageous in that environment, it will get selected for. To see how human skin variation evolved, we must first look to chimpanzees since they are our closest relative.
The evolution of black skin
Humans and chimps diverged around 6-12 mya. Since we share 99.8 percent of our genome with them, it’s safe to say that when we diverged, we had pale skin and a lot of fur on our bodies (Jablonski and Chaplin, 2000). After we lost the fur on our bodies, we were better able to thermoregulate, which then primed Erectus for running (Liberman, 2015). The advent of fur loss coincides with the appearance of sweat glands in Erectus, which would have been paramount for persistence hunting in the African savanna 1.9 mya, when a modern pelvis—and most likely a modern gluteus maximus—emerged in the fossil record (Lieberman et al, 2006). This sets the stage for one of the most important factors in regards to the ability to persistence hunt—mainly, the evolution of dark skin to protect against high amounts of UV radiation.
After Erectus lost his fur, the unforgiving UV radiation beamed down on him. Selection would have then occurred for darker skin, as darker skin protects against UV radiation. Dark skin in our genus also evolved between 1 and 2 mya. We know this since the melanocortin 1 receptor promoting black skin arose 1-2 mya, right around the time Erectus appeared and lost its fur (Lieberman, 2015).
However, other researchers reject Greaves’ explanation for skin cancer being a driver for skin color (Jablonksi and Chaplin, 2014). They cite Blum (1961) showing that skin cancer is acquired too late in life to have any kind of effect on reproductive success. Skin cancer rates in black Americans are low compared to white Americans in a survey from 1977-8 showing that 30 percent of blacks had basal cell carcinoma while 80 percent of whites did (Moon et al, 1987). This is some good evidence for Greaves’ hypothesis; that blacks have less of a rate of one type of skin cancer shows its adaptive benefits. Black skin evolved due to the need for protection from high levels of UVB radiation and skin cancers.
Highly melanized skin also protects against folate destruction (Jablonksi and Chaplin, 2000). As populations move away from high UV areas, the selective constraint to maintain high levels of folate by blocking high levels of UV is removed, whereas selection for less melanin prevails to allow enough radiation to synthesize vitamin D. Black skin is important near the equator to protect against folate deficiency. (Also see Nina Jablonski’s Ted Talk Skin color is an illusion.)
The evolution of white skin
The evolution of white skin, of course, is much debated as well. Theories range from sexual selection, to diet, to less UV radiation. All three have great explanatory power, and I believe that all of them did drive the evolution of white skin, but with different percentages.
The main driver of white skin is living in colder environments with fewer UV rays. The body needs to synthesize vitamin D, so the only way this would occur in areas with low UV rays.
White skin is a recent trait in humans, appearing only 8kya. A myriad of theories have been proposed to explain this, from sexual selection (Frost, 2007), which include better vitamin D synthesis to ensure more calcium for pregnancy and lactation (which would then benefit the intelligence of the babes) (Jablonski and Chaplin, 2000); others see light skin as the beginnings of more childlike traits such as smoother skin, a higher pitched voice and a more childlike face which would then facilitate less aggressiveness in men and more provisioning (Guthrie, 1970; from Frost, 2007); finally, van den Berghe and Frost (1986) proposed that selection for white skin involved unconscious selection by men for lighter-skinned women which is used “as a measure of hormonal status and thus childbearing potential” (Frost, 2007). The three aforementioned hypotheses have sexual selection for lighter skin as a proximate cause, but the ultimate cause is something completely different.
The hypothesis that white skin evolved to better facilitate vitamin D synthesis to ensure more calcium for pregnancy and lactation makes the most sense. Darker-skinned individuals have a myriad of health problems outside of their ancestral climate, one of which is higher rates of prostate cancer due to lack of vitamin D. If darker skin is a problem in cooler climates with fewer UV rays, then lighter skin, since it ensures better vitamin D synthesis, will be selected for. White skin ensures better and more vitamin D absorption in colder climates with fewer UV rays, therefore, the ultimate cause of the evolution of white skin is a lack of sunlight and therefore fewer UV rays. This is because white skin absorbs more UV rays which is better vitamin D synthesis.
Peter Frost believes that Europeans became white 11,000 years ago. However, as shown above, white skin evolved around 8kya. Further, contrary to popular belief, Europeans did not gain the alleles for white skin from Neanderthals (Beleza et al, 2012). European populations did not lose their dark skin immediately upon entering Europe—and Neanderthal interbreeding didn’t immediately confer the advantageous white skin alleles. There was interbreeding between AMH and Neanderthals (Sankararaman et al, 2014). So if interbreeding with Neanderthals didn’t infer white skin to proto-Europeans, then what did?
A few alleles spreading into Europe that only reached fixation a few thousand years ago. White skin is a relatively recent trait in Man (Beleza et al, 2012). People assume that white skin has been around for a long time, and that Europeans 40,000 ya are the ancestors of Europeans alive today. That, however, is not true. Modern-day European genetic history began about 6,500 ya. That is when the modern-day European phenotype arose—along with white skin.
Furthermore, Eurasians were still a single breeding population 40 kya, and only diverged recently, about 25,000 to 40,000 ya (Tateno et al, 2014). The alleles that code for light skin evolved after the Eurasian divergence. Polymorphisms in the genes ASIP and OCA2 may code for dark and light skin all throughout the world, whereas SLC24A5, MATP, and TYR have a predominant role in the evolution of light skin in Europeans but not East Asians, which suggests recent convergent evolution of a lighter pigmentation phenotype in European and East Asian populations (Norton et al, 2006). Since SLC24A5, MATP, and TYR are absent in East Asian populations, then that means that East Asians evolved light skin through completely different mechanisms than Europeans. So after the divergence of East Asians and Europeans from a single breeding population 25-40kya, there was convergent evolution for light pigmentation in both populations with the same selection pressure (low UV).
Some populations, such as Arctic peoples, don’t have the skin color one would predict they should have based on their ancestral environment. However, their diets are high in shellfish which is high in vitamin D, which means they can afford to remain darker-skinned in low UV areas. UV rays reflect off of the snow and ice in the summer and their dark skin protects them from UV light.
Black-white differences in UV absorption
If white skin evolved to better synthesize vitamin D with fewer (and less intense) UV rays, then those with blacker skin would need to spend a longer time in UV light to synthesize the same amount of vitamin D. Skin pigmentation, however, is negatively correlated with vitamin D synthesis (Libon, Cavalier, and Nikkels, 2013). Black skin is less capable of vitamin D synthesis. Furthermore, blacks’ skin color leads to an evolutionary environmental mismatch. Black skin in low UV areas is correlated with rickets (Holick, 2006), higher rates of prostate cancer due to lower levels of vitamin D (Gupta et al, 2009; vitamin D supplements may also keep low-grade prostate cancer at bay).
Libon, Cavalier, and Nikkels, (2013) looked at a few different phototypes (skin colors) of black and white subjects. The phototypes they looked at were II (n=19), III (n=1), and VI (n-11; whites and blacks respectively). Phototypes are shown in the image below.
To avoid the influence of solar UVB exposure, this study was conducted in February. On day 0, both the black and white subjects were vitamin D deficient. The median levels of vitamin D in the white subjects was 11.9 ng/ml whereas for the black subjects it was 8.6 ng/ml—a non-statistically significant difference. On day two, however, concentrations of vitamin D in the blood rose from 11.9 to 13.3 ng/ml—a statistically significant difference. For the black cohort, however, there was no statistically significant difference in vitamin D levels. On day 6, levels in the white subjects rose from 11.6 to 14.3 ng/ml whereas for the black subjects it was 8.6 to 9.57 ng/ml. At the end of day 6, there was a statistically significant difference in circulating vitamin D levels between the white and black subjects (14.3 ng/ml compared to 9.57 ng/ml).
Different phototypes absorb different amounts of UV rays and, therefore, peoples with different skin color absorb different levels of vitamin D. Lighter-skinned people absorb more UV rays than darker-skinned people, showing that white skin’s primary cause is to synthesize vitamin D.
UVB exposure increases vitamin D production in white skin, but not in black skin. Pigmented skin, on the other hand, hinders the transformation of 7-dehydrocholesterol to vitamin D. This is why blacks have higher rates of prostate cancer—they are outside of their ancestral environment and what comes with being outside of one’s ancestral environment are evolutionary mismatches. We have now spread throughout the world, and people with certain skin colors may not be adapted for their current environment. This is what we see with black Americans as well as white Americans who spend too much time in climes that are not ancestral to them. Nevertheless, different-colored skin does synthesize vitamin D differently, and knowledge of this will increase the quality of life for everyone.
Even the great Darwin wrote about differences in human skin color. He didn’t touch human evolution in On the Origin of Species (Darwin, 1859), but he did in his book Descent of Man (Darwin, 1871). Darwin talks about the effects of climate on skin color and hair, writing:
It was formerly thought that the colour of the skin and the character of the hair were determined by light or heat; and although it can hardly be denied that some effect is thus produced, almost all observers now agree that the effect has been very small, even after exposure during many ages. (Darwin, 1871: 115-116)
Darwin, of course, championed sexual selection as the cause for human skin variation (Darwin, 1871: 241-250). Jared Diamond has the same view, believing that natural selection couldn’t account for hair loss, black skin and white skin weren’t products of natural selection, but female mate preference and sexual selection (Greaves, 2014).
Parental selection for white skin
Judith Rich Harris, author of the book The Nurture Assumption: Why Kids Turn Out the Way They Do (Harris, 2009), posits another hypothesis for the evolution of light skin for those living in northern latitudes—parental selection. This hypothesis may be controversial to some, as it states that dark skin is not beautiful and that white skin is.
Harris posits that selection for lighter skin was driven by sexual selection, but states that parental selection for lighter skin further helped the fixation of the alleles for white skin in northern populations. Neanderthals were a furry population, as they had no clothes, so, logic dictates that if they didn’t have clothes then they must have had some sort of protection against the cold Ice Age climate, therefore they must have had fur.
Harris states that since lighter skin is seen as more beautiful than darker skin, then if a woman birthed a darker/furrier babe than the mother would have committed infanticide. Women who birth at younger ages are more likely to commit infanticide, as they still have about twenty years to birth a babe. On the other hand, infanticide rates for mothers decrease as she gets older—because it’s harder to have children the older you get.
Harris states that Erectus may have been furry up until 2 mya, however, as I’ve shown, Erectus was furless and had the ability to thermoregulate—something that a hairy hominin was not able to do (Lieberman, 2015).
There is a preference for lighter-skinned females all throughout the world, in Africa (Coetzee et al, 2012); China and India (Naidoo et al, 2016; Dixson et al, 2007); and Latin America and the Philipines (Kiang and Takeuchi, 2009). Light skin is seen as attractive all throughout the world. Thus, since light skin allows better synthesize of vitamin D in colder climes with fewer UV rays, then there would have been a myriad of selective pressures to push that along—parental selection for lighter-skinned babes being one of them. This isn’t talked about often, but infanticide and rape have both driven our evolution (more on both in the future).
Harris’ parental selection hypothesis is plausible, and she does use the right dates for fur loss which coincides with the endurance running of Erectus and how he was able to thermoregulate body heat due to lack of fur and more sweat glands. This is when black skin began to evolve. So with migration into more northerly climes, lighter-skinned people would have more of an advantage than darker-skinned people. Infanticide is practiced all over the world, and is caused—partly—by a mother’s unconscious preferences.
Skin color and attractiveness
Lighter skin is seen as attractive all throughout the world. College-aged black women find lighter skin more attractive (Stephens and Thomas, 2012). It is no surprise that due to this, a lot of black women lighten their skin with chemicals.
In a sample of black men, lighter-skinned blacks were more likely to perceive discrimination than their darker-skinned counterparts (Uzogara et al, 2014). Further, in appraising skin color’s effect on in-group discrimination, medium-skinned black men perceived less discrimination than lighter- and darker-skinned black men. Lastly—as is the case with most studies—this effect was particularly pronounced for those in lower SES brackets. Speaking of SES, lighter-skinned blacks with higher income had lower blood pressure than darker-skinned blacks with higher income (Sweet et al, 2007). The authors conclude that a variety of psychosocial stress due to discrimination must be part of the reason why darker-skinned blacks with a high SES have worse blood pressure—but I think there is something else at work here. Darker skin on its own is associated with high blood pressure (Mosley et al, 2000). I don’t deny that (perceived) discrimination can and does heighten blood pressure—but the first thing that needs to be looked at is skin color.
Lighter-skinned women are seen as more attractive (Stephen et al, 2009). This is because it signals fertility, femininity, and youth. One more important thing it signals is the ability to carry a healthy child to term since lighter skin in women is associated with better vitamin D synthesis which is important for a growing babe.
Skin color and intelligence
There is a high negative correlation between skin color and intelligence, about –.92 (Templer and Arikawa, 2006). They used the data from Lynn and Vanhanen’s 2002 book IQ and the Wealth of Nations and found that there was an extremely strong negative correlation between skin color and IQ. However, data wasn’t collected for all countries tested and for half of the countries the IQs were ‘estimated’ from other surrounding countries’ IQs.
Jensen (2006) states that the main limitation in the study design of Arikawa and Templer (2006) is that “correlations obtained from this type of analysis are completely non-informative regarding any causal or functional connection between individual differences in skin pigmentation and individual differences in IQ, nor are they informative regarding the causal basis of the correlation, e.g., simple genetic association due to cross-assortative mating for skin color and IQ versus a pleiotropic correlation in which both of the phenotypically distinct but correlated traits are manifested by one and the same gene.”
Lynn (2002) purported to find a correlation of .14 in a representative sample of American blacks (n=430), concluding that the proportion of European genes in African Americans dictates how intelligent that individual black is. However, Hill (2002) showed that when controlling for childhood environmental factors such as SES, the correlation disappears and therefore, a genetic causality cannot be inferred from the data that Lynn (2002) used.
Since Lynn found a .14 correlation between skin color and IQ in black Americans, that means that only .0196 percent of the variation in IQ within black American adults can be explained by skin color. This is hardly anything to look at and keep in mind when thinking about racial differences in IQ.
However, other people have different ideas. Others may say that since animal studies find that lighter animals are less sexually active, are less aggressive, have a larger body mass, and greater stress resistance. So since this is seen in over 40 species of vertebrate, some fish species, and over 30 bird species (Rushton and Templer, 2012) that means that it should be a good predictor for human populations. Except it isn’t.
we know the genetic architecture of pigmentation. that is, we know all the genes (~10, usually less than 6 in pairwise between population comparisons). skin color varies via a small number of large effect trait loci. in contrast, I.Q. varies by a huge number of small effect loci. so logically the correlation is obviously just a correlation. to give you an example, SLC45A2 explains 25-40% of the variance between africans and europeans.
long story short: it’s stupid to keep repeating the correlation between skin color and I.Q. as if it’s a novel genetic story. it’s not. i hope don’t have to keep repeating this for too many years.
Finally, variation in skin color between human populations are primarily due to mutations on the genes MC1R, TYR, MATP (Graf, Hodgson, and Daal, 2005), and SLC24A5 (also see Lopez and Alonso, 2014 for a review of genes that account for skin color) so human populations aren’t “expected to consistently exhibit the associations between melanin-based coloration and the physiological and behavioural traits reported in our study” (Ducrest, Keller, and Roulin, 2008). Talking about just correlations is useless until causality is established (if it ever is).
The evolution of human skin variation is complex and is driven by more than one variable, but some are stronger than others. The evolution of black skin evolved—in part—due to skin cancer after we lost our fur. White skin evolved due to sexual selection (proximate cause) and to better absorb UV rays for vitamin D synthesis in colder climes (the true need for light skin in cold climates). Eurasians split around 40kya, and after this split both evolved light skin pigmentation independently. As I’ve shown, the alleles that code for skin color between blacks and whites don’t account for differences in aggression, nor do they account for differences in IQ. The genes that control skin color (about a dozen) pale in comparison to the genes that control intelligence (thousands of genes with small effects). Some other hypotheses for the evolution of white skin are on par with being as controversial as the hypothesis that skin color and intelligence co-evolved—mainly that mothers would kill darker-skinned babies because they weren’t seen as beautiful as lighter-skinned babies.
The evolution of human skin variation is extremely interesting with many competing hypotheses, however, to draw wild conclusions based on just correlations in regards to human skin color and intelligence and aggression, you’re going to need more evidence than just correlations.
Bang KM, Halder RM, White JE, Sampson CC, Wilson J. 1987. Skin cancer in black Americans: A review of 126 cases. J Natl Med Assoc 79:51–58
Beleza, S., Santos, A. M., Mcevoy, B., Alves, I., Martinho, C., Cameron, E., . . . Rocha, J. (2012). The Timing of Pigmentation Lightening in Europeans. Molecular Biology and Evolution,30(1), 24-35. doi:10.1093/molbev/mss207
Blum, H. F. (1961). Does the Melanin Pigment of Human Skin Have Adaptive Value?: An Essay in Human Ecology and the Evolution of Race. The Quarterly Review of Biology,36(1), 50-63. doi:10.1086/403275
Coetzee V, Faerber SJ, Greeff JM, Lefevre CE, Re DE, et al. (2012) African perceptions of female attractiveness. PLOS ONE 7: e48116.
Darwin, C. (1859). On the origin of species by means of natural selection, or, the preservation of favoured races in the struggle for life. London: J. Murray.
Darwin, C. (1871). The descent of man, and selection in relation to sex. London: John Murray, Albemarle Street.
Dixson, B. J., Dixson, A. F., Li, B., & Anderson, M. (2006). Studies of human physique and sexual attractiveness: Sexual preferences of men and women in China. American Journal of Human Biology,19(1), 88-95. doi:10.1002/ajhb.20584
Ducrest, A., Keller, L., & Roulin, A. (2008). Pleiotropy in the melanocortin system, coloration and behavioural syndromes. Trends in Ecology & Evolution,23(9), 502-510. doi:10.1016/j.tree.2008.06.001
Frost, P. (2007). Human skin-color sexual dimorphism: A test of the sexual selection hypothesis. American Journal of Physical Anthropology,133(1), 779-780. doi:10.1002/ajpa.20555
Graf, J., Hodgson, R., & Daal, A. V. (2005). Single nucleotide polymorphisms in theMATP gene are associated with normal human pigmentation variation. Human Mutation,25(3), 278-284. doi:10.1002/humu.20143
Greaves, M. (2014). Was skin cancer a selective force for black pigmentation in early hominin evolution? Proceedings of the Royal Society B: Biological Sciences,281(1781), 20132955-20132955. doi:10.1098/rspb.2013.2955
Gupta, D., Lammersfeld, C. A., Trukova, K., & Lis, C. G. (2009). Vitamin D and prostate cancer risk: a review of the epidemiological literature. Prostate Cancer and Prostatic Diseases,12(3), 215-226. doi:10.1038/pcan.2009.7
Guthrie RD. 1970. Evolution of human threat display organs. Evol Biol 4:257–302.
Harris, J. R. (2006). Parental selection: A third selection process in the evolution of human hairlessness and skin color. Medical Hypotheses,66(6), 1053-1059. doi:10.1016/j.mehy.2006.01.027
Harris, J. R. (2009). The nurture assumption: why children turn out the way they do. New York: Free Press.
Hill, Mark E. 2002. Skin color and intelligence in African Americans: A reanalysis of Lynn’s data. Population and Environment 24, no. 2:209–14
Holick, M. F. (2006). Resurrection of vitamin D deficiency and rickets. Journal of Clinical Investigation,116(8), 2062-2072. doi:10.1172/jci29449
Jablonski, N. G., & Chaplin, G. (2000). The evolution of human skin coloration. Journal of Human Evolution,39(1), 57-106. doi:10.1006/jhev.2000.0403
Jablonski, N. G., & Chaplin, G. (2014). Skin cancer was not a potent selective force in the evolution of protective pigmentation in early hominins. Proceedings of the Royal Society B: Biological Sciences,281(1789), 20140517-20140517. doi:10.1098/rspb.2014.0517
Jensen, A. R. (2006). Comments on correlations of IQ with skin color and geographic–demographic variables. Intelligence,34(2), 128-131. doi:10.1016/j.intell.2005.04.003
Kiang, L., & Takeuchi, D. T. (2009). Phenotypic Bias and Ethnic Identity in Filipino Americans. Social Science Quarterly,90(2), 428-445. doi:10.1111/j.1540-6237.2009.00625.x
Libon, F., Cavalier, E., & Nikkels, A. (2013). Skin Color Is Relevant to Vitamin D Synthesis. Dermatology,227(3), 250-254. doi:10.1159/000354750
Lieberman, D. E. (2015). Human Locomotion and Heat Loss: An Evolutionary Perspective. Comprehensive Physiology, 99-117. doi:10.1002/cphy.c140011
Lieberman, D. E., Raichlen, D. A., Pontzer, H., Bramble, D. M., & Cutright-Smith, E. (2006). The human gluteus maximus and its role in running. Journal of Experimental Biology,209(11), 2143-2155. doi:10.1242/jeb.02255
López, S., & Alonso, S. (2014). Evolution of Skin Pigmentation Differences in Humans. ELS. doi:10.1002/9780470015902.a0021001.pub2
Lynn, R. (2002). Skin color and intelligence in African Americans. Population and Environment, 23, 365–375.
Mosley, J. D., Appel, L. J., Ashour, Z., Coresh, J., Whelton, P. K., & Ibrahim, M. M. (2000). Relationship Between Skin Color and Blood Pressure in Egyptian Adults : Results From the National Hypertension Project. Hypertension,36(2), 296-302. doi:10.1161/01.hyp.36.2.296
Naidoo, L.; Khoza, N.; Dlova, N.C. A fairer face, a fairer tomorrow? A review of skin lighteners. Cosmetics 2016, 3, 33.
Norton, H. L., Kittles, R. A., Parra, E., Mckeigue, P., Mao, X., Cheng, K., . . . Shriver, M. D. (2006). Genetic Evidence for the Convergent Evolution of Light Skin in Europeans and East Asians. Molecular Biology and Evolution,24(3), 710-722. doi:10.1093/molbev/msl203
Rushton, J. P., & Templer, D. I. (2012). Do pigmentation and the melanocortin system modulate aggression and sexuality in humans as they do in other animals? Personality and Individual Differences,53(1), 4-8. doi:10.1016/j.paid.2012.02.015
Sankararaman, S., Mallick, S., Dannemann, M., Prüfer, K., Kelso, J., Pääbo, S., . . . Reich, D. (2014). The genomic landscape of Neanderthal ancestry in present-day humans. Nature,507(7492), 354-357. doi:10.1038/nature12961
Stephen, I. D., Smith, M. J., Stirrat, M. R., & Perrett, D. I. (2009). Facial Skin Coloration Affects Perceived Health of Human Faces. International Journal of Primatology,30(6), 845-857. doi:10.1007/s10764-009-9380-z
Stephens, D., & Thomas, T. L. (2012). The Influence of Skin Color on Heterosexual Black College Women’s Dating Beliefs. Journal of Feminist Family Therapy,24(4), 291-315. doi:10.1080/08952833.2012.710815
Sweet, E., Mcdade, T. W., Kiefe, C. I., & Liu, K. (2007). Relationships Between Skin Color, Income, and Blood Pressure Among African Americans in the CARDIA Study. American Journal of Public Health,97(12), 2253-2259. doi:10.2105/ajph.2006.088799
Tateno, Y., Komiyama, T., Katoh, T., Munkhbat, B., Oka, A., Haida, Y., . . . Inoko, H. (2014). Divergence of East Asians and Europeans Estimated Using Male- and Female-Specific Genetic Markers. Genome Biology and Evolution,6(3), 466-473. doi:10.1093/gbe/evu027
Templer, D. I., & Arikawa, H. (2006). Temperature, skin color, per capita income, and IQ: An international perspective. Intelligence,34(2), 121-139. doi:10.1016/j.intell.2005.04.002
Uzogara, E. E., Lee, H., Abdou, C. M., & Jackson, J. S. (2014). A comparison of skin tone discrimination among African American men: 1995 and 2003. Psychology of Men & Masculinity, 15(2), 201–212. http://doi.org/10.1037/a0033479
van den Berghe PL, Frost P. 1986. Skin color preference, sexual dimorphism and sexual selection: a case of gene-culture coevolution? Ethn Racial Stud 9:87–113.