“There Remains the Female”: How the Theory of Avian Territory Eclipsed Sexual Selection after World War One

by David Hoyt 

Do birds have a sense of beauty? Do they, or does any animal, have an aesthetic sense? Do they respond to beauty in ways we might find familiar – with a feeling of awe, suffused with attraction, mixed with joy? Do they seek it out, and perhaps even work to fashion it from their surroundings? Darwin thought so, and made the idea the subject of his second major work, The Descent of Man (1871). In it, he outlined a mechanism by which the sense of beauty might, by shaping mating preferences, work to shape the form of insects, fish, and birds in a manner parallel to the better known process of natural selection. The resulting beauty of form, sound, or movement, Darwin argued, is neither the result of intelligent design, nor a necessary indication of superior fitness. Beauty, as Richard Prum has put it, simply happens – but in an organized, understandable way that leads to the dazzling primary feathers of a Peacock, or the bewitching song of a Wood Thrush.

The idea was, as ornithologist Matt Ridley calls it in the subtitle of his recent book, Darwin’s “strangest idea.” It is probably safe to say that the idea, known as sexual selection, was also his most original, and perhaps even most controversial, even more so than the idea of a blind transformism capable of achieving astounding organic adaptations over time. Though it made steady headway after it was introduced, natural selection was challenging enough for many to accept in the second half of the nineteenth century. Sexual selection, on the other hand, premised as it was on idiosyncratic female preference driving the evolution of ever-more varied and harmonious displays among male suitors, was, in the late Victorian period, simply a step too far. As a number of recent authors on the subject have noted, including Ridley and Prum, by the early twentieth century the idea was ushered out the door like an embarrassing dinner guest.

How this happened is a story that has not been fully told outside the confines of ornithology or the history of biology. For those who may recall Michel Foucault’s proposition that the Victorian period, rather than being characterized by the repression of all things sexual, in fact made sex the subject of boundless discourse, the story of Darwin’s theory of sexual selection rings true, but with a twist: Yes, nineteenth-century Nonconformist churchmen could fulminate against the ascription of sexual meaning to the Peacock’s  spotted or the ritual display of the Malay Bird of Paradise; Yes, gentlemen naturalists could lavish attention on the polyamorous mating displays of the Black Grouse on the leks of Britain, Germany, or Scandinavia; and Yes, they could detail the timing mechanism, and ritual of mate pairing as if matters of civilizational import depended on it. But, No, this did not continue beyond the Edwardian period, and in fact was replaced – or displaced – with something like the very repression that Foucault dismissed.

Or, perhaps a better way of putting it is to say that the problem of sexual selection was left unresolved, as often happens, and that attention was, not without relief, turned elsewhere. That this occurred between roughly 1900 and the end of the First World War is in keeping with the transformative effect of that conflict, but also with certain large-scale developments that led up to it. These include the changing status and mobilization of women, the growing scale and concentration of British industry, and the solidification of the modern nation state as the institutional grounds of sovereignty.

What arose in place of sexual selection was, as amateur ornithologist and assiduous birdwatcher Henry Eliot Howard titled his paradigmatic work of 1920, a theory of “territory in bird life.” If you pay any attention to birds, you are probably aware that they are all ‘territorial’ to some degree. Even in Howard’s day, this was common knowledge. As numerous commentators pointed out in response to Howard’s work, including mid-century fans such as Julian Huxley and Ernst Mayr, the idea itself was nothing new. Aristotle had remarked upon it. What was new about Howard’s formulation was that it used the concept of territory to gather together all the different aspects of a bird’s life history and organize them around one central biological imperative: the necessity, for the sake of reproduction, of securing a piece of ground upon which to “discharge of the sexual function.” With Howard, sex became secondary to geography, and Darwin was led to the penalty box.

In this sense, the theory of territory was a functionalist model of the sort that swept across the human sciences during the immediate post-war period (Bronislaw Malinowski’s early functionalist ethnography, for example, first appeared in the early 1920’s), including the emerging field of ethology, or the study of animal behavior. Within the territory model, the male did almost all of the work. He got there first during migration. He located and defended a breeding territory. He then advertised its existence to nearby females. All the while, he battled off contending males, intimidating or signaling dominance to them by means of his colors and plumage. The female showed up later, was appropriately stimulated by everything the male had done, and mated. Elaborate song, visually arresting plumage, and ostentatious behavior on the part of the breeding male were disengaged from the effort of seduction and any evolutionary implication it may have had, and attached instead to the establishment and defense of territory. “There remains the female,” Howard wrote, with a touch of dissatisfaction, in a chapter devoted to birdsong. “I place her last in order of importance.” Not without qualification, of course, but still. Or, further on, “the appearance of the female on the scene marks the opening of a new stage in the life-history of the male.”

This is not to deny that territory is a salient factor in the behavior of birds, as had long been recognized. Nor should it be taken to imply that Howard was insufficiently observant or rigorous. The opposite was acknowledged by his peers both then and now. In fact, it is arguable that Howard represented the last in a great tradition of British field naturalists, reaching back to before and including Darwin himself, characterized by somewhat retiring individuals who were more comfortable in bog boots than drawing rooms, and who were not afraid to put in chilly pre-dawn hours in the “hide”, or duck blind (something the unhappily married Julian Huxley famously did on his honeymoon). Among Howard’s papers at the Bodleian Library are 29 field notebooks, not counting diaries and miscellaneous writings, themselves also containing field notes. In its way, the theory of territory is an impressive case of ecological thinking avant la lettre, framed as it was in the context of the hemispheric, annual migration of warblers to and from the British Isles, and how all their behavior must be considered with functional reference to the execution of reproduction within this overarching geographic feat.

That said, it is striking how the idea of territory served as a felicitous off-ramp from the Victorian impasse of sexual selection. It emerges from the nexus of Howard’s family history and career experience. Tellingly, the British ornithologist most responsible for shutting down interest in sexual selection in the early 20th century, Julian Huxley, co-authored a brief paper with an elderly Howard, and wrote the introduction to a reprint of Territory in Bird Life in the 1960s. Howard’s particular solution in 1920 derived from the intellectual heritage of an accomplished grandfather who was a voluble creationist, and Howard’s own stewardship of one of England’s largest industrial firms at a time of cutthroat international competition – first in the global marketplace, then on the battlefields of Europe. Howard’s own inherited diffidence regarding Darwin, and his direct experience of an increasingly managed and partitioned world, helped him to synthesize aspects of bird behavior in a way that resonated widely.

Henry Eliot Howard (1873-1940) was a director of the largest steel tube manufacturer in the British Isles, Stewarts & Lloyds, and came from an accomplished family with deep Quaker roots. His great-grandfather Luke and grandfather John Eliot had both left the Society of Friends in favor of the less inner-directed, more strident Evangelical Christianity then gaining popularity in the first half of the 19th century. They were successively drawn to an informal group known as the Plymouth Brethren, who stressed a greater attention to the text of Scripture. Both his grandfather John Eliot (the creationist) and his great grandfather Luke were Fellows of the Royal Society – John Eliot for work on the acclimatization of quinine-providing cinchona trees to plantations in the British Raj, and Luke for having come up with a classification system for clouds that is used to this day. Luke corresponded with Goethe; John Eliot maintained a global network of contacts and colleagues in the emerging field of quinology, and acted as an ‘outside expert’ on cinchona bark in relation to Kew Gardens.

Luke Howard had been, like many Quakers in the early 1800s, active in the cause of anti-slavery. John Eliot, by the third quarter of the nineteenth-century, was an Evangelical imperialist. Both men expressed a commitment to scientific truth and inquiry, for which they were each decorated. They also shared a conviction in the ultimate harmony of scientific knowledge with religious truth. It was the mid-Victorian John Eliot, however, who felt the full force of Darwinism’s impact on this placid equilibrium. Not long before his death in 1883, he penned a thundering denunciation of Darwinism in response to a speech given at the Royal Society. Titled “Creation and Providence, with especial reference to The Evolutionist Theory” (1878), it singled out, among the various objectionable elements of Darwinism, the theory of sexual selection:

“The perception of beauty in Creation is a reflection of an attribute of the Infinite Mind, and, like the perception of harmony, is intuitive, belonging to man in his original perfection…But if this last statement be admitted, much less ought we to extend to the lower animals these aesthetic tastes. Can we suppose any sense of abstract beauty to influence the mental emotions of the swine? Or have we any reason to think more highly of the taste of the peahen? … The thrill of pleasure accompanying the expansion of his [the peacock’s] tail is in no way dependent on her stolid regard; nor do I believe that the range of her visual organs is sufficient to take in at once, as we do, the superb spectacle. Certainly the propagation of the race would have gone on just as well if the male had been as plain in his plumage as the female…What then becomes of the theory of “sexual selection” in reference to beauty? It presupposes aesthetic tastes which we have no right to presuppose exist….”

Henry Eliot would have been ten years old at his grandfather’s passing, and so not immediately aware of this creationist polemic. There is reason to believe, however, that Howard’s later theoretical formulation arose in partial response to this emotional controversy, and represented his attempt to bridge the widening gap between science and an ambiguous religiosity.

His own father, Henry Howard, was not among the London-based line of commercial chemists (Luke and John Eliot), but was rather in the “iron trade” centered in the English Midlands around Birmingham. Like medicinal chemistry, the iron trade was a sector in which Quakers had long been active and benefited from extensive networks. Henry Eliot’s uncle, Joseph, was also in the iron trade, and entered politics as a Tory representative for a London suburb from 1885 to 1906. Conservative politics, together with a passion for observing the natural world, and a blend of older Quaker and newer Evangelical religiosity, were thus significant legacies in the heritage of the Howard family.

Henry Eliot’s eventual engagement with Darwinism would not be a head-on battle, as it had been for his grandfather. By the time he was at university, the ground had been largely ceded to evolutionary theory at such nodes of industrial, middle class affluence as Birmingham’s new Mason College. Henry Eliot’s alma mater opened its doors in 1880 with a speech given by Darwin’s great advocate, Thomas Huxley. A private school financed by a local manufacturer, Mason College was intended to offer scientific instruction with a view to its practical applications among the “thousand trades” of Birmingham, second only to Manchester in the measure of its diverse and innovative industrial economy. There was, as reported in the journal Nature in 1880, to be “no restriction of the advantages of the college as to sex, creed, or birthplace” at Mason.

An example from the institution’s first decade conveys the ambience of the school. Historian Anne Rodrick describes how a group of students, about one third of which were female, launched a journal, the Mason College Magazine, the first editor of which was an ambitious and talented young woman, Constance Naden. She was tragically short-lived, stricken with complications from ovarian cysts at age 31, yet she read widely and was a prolific writer. She has been rediscovered of late as a poet. Among her philosophical and poetic writings are a collection of verses given the title “Evolutional Erotics. A stanza from one of these, “Natural Selection,” reads:

And we know the more dandified males
By dance and by song win their wives –
It’s a law that with Aves prevails,
And even in Homo survives.

In 1885, a work of more substantial tenor appeared in the journal of popularization, Knowledge, titled “The Evolution of Beauty.” Here, Naden offered her own understanding of the problem that had vexed Darwin and infuriated, among others, John Eliot Howard:

“We now begin to see why the hen is pleased where one of her lovers develops a new streak or spot or eye of colour, or a tuft of feathers arranged in an unhackneyed and conspicuous fashion. She likes vivid sensations and vivid ideas. It is a joy to be alive, and to be as much alive as possible she must have her faculties brought into vigorous play. So she chooses the mate who will best satisfy this craving for fullness of life…She is a healthy living being, who instinctively takes measures to be as healthy and living as she can. In fact, she is a flying refutation of pessimism.”

Such enthusiasm was not to last, as Naden’s biographer Claire Stainthorp illustrates. In a letter of condolence written on the occasion of Naden’s passing, the philosopher Herbert Spencer, after acknowledging Naden’s gifts, concluded “that the mental powers so highly developed in a woman are in some measure abnormal, and involve a physiological cost which the feminine organization will not bear without injury more or less profound.” The flying refutation of pessimism that had taken wing at Mason College, the healthy living being seeking to bring her faculties into play, was posthumously grounded.

By 1903, it was Henry Eliot Howard’s turn. In an article published in The Zoologist, entitled “On Sexual Selection and the Aesthetic Sense in Birds,” he wrote what could be read as a deferred reply to his grandfather, though in a more quietist key than his elder’s aggressive evangelism, and based on far more observational experience with birds. In the text, the truth of evolution is presumed. All that is disputed is the mechanism of one process, that of mate selection. It is a confusing essay. Howard critiques both Darwin and his antagonist on this subject, Alfred Russel Wallace. Darwin is faulted for studying mostly foreign birds; Howard will study British ones. The northern ruff and grouse that provide so much evidence of male display and female mate choice, Howard claims, simply go at it for too long to provide evidence of any real female choice occurring. As for Wallace, “more Darwinian than Darwin”, he can’t explain beauty through the operation of natural selection alone.

What, then, is to be done? Howard concludes, “[f]rom the beauty of the colours…and from the harmony with which variations tend to develop, it is evident that there is in nature some direct power continuously at work fostering and developing all that is beautiful.” There is male beauty, and there are females ready to mate, but what brings them together remains shrouded in modesty and mystery. Perhaps unsatisfied with his own attempt to resolve the conflict that had roused his grandfather and which clearly troubled him, he went on to drop it entirely. As an epitaph to his attempt, he concludes by quoting, without attribution, Tennyson’s famous lament on the subject of knowledge and beauty, “Flower in the Crannied Wall.” In the book he would write 17 years later, Territory in Bird Life, there are only two mentions of beauty.

The theory of territoriality which he subsequently developed was, as indicated earlier, Howard’s way out of this Victorian problem. But how did he get there? For this, we have to look at the other half of his life, much less well-known than the birdwatching half. His adult career spent at what we might reasonably call the commanding heights of the British manufacturing economy provided him with certain intellectual resources with which to think about populations in competition across space, and how they might be brought into stable relations so as to assure their reproduction over time. For these were exactly the concerns he dealt with as a director on the board of the tube manufacturer Stewarts & Lloyds as it navigated a changing world of corporate concentration, cut-throat competition, and state management of the British economy during the First World War.

A growing though minority consensus began to emerge within British industry by the year 1900 that some form of managed capitalism was preferable to increasingly unchecked competition, which was leading to ruinous price wars and declining profits in industries exposed to international trade. The Germans, among the first to head down this road, came up with a word to describe one of their solutions: Rayonierungskartelle, or ‘territorial cartels’. These were simply agreements between manufacturers to restrict their marketing and sales activities to separate geographic areas – or territories – so as to avoid direct competition. The practice soon found imitators.

If the crisis of revealed religion induced by Darwinism and other sciences was among those that defined the Victorian period (a brand-new biography of Tennyson centers on just this), another was the crisis of free trade ideology brought on by the intensified globalization of world capitalism beginning in the 1870s. Howard refers to his theory of territory as a ‘system’, and it is a system designed to increase the efficiency of mating, so that time and energy are not lost in the search for a mate on the breeding grounds. The system of male-dominated territory is a way of efficiently allocating resources so that the population as a whole, in the grand biotic cycles of migration and reproduction within restricted seasonal windows, may optimally reproduce itself. Greg Mitman has demonstrated the surge of research interest involving animal and human ‘populations’ following the enormous territorial dislocations of the First World War. In this context, what Howard called the “discharge of the sexual function” became a somewhat minor detail.

Howard’s language carried somewhat economistic connotations. I argue that it is no coincidence. Well-defined territory on the breeding grounds optimized reproductive efficiency, as well defined political borders were expected to guarantee international peace. As such, territorial allotment is the opposite of unrestricted competition. The effect of the territorial system within a bird population is comparable to the effect of a cartelized or planned economy on production. Allocation of resources is rationalized. As Howard writes,

“The advantage of this territorial system is therefore apparent. Instead of this district being overcrowded and that one deserted; instead of there being too many of one sex here and too few of the other sex there; instead of a high percentage of individuals failing to procreate their kind, just because circumstances over which they have no control prevent their discovering one another at the appropriate time – each sex has its allotted part to play, each district has its allotted number of inhabitants, and the waste of energy and the loss of time incurred in the process of mating is reduced to a minimum.”

The notion of “circumstances over which they have no control prevent their discovering one another at the appropriate time” is one that had informed the critique of chronic British unemployment since the 1880’s, and the challenge of matching the supply and demand of labor within a cyclical national economy. Howard repeatedly suggests that, in the hypothetical absence of the territorial system, “needlessly severe competition” would ensue, making it unnecessarily difficult for males and females to find each other. “[T]he less severe…the competition, the more uniformly successful would the mating of all the individuals in a given district tend to become.”

This is hardly the language of hackneyed Victorian evolutionism, red in tooth and claw. Coming from the pen of a director of one of Britain’s leading manufacturing concerns, who had helped navigate Stewarts & Lloyds through the firm’s effective appropriation by the state for wartime production, it differs fundamentally from classical statements of Manchester School doctrine. But the regulatory overtones of Howard’s system of territory did not await the rationalization of industrial production that accompanied the First World War. Analysts of Britain’s economic position around the turn of the 20th century were keenly aware of the problems posed by competition now that Great Britain was no longer the sole industrial power. One well-known analyst used Howard’s own industry – tube manufacturing – to highlight the problem of excessive capacity leading to overproduction. In a treatise on the widespread emergence of business trusts, Fabian activist Henry Macrosty wrote in 1907:

“The tube trade has been for years ravaged by a fierce competition, both in the home and foreign trade. Many attempts at regulation by means of price associations have met with no success, but out of the turmoil has arisen one large combination which, though it does not completely dominate the trade, is certainly the most conspicuous within it. Three Coatbridge firms combined in 1890 as A.J. Stewart & Clydesdale, becoming A.J. Stewart and Menzies and the largest tube company in Scotland on the inclusion of a fourth in 1894… Late in 1902 an amalgamation was achieved with Lloyd and Lloyd, of Birmingham, the largest tube makers in England. The express reason was the extinction of competition.”

The source for this stated rationale was none other than Henry Eliot Howard’s father, Henry Howard, as of 1903 head of the newly amalgamated Stewarts & Lloyds (the same year as his thirty-year old son Eliot Howard’s publication in The Zoologist). As Eliot Howard’s father put it, the two largest English and Scottish tube firms “had continually crossed each other’s paths in most corners of the globe. The fight had been severe … This competition would [after amalgamation] for the future cease to exist.”

What amalgamation and eventual state direction did to reduce or eliminate competition between industrial firms, wartime legislation did in the field of labor. The Munitions War Act of 1915 tightly controlled the mobility of labor while also fixing wages, thereby essentially locking workers in place and eliminating the free market. Given the size of Stewarts and Lloyds, it is not surprising that a number of cases involving their employees came before the special wartime Munitions Tribunals, an institution established for the purpose of arbitrating conflicts over of wages and workers attempting to change jobs. Workers, like their concentrated employers, were rooted to fixed locales.

Episodes of labor conflict erupted during and immediately after the war, especially in the heavily industrialized area around Glasgow known as Clydeside, where Stewarts and Lloyds ran the largest tube factory in Scotland. Strikes in 1917 involved some 200,000 Clydeside workers. Further research would be required to determine the precise impact on Stuarts & Lloyds, and their response. We do know that, as with virtually all wartime production, the government-sanctioned practice of hiring and training women in the Glasgow factories to substitute for men sent to the front led to high expectations of post-war reform in areas such as housing, childcare, and healthcare, or what Clara Mattei calls “the material conditions for women’s emancipation.” These expectations were consolidated by a report of Parliament in 1919, written with direct input from female factory workers. Mattei relates how most of the suggested reforms arising from the period of worker mobilization were stymied. In the period of conservative retrenchment following the militancy of 1919, at least at the level of policy, it was made clear that the women who had experienced wartime factory employment were expected to return to lower-paid, lower status, and less visible domestic jobs. As with territory, so on the shop floor: “There remains the female.”

Henry Eliot Howard thus carried out the fieldwork for and composition of his influential book on territory during a prolonged period of upheaval with which he was directly involved. His image among ornithologists remains one of a meticulous, dedicated, and isolated fieldworker, alone with his notebooks on the northwest coast of Ireland or along the marshes of the River Severn. That his personal experience with the spiritual dilemmas of the late Victorian era, or his professional life at the commanding heights of British capitalism, might have influenced his ideas about birds has not, to my knowledge, been previously considered.

This is not to say that the theory of territory is merely ideological, nor incorrect. Howard formulated a problem in a novel and comprehensive way, and at a level (of populations rather than individuals) that gave people a fresh way of thinking about bird behavior. That an idea with such an ancient pedigree as that of territory became so suddenly popular, however, suggests the influence of factors other than those purely empirical. Howard also made mistakes, and missed things. Female birds do sing, even in the Northern Hemisphere. Female choice in mate selection might have been more visible if he had studied a smaller population over several generations, and identified individuals and their offspring through banding, as was just then being done by amateurs such as Margaret Morse Nice in the United States. A close, first-hand study of the Black Grouse of the nearby Scottish moorlands, such as that undertaken just over a century ago by Edmund Selous, might have made it more difficult to dismiss the role of male display and female choice. Despite the best efforts of mentors such as Julian Huxley to recruit Howard’s material for deployment against the theory of sexual selection, a century later it is now more widely accepted than ever that female mate choice does happen, to beautiful effect, even among British birds.