Red Hair in Art: Thomas Cooper Gotch

Thomas Cooper Gotch (1854–1931) was an English painter and book illustrator loosely associated with the Pre-Raphaelite movement; he was the brother of John Alfred Gotch, the architect.

Gotch studied art in London and Antwerp before he married and studied in Paris with his wife, Caroline, a fellow artist. Returning to Britain, they settled into the Newlyn art colony in Cornwall. He first made paintings of natural, pastoral settings before immersing himself in the romantic, Pre-Raphaelite romantic style for which he is best known. His daughter Phyllis was often a model for the colourful depictions of young girls.

His works have been exhibited at the Royal Academy, Royal College of Art and the Paris Salon.

Self-portrait

The Heir to All Ages

Portrait of a Redhead

The Mother Enthroned

My Crown and My Sceptre

The Child Enthroned

Alleluia

A Pageant of Childhood

The Dancing Lesson

Red Hair in Art: Frank Dicksee

Sir Francis Bernard Dicksee (27 November 1853 – 17 October 1928) was an English Victorian painter and illustrator, best known for his pictures of dramatic literary, historical, and legendary scenes. He also was a noted painter of portraits of fashionable women, which helped to bring him success in his own time.

His father Thomas and his sister MArgaret were painters as well.

Red hair is featured in many of his paintings.

A Knight Being Crowned

Harmony

Chivalry

The Two Crowns

La belle dame sans merci

Passion

Stella

The Ideal or In Quest of Truth

Startled

Yseult

Red Hair in Art: Ferdinand Hodler

Ferdinand Hodler (March 14, 1853 – May 19, 1918) was a Swiss painter. He is one of the best-known Swiss painters of the nineteenth century. His early works were portraits, landscapes, and genre paintings in a realistic style. Later, he adopted a personal form of Symbolism which he called "parallelism".

Here are some of his painting featuring redheads.

Portrait of an Unknown

Portrait of Madame Loup

Spring

Spring

Head of a Red-haired Woman

Portrait of an Unknown Woman from Wien

Abendruhe

Portrait of James Vibert

Portrait of James Vibert

Portrait eines lesenden Mannes

Die Empfindung

The Dream. Cover for the novel Die Mittagsgöttin

Is red hair the result of sexual selection?

As we have seen in a previous post, in the study by Rosalind Harding and colleagues Evidence for Variable Selective Pressures at MC1R (2000) the authors argued that MC1R gene didn’t show any selective pressure.

The study by Peter Frost, published in 2006, European hair and eye color: a case of frequency-dependent sexual selection?  is a sort of reply to Harding’s paper. 

Frost writes that, without any selection, the current level of hair-colour diversity would have taken 850,000 years to develop, but modern humans have been in Europe for approximately 35,000 years according to Out of Africa II (for which I refer you to the final part of the post I linked above). So, Frost argues that there was indeed a selection: a sexual selection, which, however, acted primarily on women.

He proposes the concept of frequency-dependent sexual selection, according to which much rarer traits are more attractive and desirable and have a reproductive advantage as long as they remain rare. 

This process was stronger during the Upper Paleolithic and the last glacial period, when many men died or were absent for long periods due to long-distance hunting and extreme environmental conditions. This would have created fewer available men, less polygyny and a greater difficulty for women in finding a stable partner. In this situation, women had to compete for male partners, thus increasing the sexual selection’s pressure at more conspicuous feminine traits such as hair, eyes and skin pigmentation. Fairer traits would become a sort of visual signal, so women with these traits had an advantage in finding a partner. 

Europe during the last glacial period

This would also explain the rapid spreading of MC1R and its many mutations. Normally, more "silent" mutations (which do not change the phenotype) accumulate in genes than visible mutations. In MC1R, however, the opposite occurs: there are numerous variants that actually change hair colour. Frost interprets this as a sign that these variants have been favoured by selection (sexual selection, in this case) and not simply tolerated. The genetic structure of MC1R seems incompatible with simple random genetic drift and instead suggests a selective pressure that favoured visible hair diversity.

On the contrary, according to Frost, in many other traditional societies the situation was the opposite: thanks to a better climate, mortality rate among men was lower and they had to compete among them for women. As a consequence, sexual selection would affect men primarily. Furthermore, due to the very strong UV radiation, environmental natural selection tended to maintain more stable pigmentations.

This theory of sexual selection seems to be supported (at least in part) by other studies cited by Frost, according to which during adulthood blond hair darkens with age more slowly in women than in men, and that in all human populations women are paler than men after puberty. 

Besides, the 2008 paper Spectrophotometric Methods for Quantifying Pigmentation in Human Hair—Influence of MC1R Genotype and Environment also supports these findings, because the authors argue that hair colour varies more in women than in men and that redheads are more frequent among women. They don’t identify the genes causing this variation, but in his study Frost writes that, according to an unpublished work by Mather and colleagues, prenatal exposure to estrogen appears to be higher in individuals with blond hair or non-brown eyes. 

Many geneticists and evolutionists consider Frost’s hypothesis interesting, but not definitively proven. The main criticisms are: difficulty in empirically demonstrating prehistoric sexual preferences, speculative reconstruction of demographic relationships, possible underestimation of genetic drift and migration and risk of “adaptationism” (each trait must have a specific selective function).

Today, it is believed that skin, eye, and hair colour are likely the result of multiple combined factors, such as sexual selection, adaptation to UV rays and vitamin D, genetic drift, migrations and demographic history.

PS: as far as the last glacial period is concerned, in July 2025 a study was published which argued that even during the coldest ice ages the Arctic remained partially open, with seasonal sea ice allowing life to survive in the harshest climates. Here’s a short article and the original study. 

Do redheads synthesize vitamin D even without sun exposure?

The study Increased 25(OH)D3 level in redheaded people: Could redheadedness be an adaptation to temperate climate?  was conducted in 2020 in the Czech Republic on a sample of 203 individuals: 73 with red hair and 130 with different hair colours. Its aim was to investige the correlation between red hair and vitamin D, in particular the vitamin D precursor 25(OH)D3 (calcidiol) and folic acid.  

The results were quite interesting: 

Redheaded subjects had higher 25(OH)D3 concentrations and approximately the same folic acid concentrations as non-redheaded subjects. […] Redheaded subjects also reported that they used more intensive chemical and mechanical sun protection than their non-redheaded peers. In contrast to the situation in non-redheaded persons, redheaded persons' 25(OH)D3 concentrations seemed independent of the intensity of sun exposure or protection from solar radiation. […] This suggests that a factor other than eumelanin concentration (and skin fairness) is responsible for higher concentrations of 25(OH)D3 in redheaded individuals and that both traits, that is both red hair and fair skin, may well be two independent adaptations for life in environments with low UVB radiation.

 

Basically, they are saying that red-haired people are capable of synthesizing sufficient amounts of 25(OH)D3 even when their sun exposure is minimal. However, the authors warn that this observation cannot be generalised, and add:

This phenomenon was observed in two medium-sized samples of 93 men and 110 women who passed a relatively stringent self-selection process. Until this phenomenon is demonstrated in other, more representative populations, especially those living north of the 55° parallel, such as Scotland or Sweden, our conclusions must be considered merely preliminary.

In the years that followed, the study was not refuted, but it was not definitively confirmed either. There are several studies about skin pigmentation and vitamin D (like this one), but none of them focuses on the difference between red hair and other hair colours.

In any case, this Czech study is very interesting nonetheless, and we can verify these results by ourselves. If you have red hair and don’t expose yourself to the sun too much, you can have you level of vitamin D tested and see if you have deficiency of it or not. 

Also, should the study be correct, one wonders why red hair is not more common, at least in northern countries. If you have a hair colour that gives such an advantage, one expects for this hair colour to be selected by evolution over all the other hair colours. 

An original theory about the origin of red hair: boglands and nitrogen!

I recently came across an original theory about the origin of red hair, which links this hair colour to boglands. You can download the paper here. 

A bog (or bogland) is a wetland that accumulates peat as a deposit of dead plant materials, often mosses, typically sphagnum moss. They are often covered in heath or heather shrubs rooted in the sphagnum moss and peat. Bogs occur where the water at the ground surface is acidic and low in nutrients. They are generally found in cooler northern climates and are formed in poorly draining lake basins. In general, the low fertility and cool climate result in relatively slow plant growth, but decay is even slower due to low oxygen levels in saturated bog soils. Hence, peat accumulates. Large areas of the landscape can be covered many meters deep in peat. Bogs are widely distributed in cold, temperate climes, mostly in boreal ecosystems in the Northern Hemisphere. The world's largest wetland is the peat bogs of the Western Siberian Lowlands in Russia, which cover more than a million square kilometres.

A bog in Lauhanvuori National Park, Isojoki, Finland

The anaerobic environment and presence of tannic acids within bogs can result in the remarkable preservation of organic material. Finds of such material have been made in Slovenia, Denmark, Germany, Ireland, Russia, and the United Kingdom. They have yielded extremely well-preserved bog bodies, with hair, organs, and skin intact, buried there thousands of years ago after apparent Germanic and Celtic human sacrifice. Excellent examples of such human specimens include the Haraldskær Woman and Tollund Man in Denmark, and Lindow man found at Lindow Common in England (here a list of the most important bog bodies). The Tollund Man was so well preserved that when the body was discovered in 1950, the discoverers thought it was a recent murder victim and researchers were even able to tell the last meal that the Tollund Man ate before he died: porridge and fish. This process happens because of the low oxygen levels of bogs in combination with the high acidity.

As you can see in the photo below, the high levels of acidity darken the skin of these bodies, which often exhibit red hair. This led to the hypothesis that the high nitrogen level in boggy environments may be the cause of the red hair colour. The interaction between nitrogen and keratin in hair can lead to a chemical reaction that results in red pigmentation, regardless of the individual’s original hair colour.

Grauballe Man

I quote from the paper:

Building on these observations, this study posits that prolonged exposure to such environments could lead to genetic adaptations over generations. The hypothesis suggests that the nitrogen-rich conditions could exert selective pressure, favoring individuals with genetic mutations that produce red hair. These mutations might confer some survival advantages in boggy environments, possibly through mechanisms related to UV protection or thermoregulation. As a result, red hair could become a heritable trait within populations living in or around these regions.

[...]

In addition to direct genetic effects, nitrogen exposure may have also influenced epigenetic mechanisms. Epigenetics involves changes in gene expression that do not alter the DNA sequence but can be heritable. Prolonged exposure to high nitrogen levels could have triggered epigenetic modifications that enhanced the survival and reproductive success of individuals in these environments. Over time, these epigenetic changes could become stabilized in the population, contributing to the genetic adaptation observed today.

However, the author concludes by saying that the exact mechanisms and pathways of this process remain to be fully elucidated. 

Was red hair selected for fair skin? And how old are MC1R mutations?

In this post we are going to see a couple of points of the study Evidence for Variable Selective Pressures at MC1R, by Rosalind Harding et al.,

This study is from the year 2000, which means it’s now old, but it’s interesting nonetheless, because its conclusions were not entirely wrong.

In 2000, geneticists believed that MC1R mutations gave a major contribution to the evolution of fair skin in Europeans, and for that reason they expected to find a strong selective pressures at this gene. That is, they expected that human evolution selected MC1R because it was useful for people living in areas with low exposure to the sun.

So, they examined several European and non-European individuals. Europeans were from England, Ireland, Sweden, Finland, and Italy. Non-Europeans were from Africa, Japan, Papua New Guinea and southern India.

Here’s what they found out:

Although many of the MC1R amino acid variants observed in non-African populations do affect MC1R function and contribute to high levels of MC1R diversity in Europeans, we found no evidence, in either the magnitude or the patterns of diversity, for its enhancement by selection; rather, our analyses show that levels of MC1R polymorphism simply reflect neutral expectations under relaxation of strong functional constraint outside Africa.

Their models found no visible selection for MC1R. In particular, before conducting the study, the authors believed that the mutation Arg163Gln had an important role in altering MC1R, but didn’t find any evidence for that.

So, they came to the conclusion that probably MC1R had been the first gene to mutate (as far as human pigmentation is concerned), because there are many mutations that can alter this gene, but, after that, more effective mutations arised in other genes.

As a matter of fact, few years later this idea was confirmed by new discoveries.

Geneticists found out that genes SLC24A5 and SLC45A2 show a far stronger selection than MC1R and they are now considered the main contributors to skin variation in Europeans.

Origins of the SLC24A5 and SLC45A2 based on Archaeogenetics. Derived variants are said to have come from the Caucasus 28,000 years ago

Basically, MC1R underwent what geneticists call soft sweep selection, that is, not one main mutation that replaces all the others, but small mutations that grow all together. Not all of them are effective, though: some may be neutral, or very weak. On the contrary, a gene like SLC24A5 has a main mutation (A111T) showing a hard sweep selection, with a strong signal easy to detect in genomic analyses. The main mutation of SLC45A2 (L374F) behaves similarly.

The 2025 study The Genetics and Evolution of Human Pigmentation (Guermazi and Saliba) is about that too.

The authors write that in Eurasia selective pressure at MC1R had been weak, allowing for a great number of mutations, while in Europe a lighter skin pigmentation is due to genes SLC24A5 and SLC45A2 (in Asia, other genes are involved in skin pigmentation).

So, basically, MC1R is not the main driver for skin variations, but, unlike other genes involved in pigmentation, it shows a great variability.

To put it simple, there must be other reasons why red hair exists.

Let’s now come to the second part of the study,

First of all, the authors argue that both African and non-African data suggest that the time to the most recent common ancestor (TMRCA) of MC1R is ∼1 million years: that is, they are referring to the common ancestor of current MC1R mutations (see also Coalescent theory).

Second, the authors estimate the ages of MC1R alleles.

To estimate the ages of MC1R alleles, we assumed that their observed frequencies simply reflect genetic drift in constant-size, randomly mating populations, after allowing for different patterns of functional constraint.

Basically, researchers assumed that the size of ancient populations remained constant over time, but this may have biased the results given by the mathematical models.

Here are the estimated ages:

variants Val60Leu, Val92Met, and Arg163Gln: 250,000–100,000 years ago;

variants Arg151Cys and Arg160Trp: 80,000 ago;

African silent variants Leu106Leu, Cys273Cys, and Phe300Phe: 110,000–40,000 years ago;

variants Asp294His and Ser316Ser: ⩽30,000 years.

I quote from the study:

However, an incompatibility arises between estimated ages in the range of 250,000–100,000 years, for non-African MC1R allelic variation, and ages, from fossil evidence, of ⩽100,000 years for the dispersal of modern humans outside Africa and the Middle East (Stringer and Andrews 1988). One possible explanation for incompatible ages from genetic data is that they have been overestimated under an assumption of levels of genetic drift that are consistent with constant population size. These age estimates would be younger under an alternative model with a high rate of population expansion out of a Eurasian founding population.

That is, they argue that, since, according to the theory Out of Africa II, the dispersal out of Africa took place around 80/70,000 years ago, these ages are too old, so to speak, and this probably happened because they assumed a constant-size population.

In subsequent years, other studies have been released that have revised these estimates downwards. This time they used a different demographic model, involving so-called bottlenecks and a rapid growth just after the dispersal out of Africa (although I don’t understand why a population should grow rapidly just after the dispersal out of Africa).

With this new model, the estimates for the first MC1R mutations lowered to 20,000/15,000 years ago.

However, it is important to notice that these new studies are not focused on the MC1R gene and its mutations (like the 2000 study by Harding et al), but rather on multiple genes involved in human pigmentation, and on ancient DNA in general (see, for example, The Timing of Pigmentation Lightening in Europeans, by Beleza et al, 2012).

Besides, since MC1R has got many mutations, dating all of them can be difficult. In any case, since MC1R too is involved in pigmentation, a study like the one by Beleza et al. can be useful to understand MC1R as well.

As for the estimate of 1 million years for the common ancestor of current mutations, modern studies prefer avoiding such a precise date (1 million years), in favour of a more nuanced estimate (500,000/1 million), but they do recognise that MC1R alleles are more ancient than other genes involved in human pigmentation.



Obviously, all the dates we have seen so far shouldn’t be taken as the gospel, and not only because (as we have seen in a previous article) a complete genomic analysis of ancient remains is often impossible.

As far as archaeogenetics and archaeology are concerned, the theory of Out of Africa II has become a sort of dogma. So, should evidence emerge that disproves the OOA (like MC1R mutations dating back to 250,000 years ago), the evidence is considered wrong and the OOA is never questioned.


Below, you’ll find a book and three articles about this dogma.


Not Out Of Africa, by Mary Lefkowitz (free download here)


African Eve: Hoax or Hypothesis? This article is a bit technical in certain points and features a lot of citations, but if you have the patience to read throughit, you’ll see it is very informative.


The two articles below are a bit technical too.

Re-Examiningthe “Out of Africa” Theory and the Origin of Europeoids(Caucasoids) in Light of DNA Genealogy


Re-Examining the Out-of-Africa Theory and the Origin of Europeoids (Caucasoids). Part 2. SNPs, Haplogroups and Haplotypes in the Y-Chromosome of Chimpanzee and Humans