Beauty in Nature.
Desire, Evolution, and Sexual Selection
Mats Dahllöv
Differens Magazine, autumn 24
The question of beauty and nature is one of the more normative and ideologically laden. But being in general an interest from conservative and reactionary viewpoints, it is also one of surprising perspectives. This text will focus on a perspective I first met in the continental feminist philosopher Elisabeth Grosz’ Becoming Undone. Darwinian Reflections on Life, Politics, and Art (2011), that I later found more expanded and scientifically thoroughgoing in the ornithologist Richard O. Prum’s The Evolution of Beauty. How Darwin’s Forgotten Theory of Mate Choice Shapes the Animal World – and Us (2017), and, in writing this article, in the evolutionary psychologist Geoffrey Miller’s The Mating Mind. How Sexual Choice Shaped the Evolution of Human Nature (2000). A perspective that for me presented a radically new and different view of nature and evolution – one in which desire, beauty, and the artistic plays an essential role. It all goes back to Darwin’s so-called ”second idea,” his theory of sexual selection. A theory that when first presented was fiercely contested for its radicality, for a hundred years forgotten, still today met with skepticism, and to the public little known compared to his theory of natural selection and the adaptationist formula ”survival of the fittest.” From these two theories, different views of nature have been extrapolated that can be expressed as the difference between mechanism, reductionism, and the supposedly rational on the one hand, creativity, abundance, and the allegedly irrational on the other. The latter has all to do with desire, beauty, and the artistic.
This article will begin with Darwin’s formulation of his theory of sexual selection and examine how sexual selection can account for beauty and artistry in some bird species. It will thereafter deal with the historical and contemporary reductionist critique of Darwin’s theory, before exploring how other aspects of species, relating to but sometimes moving beyond the question of beauty, can be explained from the perspective of sexual selection. We will here also encounter what can be seen as the opposite of beauty – violence – and explore in what way evolution can be seen as a struggle between the sexes over autonomy and reproductive choice, before ending up with our own species to see how a desire for pleasure and artistry (more than mere beauty) can account of many of the peculiarities of our species. I will end with some reflections on how the idea of beauty in nature put forth in this article can be understood in relation to Kant’s aesthetics.
Kris Lemsalu, Fine Sin, 2022 Peace at 295 Church, 2022, Margot Samel Gallery, photo by Dawn Blackman.
If Kant’s philosophy has been criticised for being sexless and supposedly gender neutral, this article will be far from it. We will be mostly concerned with birds – Kant’s favourite example in his aesthetics – but we will also venture not only into sexed reproductive strategies but also into the looks and particularities of penises and vaginas. And although this article will present ideas that could be characterised as ”progressive,” and according to two of the authors mentioned above even ”queer” or ”feminist,” evolutionary biology is a delicate field, and some words of caution might be in place. First, it should be underlined that these are some theories on evolution, beauty, sex, and nature, with both merits and flaws, and in no way intended to give some sort of exhaustive or final answer (and considering our species, culture, in all its possible varieties, is an essential part of our nature). If, to follow the early German Romantics, nature and the human being is to be seen as a work of art, then one should perhaps also apply the principle of the artwork’s inexhaustibility: no interpretation can exhaust it, and seemingly conflicting interpretations can each have their truth.
Secondly, the focus of sexual selection and sexual reproduction is a focus on males and females and their respective mate choices and reproductive strategies. It is therefore worth to emphasise nature’s sexual variety, including both hermaphrodites and species with more than two sexes, and, not least, same-sex sexual behaviour being found en masse – as of 2019 recorded in over 1500 species, a number that will most likely grow. If one wants to seek an evolutionary explanation to what has been termed a ”Darwinian paradox,” one could in fact look to mate choice favouring same-sex sexual behaviour (which is what Prum do) or find it in social evolutionary selection favouring group affinity. However, the search for an explanation might be made from a false premise. Given the widespread occurrence of same-sex sexual behaviour across various branches of the tree of life, it could be that indiscriminate sexual behaviour directed towards all sexes is the evolutionary basis, with exclusive different-sex sexual behaviour being an evolved trait in need of evolutionary explanation.
A last introductory point I would like to make concerns the merits of turning to the natural sciences. Here I agree with Grosz who argues that critical thinking, and specifically feminist philosophy in her case, has all to win in turning to nature and ”the relentless force of the real” and develop a ”new metaphysics.” Grosz, polemically, contrasts this with postmodern feminism, which has been preoccupied with representations of the real and, as a result, has neglected discussions about matter and nature. With new materialism and posthumanism, this is no longer the case. However, I think it is safe to say that posthumanism has been more interested in bacteria, fungi, and co-evolution than evolutionary biology and sexual selection. Though radical in many ways, one could argue that it is a ”safer” way to decentre the human and underline our embeddedness in nature than to look at our animal relatives. Yes, the human is entangled with bacteria and viruses, but we are also primates formed by, among other things, sexual selection. I would also argue that the shunning and critique of evolutionary biology on the basis of a reductionist and determinist – and as we will see anti-Darwinian – understanding of it, can also work as a confirmation of it since no alternative is presented. With this said, let us turn to one of the most powerful and interesting theories of an abundant and non-deterministic nature.
Sexual Selection and Beauty
Charles Darwin might seem an unexpected ally in thinking nature and beauty anew. But ever since the publication of The Descent of Man, and Selection in Relation to Sex in 1871, Darwin’s theory of sexual selection and the importance it accords to beauty, desire and what is mostly female mate choice has been fiercely suppressed among evolutionary biologists – in the name of a ”pure Darwinism” – and is largely unknown to the general public. Yet the theory of sexual selection was something that occupied his thinking more than the one of natural selection, and it was a radical theory entirely unique to Darwin.
Before entering the abundant world of sexual selection, I will briefly present its ”competitor”: natural selection. As we will see concerning sexual selection, birds make out a wonderful example and so also here. At the age of 22, Darwin set out for what would be a five year long and crucial expedition on the navy ship HMS Beagle. Passing the Galápagos Islands, Darwin collected several specimens of different finches – they would turn out to be important in the elaboration of his theory of natural selection. Darwin recognised the finches’ affinity to birds of the nearest mainland, though there were important differences: their beaks. It seemed, therefore, that the finches from the different islands had a common ancestor but that their beaks had evolved in different directions according to the environment of the islands. Later, this has been termed adaptive radiation: the beaks have evolved to allow the finches to eat different types of food, thus creating new species. The finches are therefore an excellent example of natural selection: you either eat or you die. Note that this evolutionary process is without any aspect of choice and that it leads to a functional result in the form of beaks adapted to eat different types of food. The birds of Galápagos now go by the name ”Darwin’s finches.”
”The sight of a feather in a peacock’s tail, whenever I gaze at it, makes me sick!” So wrote Darwin in a letter in 1860, the year after the publication of one of the most famous books in the history of science, On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. Darwin’s unpleasant feelings in front of the peacock’s tail was not due to a peculiar taste in front of something generally regarded as an expression of beauty in nature; the problem was precisely how to account for the development of the peacock’s tail in relation to the theory of natural selection, often described as the ”survival of the fittest.” From the viewpoint of natural selection and survival, the peacock male’s elaborate tail seems instead unfit: the brilliance of the tail making it more visible to predators and its length a hinder to a quick escape.
So, if it does not have any survival value, why had it developed and what could be its advantage? The answer: its beauty. ”Many female progenitors of the peacock,” Darwin writes in The Descent of Man, and Selection in Relation to Sex, ”must during a long line of descent, have appreciated this superiority [in beauty]; for they have unconsciously, by the continued preference of the most beautiful males, rendered the peacock the most splendid of living birds.” Where the beak of the different Galápagos finches had evolved in adjustment to different types of food, the male peacock had developed its tail because female peacocks had chosen to reproduce with what they aesthetically desired, something that through evolution has led to the extraordinary appearance of the male peacock.
Darwin, however, does not merely describe this as a process of unconscious preference. More often, he explains it as a sophisticated judgement of taste. In the case of the Argus pheasant, whose elaborate feathers are only displayed during the mating dance, Darwin not only sees a ”good evidence that the most refined beauty may serve as a sexual charm, and for no other purpose” (my italics). He also describes the mating choice: ”Many,” Darwin writes, ”will declare that it is utterly incredible that a female bird should be able to appreciate fine shading and exquisite patterns. It is undoubtedly a marvellous fact that she should possess this almost human degree of taste.”
At the heart of the theory of sexual selection therefore stands desire and subjective judgements. An understanding of animals that is far from the idea that they are mere machines; animals are subjects with mental capacities that enable them to make judgments and choices. Furthermore, it is only by granting them these mental capacities and seeing their choices as subjective that we can understand nature and evolution. As Richard O. Prum puts it: if natural selection is described as a relentless and impersonal force that, as in the case of the ”Darwin’s finches,” produces functional design – a blind watchmaker with Richard Dawkins’ metaphor – in sexual selection, nature is not blind but has in its myriads of organisms developed its own eyes and ears and the mental capacities to make decisions based on this sensory information, producing ”unfunctional” design, that is beauty.
It is not only the visual beauty of birds that has been taken to what in some cases could be called, in Darwin’s words, ”a wonderful extreme.” There is of course bird song, an ”art,” as Darwin calls it, that have developed so elaborately out of the same desire for beauty and pleasure. The ornithologist and composer Olivier Messiaen – who had a great influence on 20th-century music, especially on the French serialists and Pierre Boulez – took a great interest in birdsong, culminating musically in his huge piano work Catalogue des oiseaux (”Catalogue of birds”). As Messiaen notes, the rivalry over territory – crucial for mating – is sometimes settled by song duels: ”if the predator wants to occupy a place that does not belong to him, the true owner sings, sings so well that the other one leaves […] [but] if the thief sings better, the owner gives him his place.”
An even more peculiar art is found among the bowerbirds. Here, beauty is expressed in elaborate and, across different species with their particular beauty norms, varied architectural and sculptural creations. There are two general types: the avenue bower and the maypole bower. In the former case the male build two walls of vertically placed sticks. In the latter, sticks are places around a support creating a pole that can rise several feet. The court, placed at some distance, is then decorated with hundreds of colourful objects of different sorts – shells, flowers, berries, even plastic items and coins – on which the male performs his courting, consisting of display and singing. It is furthermore one of the bower birds, the Tooth-billed bowerbird, that is Gilles Deleuze and Félix Guattari’s favourite example in their (post-humanistic) theory on the origin of art. The Tooth-billed bowerbird does not build a bower but only a ”stage,” consisting of leaves beneath the branch it sits upon and on which it performs its courtship consisting of advanced songs that include mimicry of other bird species. ”[U]n artiste complet,” ”une oeuvre d’art totale,” as they write. It is doubtful that Deleuze and Guattari knew of Darwin’s theory of sexual selection, but if they would, in it they could also have found an evolutionary theory in line with their idea of desire as productive rather than as feeling a lack: behind the ”complete artist” stands the female bowerbird’s desire for such courtship artistry.
As if these diverse forms of artistic expression were not enough, recent research suggests that the evolutionary development of wings – the defining feature of birds – may in fact have its cause in sexual selection. Long before wings for flying had evolved, feathers were enjoyed for their ability to be expressive. In what can be seen as a parallel to the discovery that the ancient Greek sculptures were in fact coloured, thus confounding later times idealisation of antiquity, a recent discovery is that most dinosaurs bore feathers. As with the coloured sculptures, feathered dinosaurs give quite a different feel – just imagine a colourfully feathered Tyrannosaurus Rex in the 1990s classic Jurassic Park. Prum was part of the research team that in 2010 developed the first representation of what this might have looked like in the case of the Anchiornis huxleyi. The hypothesis is that the development of more advanced forms of feathers has to do with their ability to be more expressive. As Prum writes about the fourth and final stage: “the planar vane of the feather might have evolved through aesthetic selection to create a two-dimensional canvas upon which to depict complex pigment patterns—including stripes, spots, dots, and spangles.” He concludes: “the potentially aesthetic innovation of planar feathers facilitated the evolution of flight and the avian dinosaur survival of the Cretaceous-Paleogene extinction. It’s harder to imagine a bigger possible impact for the role of beauty and desire in the history of life.”
What Darwin calls ”the taste for the beautiful” is however not the only principle of sexual selection in Darwin’s theory. There is also what he calls the ”law of battle”: the struggle between individuals of one sex, most often males, for getting the dominant position and the reproductive control over the individuals of the other sex. This process leads to the development of large body size and weapons of aggression, such as horns and antlers. The difference between these two principles of sexual selection can thus be boiled down to: ornaments or armaments. Another way to conceptualise the difference – and this I will come back to – is between autonomous choice and forced selection.
It might perhaps seem as if we are closer to the ”survival of the fittest” regarding the second form of sexual selection, that is the ”law of battle,” but it is still not about survival but about reproduction. To stretch the difference between survival and reproduction to the extreme, the only relation between them is that an animal must survive until it is sexually mature and able to reproduce; this is the only ”interest” in survival from the strict viewpoint of reproduction. It is also to be noted that reproduction is the bottom-line of evolution. An individual organism can be excellent in surviving, it will still sooner or later die, and if it has not reproduced, the excellent survival genes will die out as well. In reality, there is of course all sorts of possible overlaps between reproduction and survival, and sexually attractive traits may have their origin in them being also valuable for survival. This is what Ronald A. Fisher’s significant theory on runaway selection addresses: a trait that initially is preferred for some adaptive reason becomes unhinged and desired for its own sake. The ”runaway” aspect has to do with the fact that both the desire for the trait and the trait itself is inherited and it thus creates a positive feedback-loop that can drive ornaments to extremes, as in the case of the Peacock tail.
Kris Lemsalu, 2014, Aldonzo Lorenzo, Courtesy of the artist and Temnikova & Kasela Gallery.
Adaptationist and reductionist critique
Darwin’s theory of sexual selection was not well received, especially not the importance Darwin accorded to female mate choice. In a review by the biologist St. George Mivart, the law of battle was accepted but Mivart argued that it should be seen as a branch of natural selection. Concerning the ”taste for the beautiful,” Alfred Russel Wallace, famous for discovering the principle of natural selection independently of Darwin, formulated the view that has since been the leading one: ”The only way in which we can account for the observed facts is by supposing that colour and ornament are strictly correlated with health, vigor, and general fitness to survive.” And as Wallace continued: ”In rejecting that phase of sexual selection depending on female choice, I insist on the greater efficacy of natural selection. This is pre-eminently the Darwinian doctrine, and I therefore claim […] the position of being the advocate of pure Darwinism.” To be a true Darwinian then meant to oppose one of Darwin’s two theories of evolution, and precisely the one that took up most of his interest, surely as Darwin knew it was the most controversial and radically new. Wallace’s reasons were also of a moral and prejudiced nature, not wanting to grant the ”brutes” of the animal kingdom the mental powers that Darwin’s theory of sexual selection accorded them. As Geoffrey Miller writes: ”This psychologizing of evolution was Darwin’s greatest heresy. It was one thing for a generalized Nature to replace God as the creative force. It was much more radical to replace an omniscient Creator with the pebble-sized brains of lower animals lusting after one another.”
With some few exceptions, the theory of sexual selection was then absent from evolutionary biology for a hundred years. And when scientists revived the theory of sexual selection in the 1970s, it was Wallace’s view that ornaments are correlated to fitness that returned, now expressed as ”good genes” and perhaps best known in the form of the ”handicap principle”. This principle, proposed by Amotz Zahavi in 1975, argues that ornamentation signals not beauty but burden, that is, a handicap. An individual communicates that it is able to survive in spite of the handicap. In short, the greater the handicap, the better the genes. Zahavi’s idea and its later variants have proven very popular, despite having been contested for its logic, and despite decades of empirical research failing to prove it. Similarly, while Geoffrey Miller’s theory of the evolution of human nature is both mind-opening and convincing, he tends to view everything – from bird songs and extravagant plumage to marriage rituals and human beauty and art – as ultimately fitness indicators and expressions of good genes.
The reason for the popularity of the handicap principle is perhaps best understood in the reasoning of one of its principal developers, Alan Grafen, who favours its ”rhyme and reason” over the arbitrariness of aesthetic mate choice, an explanation he finds ”methodologically wicked.” To return to the opening paragraph of this article, I would therefore argue that it is a matter of worldview: the processes of nature must be rational and evolution strictly about adaptation. Richard Dawkins, the most famous evolutionary biologist of our time and an outspoken neo-Wallacean, endorses the handicap principle, and in the second edition of his reductionist classic The Selfish Gene (1976, 1989) he likens the female to a ”good diagnostic doctor” who chooses only the healthiest male as a mate. In one of the examples Dawkins is discussing – to which I will return – he adds that his theory of the diagnostic female doctor is ”less plausible than pleasing,” that is Dawkins finds his own reasoning implausible, but likes it because of its consistency with the doctrine of adaptation. Dawkins’ comment is, as Prum remarks, revealing for the whole research field on adaptive mate choice in its quest to find an adaptive explanation even when there is nothing to support it.
The resistance to Darwin’s theory of sexual selection thus has much to do with a predilection for rationality and reductionism, evident in Wallace’s expression ”pure Darwinism,” with adaptation as the single and global principle capable of explaining everything in nature. To accept the theory of sexual selection and the place of desire and beauty, is to accept an evolution that, in the words of Prum, is “far quirkier, stranger, more historically contingent, individualised, and less predictable and generalizable than adaptation can explain.” Grosz, for her part, describes sexual selection as ”the queering of natural selection”: ”that is, the rendering of any biological norms, ideals of fitness, strange, incalculable, excessive. Sexual selection, as an alternative principle to natural selection, expands the world of the living into the nonfunctional, the redundant, the artistic.” Sexual selection is, Grosz says with a nod to Gilles Deleuze, a ”difference machine.”
Kris Lemsalu, 2008 – 2022, photos by Katharina Reckendorfer, Josef Schauer-Schmidinger and Aadam Kaarma.
Mate Choice Expanded: Sociality, Female Autonomy, Sexual Pleasure, and the Creative Mind
To see how powerful the principle of sexual selection is in creating the richness and diversity of nature, we will move beyond mere beauty. Beauty may be the most striking example, but mate choice is ubiquitous – Darwin devotes considerable space to insects, fishes, reptiles, amphibians (leaving out plants however) – and can also be about favouring a particular behaviour, with enormous consequences for evolution and the development of new species. It can furthermore, to stick with pleasure but moving beyond aesthetic pleasure, be about the actual ”sex” in sexual selection, and it may be the major principle behind the development of our own essentially artistic species.
We will however stick with beauty and birds to make a bridge between aesthetic preferences and see how these can go hand in hand with the development of certain behaviours. If the peacock, the Argus pheasant, and the bowerbirds are prime examples of aesthetic evolution and beauty in nature, manakin birds are another. Although the 55 different species of manakin are also visually striking, their most characteristic feature is their advanced and varied courtship rituals that includes acrobatic flights, jumps, bowing, and wing vibration. Simply put: highly choreographed dances and show numbers. In certain manakin species, this has led to a radical change in male social behaviour as female manakins have developed a taste for coordinated group shows. The result? Male manakins are, in order to create the most impressive choreography, highly social, with daily social interactions and relationships with other males that can last for a decade or more. Adult females manakins, on the other hand, lead completely independent lives, their only social interaction being the few brief minutes a year when they visit the artistic shows, choose a mate, and copulate with him. In the Chiroxiphia genera, this entails that 90 percent of the males never get to reproduce, as the female chooses one of the alpha males in the group whose performance she prefers. Who then becomes the chosen male? Well, it is not about strength or domination. Studies on the long-tailed manakin show that the best indicator of a young male’s future reproductive success is his ability to establish social relationships, as this is what enables him to be the star in the best artistic group performance.
So how to understand this from the viewpoint of reproduction? In all the species discussed so far, the female chooses the paternity of the offspring out of her ”taste for the beautiful.” It is therefore – and this is one of Prum’s concerns – also a question of female autonomy. Sexual selection is, naturally, about sexual reproduction. This, however, does not require different sexes or sexual differences, though this is the case in most animals. On the most basic level, sexual difference is about the size of the gamete: the female sex evolved to make large packets, the egg, in which their DNA comes with nutrients, the male sex to make small almost naked package with their DNA, the sperm. The fundamental difference thus lies in the amount of energy contributed to the offspring. In mammals, this difference in ”parental investment” is further increased by pregnancy, milk production, and in most species care of the offspring. The crudest expression of this in relation to reproductive strategy is that males, especially when the investment is minimised to copulation, strive after quantity of females, and females after quality of males, which means that one male mates with several females and some males do not get to mate at all (with the Chiroxiphia genera of manakin birds being an extreme example). This is the reason why we see a general sex difference in the animal kingdom where males court and females choose, but also why there can be an evolutionary struggle between male control and female autonomy. Like Darwin, the great empiricist, we should however take note of species where the roles of the sexes are reversed, seen both regarding the law of combat, where females of some species are larger than the males ”for the sake of conquering other females and obtaining possession of the males,” and regarding the taste for the beautiful, where females are ”the more eager in courtship, the males remaining comparatively passive, but apparently selecting the more attractive females.” And there is of course species, such as ours, in which both sexes practice mate choice and courtship.
But there also exists a rather dark story of male coercion, or outright rape, versus female struggle for autonomous mate choice. This has been taken to extremes in the case of some species of waterfowl. Prum and colleagues have studied the anatomical evolution of the duck penises and vaginas which in some species show a kind of arms race between males and females ”in which each sex evolves successive behavioral, morphological, or even biochemical mechanisms to overcome the evolved efforts by the other sex to assert control or freedom of choice over reproduction.” Among the duck species, one finds the longest penises in relation to body size in the entire animal kingdom, and in the case of the mallard duck, the male has developed a long, corkscrew-shaped penis, while the female mallard duck has co-evolved a vagina with ”dead ends” and several clockwise spirals to reduce the chances of fertilisation when she has not chosen her partner freely but has been forced to copulation. But to call it a ”war of the sexes” is misleading, as Prum argues, because it is highly asymmetrical: ”Males evolve weapons of control, while females are merely coevolving defenses that create opportunity for choice. It’s not a fair fight, because only males are really at war.”
Is there then an evolutionary conflict between beauty and sexual violence? Returning to the bowerbirds, this seems in fact to be the case. In 1995, Gerald Borgia presented the risk reduction hypothesis, later experimentally verified. As the subtitle of his article ”Why Do Bowerbirds Build Bowers?” summarises it: ”Females prefer to visit courtship areas that provide easy avenues of escape, thereby protecting them from forced copulations.” The bowers thus works as protection that allows the females to safely exercise their aesthetic judgements, where research furthermore shows that females prefers to mate with the male with the most intense and aggressive courtship show. But enjoyed in safety and chosen freely. It can be understood as a case of, as Prum aptly calls his chapter on the bowerbirds, ”beauty from the beast.”
Forced copulation, an extreme form of the ”law of the battle,” should therefore be understood as a selfish male evolutionary strategy in conflict with the evolutionary interests of its female victims and the principle of mate choice and the ”taste for the beautiful.” It is, to use my distinction from above, a case of forced selection over autonomous mate choice. In some animals, this selfish male strategy to take forced control over reproduction and to spread the individual’s own genes is not only directed at females through forced copulation. In the case of many primates it gets even more sinister: infanticide. When a new male gains control of a group, he kills the lactating offspring of the group’s females to create reproductive opportunities for himself. In gorillas, it is estimated that around a third of infant mortality is due to reproductive motivated killings, and this is also well documented in chimpanzees.
With the bonobo – our closest relative along with the chimpanzee – things are notably different. Their societies have a matriarchal or at least co-dominant structure; infanticide is absent; and sex is never coerced, and serves far more purposes than just reproduction. Bonobos engage in sex to mediate conflicts, reduce social tensions, and reconcile—regardless of the individual’s sex, age, or social status. In one key respect, however, these different primates are similar: females take full responsibility for the offspring.
Turning to our own primate species, we share some important aspects with the bonobos: reproductively motivated infanticide is non-existent, forced sex exists but is far from being the norm, and sex is much more than just reproductive. However, humans differ from bonobos in two important ways: males invest in the care of the offspring and males also practice mate choice; the two are most likely interrelated. Following Prum – who, if it has not been clear, has a feminist viewpoint in his theorising on evolution and mate choice – he argues that our ancestral hominid females succeeded in what he calls an ‘aesthetic de-weaponization’ of males. This can be seen in the loss of elongated, razor-sharp canine teeth and the relatively small difference in body size between males and females. Once this was achieved, females used mate choice to expand their autonomy and drive a broader social transformation, turning males into fathers. To understand it in terms of autonomy might perhaps be to stretch it, but that male childcare was an aspect in mating success – that is, hominid males that practiced childcare were more often repeatedly chosen by females – is more than likely. Thus, not only did male care for the offspring increase the chances of the offspring to survive to a reproductive age, thereby spreading those genes, but it was also a reason of male mating success. And one thing is clear: male care of offspring is an essential part in the hominid evolution that led to homo sapiens.
That hominid males made substantial investments for the care of the offspring goes hand in hand with male mate choice, unknown among our primate relatives that never misses a fertile sexual opportunity. The perhaps most conspicuous ornament evolved out of the hominid male ”taste for the beautiful” is permanent breast, a unique human visual ornament that does not exist among any of the worlds other 5000 mammals. Natural selection obviously cannot account for this; breastfeeding among other mammals works great. And to just clear out some well-spread adaptationist ideas about human beauty: That the female body form in its individual differences would be related to fertility and ”reproductive value” – most well-known is the waist-to-hip ratio – is a theory not backed by any convincing data, despite massive research in search of it. The same applies to the theory regarding a correspondence between facial symmetry and good genes. And just to state it out clearly: beauty norms are diversified, plural and cultural. Returning to Darwin – a forceful opponent of slavery – sexual selection, grounded in arbitrary preferences for some physical traits, has led to the diverse appearances of the human ”races.” The reason for this diversity does not lie in adaptation or the survival of the fittest. Although human beauty does not display the same level of extravagance as that of the Argus pheasant, it remains an evolved trait enjoyed for its own sake.
Another human peculiarity is the place of sexual pleasure, an all-encompassing sensory experience rather than only visual. Both Prum and Miller argue that the striving for increased sexual pleasure has been a major driving force in human evolution, both on an anatomical and mental level, and primarily driven by female mate choice. The length of human copulation is unmatched, even the promiscuous bonobos only have an average of 13 seconds, and it can hardly be the result of natural selection. Instead, a preference for sexual pleasure has led to an evolutionary development, ”an aesthetic, coevolutionary lovefest” in the words of Prum. The best visible evidence for this is the human male penis: in its length, thickness, and flexibility it is without comparison among other primates. However, theories have been made to also explain this from a reductionist viewpoint. And here is the place to return to Dawkins’ idea of the female as a ”good diagnostic doctor,” because it precisely regards the penis. Together with the spider monkey, the human male penis is unlike all other primates’ as it lacks a bone, the baculum. The reason for this, Dawkins finds in the ability for the female to discern if the male has good genes. The lack of a bone is a handicap. Shortly, the greater the boneless boner the greater the genes: ”only genuinely healthy or strong males could present a really stiff erection.” Problem is that there is no truth in it. Not only is Dawkins’ idea a good example of how far the search of an adaptive explanation might go, but it is also, as Prum puts it, ”a masterwork of phallocentric evolutionary biology.” In general, adaptationist evolutionary biology does not only have difficulties with beauty but also sexual pleasure. Several theories have also been launched to explain the seemingly unnecessary female orgasm: as an evolutionary byproduct (the male orgasm is needed for fertilisation and the female has gotten it as by a happy accident), as increasing the chances of fertilisation (the so-called ”upsuck hypothesis”). None of them have gathered any empirical support and, as both Prum and Miller shows, they just do not make sense.
The true Darwinian answer is instead to see our sexual organs, and the foremost pleasure organ, the clitoris, not the least, as evolved through mate choice with a preference for sexual pleasure. Just as beauty can be an evolutionary driving force and valued for its own sake, so can sexual pleasure. The fact that our species has concealed ovulation goes hand in hand with this: as individual acts of sexual intercourse have a low probability of leading to fertilisation – in an average couple it takes about three months of regular sex – Prum suggest that we should better think of humans as having remating preferences, where sexual pleasure, or sexual ”compatibility” with a common expression, is precisely one major aspect of remate choice.
Sexual pleasure notwithstanding, the most characteristic feature of homo sapiens is the mind. But what if the mind actually has to do with pleasure, as in the ability to charm a potential mate through humour, imagination, storytelling, art, music, ornate language? This is what Geoffrey Miller convincingly argues for in the book The Mating Mind. How Sexual Choice Shaped the Evolution of Human Nature. The human mind would then be the equivalent to the peacock’s tail. This is not to say that looks are not important among humans, which would be a standpoint far from the reality we all live in. Looks are however only the first part of attraction, then comes charm, sociability, humour, intelligence, and so on, that is, the mind. The importance of the latter is even more evident regarding remating and pair bonding, where most actual reproduction happens, and it is thus here the real (reproductive) mate choice is made.
The crucial part is that the human mind stands out in its extravagancy, just like the peacock’s tail. To find an explanation to the rapid development of the mind and its abilities – with brain size tripling in the evolutionary mind-boggling speed of just two million years – has been a difficult scientific nut to crack for adaptationist science. In evolutionary theory, the best way to explain rapid development of something radically new, is the positive-feedback process that distinguishes runaway sexual selection. For an arbitrary reason – and sexual selection is always arbitrary – our ancestors developed a desire for a creative mind, just as the peacock female developed a desire for extravagant plumage.
With humanity dominating the world in a geological epoch we have named after us, the Anthropocene, it might be hard to imagine that the mind for the great majority of time since the modern homo sapiens first came into existence 100 000 years ago did not have any particular survival value. The major event that occurred around 12 000 years ago, which made the mind’s abilities useful from a survival perspective, was the advent agriculture and civilisation. Evolution, however, does not work with such foresight. From an adaptive perspective, every step leading to a complex innovation such as the human brain – which moreover accounts for 20% of the body’s energy consumption – needs some survival advantage. Traits that develop out of sexual selection do not, all they must do is to bring a reproductive advantage. So, rather than an adaptively developed problem-solver, the human mind should be viewed as a sexually selected amusement park, Miller argues. The mind is ornamental, abundant, and creative.
A phenomenon such as art is in this scenario therefore not a strange and unnecessary byproduct whose evolutionary and adaptive value is an enigma. On the contrary, it is an essential aspect of humanity and how our species came to evolve. It is a case of the ”taste for the beautiful,” only highly developed and complexified. To once more turn to Darwin: ”In man […] the sense of beauty is manifestly a far more complex feeling, and is associated with various intellectual ideas.”
Kris Lemsalu, Moo-Ma, 2021
Philosophical Aesthetics Revisited
In the arguably most famous book in the history of aesthetics, Immanuel Kant’s so called third Critique, the Critique of Judgement of 1790, natural beauty takes precedence. In the way nature is pleasing to our senses, Kant finds support for a purposiveness of nature beyond the mere deterministic nature depicted in the first Critique, thus connecting nature with the higher sphere of God, human freedom and morality, the topics of the second Critique. Natural beauty thus provides the sought-after connection that Kant describes in the introduction to the third Critique, bridging the gulf between the worlds of the previous Critiques, the phenomenal world of law-bound nature and the noumenal world of human freedom and morality. As Kant writes in §58:
”The beautiful forms displayed in the organic world all plead eloquently on the side of the realism of the aesthetic purposiveness of nature in support of the plausible assumption that beneath the production of the beautiful there must lie an antecedent idea in the producing cause—that is to say an end acting in the interest of our imagination. Flowers, blossoms, even the shapes of plants as a whole, the elegance of animal formations of all kinds, unnecessary for the discharge of any function on their part, but chosen as it were with an eye to our taste[.]”
With Darwin’s theory of sexual selection, however, the reason for the production of the beautiful in nature lies in the taste of animals themselves. Natural beauty is not ”chosen as it were with an eye to our taste,” thus pointing at a supersensible ground connecting the moral destiny [Bestimmung] of humanity with nature, it is grounded in the aesthetic judgment of animals. What Darwin does is to endow animals with subjectivity and mental capacities and, in Kant’s terminology, a ”judgement of taste.” But it is not only, to stick with Kant, an aesthetic reflective judgement, it is an aesthetic constitutive judgement as it, through mate choice – and in the timespan of evolution – forms and creates the natural world. Mate choice is of course not disinterested, a key aspect of aesthetic judgment in Kant, its end being reproduction and desire being essential. But contrary to Kant’s idea of nature’s purposiveness, the beauty of nature treated in this article does not have any higher purpose.
A key aspect of the beauty of nature covered in this article is furthermore its intimate link to desire, once more at odds with Kant’s conception of disinterestedness. But a desire for what precisely? As quoted earlier, Darwin proposes that ”the most refined beauty may serve as a sexual charm, and for no other purpose.” Compared to the reductionist viewpoint where beauty signals fitness or good genes, beauty according to Darwin serves no other purpose than to charm. In this way it is autonomous as it does not follow any other logics, rather it does not have any logic, apart from the standards of taste that has developed. And what more is, these standards are not fixed. In what I would like to see as an essential aspect of the theory of sexual selection, Darwin writes: ”There is also reason to suspect that they [animals] love novelty, for its own sake.” This love for the beautiful, pleasurable and the new – as with feathers, as with the creative brain – can later turn out to be useful for other purposes, but first and foremost they are relished on their own terms.
As we have seen, natural beauty can not only be explained from the theory of sexual selection and the desire for the new, but it is also key in understanding the abundance of nature at large. This is also how the Swedish Idealist and post-Kantian Benjamin Höijer understands it, as seen in one of his major works in philosophy of art of around 1807:
”This sexual [attraction and] sympathy, universal in nature and visible in an infinity of guises, could, in fact, when developed further to its species and possible random causes, shed much unexpected light – and explains individuality – just as this infinity and diversity of nature in its species, and universality in its individuals, is a consequence of individuality[.]”
Sexual desire – könsattraktion and könssympati in Höijer’s vocabulary– would thus be what explains nature in its abundance. It is universal as it can be seen in all nature’s ”subdivisions and subspecies in infinitum” and in an infinity of variation, including, as Höijer adds, between individuals of the same sex. Sexual desire is therefore the creative principle of organic life and of difference, variation, and multiplicity in nature, right down to the individual level; sexual desire being in itself individual and a result of ”possible random causes.” The capricious ”taste for the beautiful” being one of them. Beauty is therefore not only something we find when aesthetically contemplating nature, but a way in which nature works. Therein lies the universality of beauty.
Mats Dahllöv has a PhD in Aesthetics from Södertörn University, Stockholm, and is currently a postdoctoral researcher.
Notes
[i] See Julia D. Monk et al., ”An alternative hypothesis for the evolution of same-sex sexual behaviour in animals”, Nature Ecology & Evolution, vol. 3, December 2019, 1622–1631, https://doi.org/10.1038/s41559-019-1019-7. Superfluous to the discussion here, but I want to add that I have never understood the logics of claiming something to be ”natural” and other things ”unnatural”: if something exists in nature, including in our own species, it is by definition natural.
[ii] Elisabeth Grosz, Becoming Undone. Darwinian Reflections on Life, Politics, and Art (Durham: Duke University Press, 2011), 85.
[iii] Furthermore, to examine and to criticise how societal norms are reproduces in the natural sciences – something not least feminist and queer scholars have been good at – is one thing, to examine alternative theories of understanding nature in these sciences is something different.
[iv] As the title suggests, the book has two themes with the largest part devoted to sexual selection in animals. The first edition of The Descent of Man, and Selection in Relation to Sex was also published in two separate volumes.
[v] Darwin to Asa Gray, April 3, 1860, quoted in Richard O. Prum, The Evolution of Beauty. How Darwin’s Forgotten Theory of Mate Choice Shapes the Animal World – and Us (New York: Doubleday, 2017), 18.
[vi] The expression “survival of the fittest” was coined by Herbert Spencer in 1864 after reading On the Origin of Species, Darwin incorporated it in the fifth edition of the book.
[vii] Charles Darwin, The Descent of Man, and Selection in Relation to Sex, 2nd edition (New York: D. Appelton and Company, 1889), 434.
[viii] Darwin, The Descent of Man, 400.
[ix] Darwin, The Descent of Man, 400.
[x] Richard Dawkins, The Blind Watchmaker. Why the Evidence of Evolution Reveals a Universe without Design (Harlow: Longman, 1986).
[xi] Prum is even talking about a certain ”decadence,” by which he refers to the process where a species by way of sexual selection develops to a point where they could actually go extinct.
[xii] For a hilarious clip that crosscuts different bird chants, Messiaen’s vocal imitation of them, and the performance of his piano composition, see https://www.youtube.com/watch?v=9QdgUJss9BU
[xiii] Claude Samuel, Entretiens avec Olivier Messiaen (Paris: Belfond, 1967), 96: ”si le prédateur veut occuper indûment un endroit qui ne lui appartient pas, le véritable propriétaire chante, chante si bien que l’autre s’en va […] [mais] si le voleur chante mieux, le propriétaire lui cède la place.”
[xiv] Gilles Deleuze and Félix Guattari, Qu’est-ce que la philosophie? (Paris: Minuit, 1991), 174f. The Tooth-billed bowerbird, or Scenopoetes dentirostris, also figures in Mille plateaux.
[xv] In his major opus Différence et répétition, Deleuze hails Darwin’s theory of natural selection as inaugurating the thought of individual difference: ”The leitmotiv of The Origin of Species is: we do not know what individual difference is capable of!” Deleuze thereafter, falsely, credits the German evolutionary biologist August Weisman for his essential contribution in showing how individual differences find a natural cause in sexual reproduction. Deleuze quotation from an 1892 French translation of Weisman on the principle of sexual reproduction is nevertheless on point: ”incessant production of varied individual differences.” See Gilles Deleuze, Difference and Repetition, transl. Paul Patton (New York: Columbia University Press, 1994 [1968]), 248f. View to the fact that Darwin’s theory of sexual selection was more or less forgotten at the time, one has to once more credit Deleuze for his use of the most diverse and obscure sources.
[xvi] Prum, The Evolution of Beauty, 147.
[xvii] Prum, The Evolution of Beauty, 148.
[xviii] See Prum, The Evolution of Beauty, 35–40. Fischer’s idea was the most important contribution to the theory of sexual selection from the first half of the 20th century – it had to wait half a century to make an impact.
[xix] This and the following quote in Prum, The Evolution of Beauty, 33f.
[xx] Geoffrey Miller, The Mating Mind. How Sexual Choice Shaped the Evolution of Human Nature (London: Vintage, 2001 [2000]), 46.
[xxi] Amotz Zahavi, ”Mate selection – A selection for a handicap”, Journal of Theoretical Biology, vol. 53, no. 1, September 1975, 205-214.
[xxii] Quoted in Prum, The Evolution of Beauty, 38f.
[xxiii] Richard Dawkins, The Selfish Gene, 30th anniversary edition (Oxford: Oxford University Press, 2006), 307.
[xxiv] Prum, The Evolution of Beauty, 250.
[xxv] Prum, The Evolution of Beauty, 11.
[xxvi] Grosz, Becoming Undone, 132.
[xxvii] For this aspect in manakins, see Prum, The Evolution of Beauty, chap. 6, ”Bromance Before Romance”.
[xxviii] Darwin, The Descent of Man, 362.
[xxix] Darwin, The Descent of Man, 225.
[xxx] Prum, The Evolution of Beauty, 168.
[xxxi] Prum, The Evolution of Beauty, 173f.
[xxxii] Gerald Borgia, ”Why Do Bowerbirds Build Bowers?”, American Scientist, vol. 83, no. 6, November-December 1995, 542-547.
[xxxiii] Bonobos (also known as pygmy chimpanzees) and chimpanzees, the two species of the genus Pan, diverged around 2 million years ago, and their common ancestor and the human ancestor around 8 million years ago.
[xxxiv] I here use ”hominid” to refer to our extinct ancestors in the Homo genera.
[xxxv] See Lee T. Gettler, ”Direct Male Care and Hominin Evolution: Why Male–Child Interaction Is More Than a Nice Social Idea”, American Anthropologist, vol. 112, no. 1, Mars 2010, 7–21.
[xxxvi] Jeanne Bovet, ”Evolutionary Theories and Men’s Preferences for Women’s Waist-to-Hip Ratio: Which Hypotheses Remain? A Systematic Review”, Frontiers in Psychology, vol. 10, June 2019, https://doi.org/10.3389/fpsyg.2019.01221
[xxxvii] Alex L. Jones and Bastian Jaeger, ”Biological Bases of Beauty Revisited: The Effect of Symmetry, Averageness, and Sexual Dimorphism on Female Facial Attractiveness”, Symmetry, vol. 11, no. 2, February 2019, https://doi.org/10.3390/sym11020279.
[xxxviii] Prum only takes one example at odds with the predominant female beauty norm in our culture to underline this, namely the Mauritanian culture where female obesity is regarded as so attractive that girls of normal body weight are sent to ”fat camps” to gain weight. See Prum, The Evolution of Beauty, 257.
[xxxix] In their introduction, James Moore and Adrian Desmond argues, with support in Darwin’s notebooks, that Darwin’s revulsion against slavery was a main reason behind the writing of The Descent of Man. See ”Introduction”, in Charles Darwin, The Descent of Man, and Selection in Relation to Sex (London: Penguin, 2004), xi–liix.
[xl] Brighter skin colour, however, is likely the result of natural selection, as it facilitates vitamin D synthesis in regions where sunlight is weaker. Regarding the smaller variations in skin colour between populations at the same latitude, sexual selection has likely played a role.
[xli] See chap. 9 in Prum, ”Pleasure Happens”, and chap. 7 in Miller, ”Bodies of Evidence”.
[xlii] Prum, The Evolution of Beauty, 239.
[xliii] It could be added that this has nothing to do with male domination, quite the opposite, as seen in the dominant male gorilla with its one-inch penis. See Miller, The Mating Mind, 233.
[xliv] Dawkins, The Selfish Gene, 308.
[xlv] Prum, The Evolution of Beauty, 250.
[xlvi] Prum, The Evolution of Beauty, 254.
[xlvii] The Latin word sapiens having the meanings wise, clever, and the like.
[xlviii] And with this: patriarchy. Prum suggests that the advent of civilisation and patriarchy can be seen as a new cultural way of waging the conflict between female autonomous mate choice and male control. See Prum, The Evolution of Beauty, 331–333.
[xlix] Darwin, The Descent of Man, 359
[l] Immanuel Kant, Critique of Judgement, transl. James Creed Meredith, revised by Nicholas Walker (Oxford: Oxford University Press, 2007), 175.
[li] Darwin, The Descent of Man, 93.
[lii] Benjamin Höijer, Ideer til den Sköna Konstens Historia, in Samlade skrifter. Tredje delen, ed. Jospeh Otto Höijer (Stockholm, 1826), 442: ”Denna öfver naturen universella och i en oändlighet af skepnader synbara könssympathi kunde tiläfventyrs, närmare utvecklad til sina arter och möjliga tilfälliga anledningar, breda öfver mycket et oförväntadt ljus – och förklarar individualiteten – äfvensom denna naturens oändlighet och mångfaldighet i sina arter, och universalitet i sina individuer, är en följd af individualiteten.” The awkward language is due to the fact that Höijer never got the time to properly edit and finish the texts. Like his other two extensive works on philosophy of art, it only exists in an unfinished manuscript state.
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