Robert Ardrey, The Territorial Imperative: A Personal Inquiry into the Animal Origins of Property and Nations, 1966.
2. Arena Behavior
The Uganda kob is among the supreme beauties of the antelope world, a photographic delicacy for antelope connoisseurs. Less graceful than the impala, less majestic than the kudu with its corkscrew horns, the kob has a sturdy elegance unlike either. His coat is a golden brown, like proper toast. There are black-and-white markings about his face, and they vary considerably, so that two kob, like two people, seldom look quite alike. He stands about three feet high at the shoulder, but his neck is so long, his curved, lyrate horns are so sweeping, his dark eyes regard his fellow kob so imperiously, that he seems much larger. He is a superb beast, and in 1960 I thought I knew all about him.
I spent the month of June of that year in the eastern Congo and western Uganda, home base for the kob. It was the last month of Belgian rule, and while things were still quiet in the Congo, it required the assistance of no witch-doctor's bones to inform me as to what would happen next. Tourists vanished. My wife and I were the last two guests at the Congo's magnificent game reserve,
the Pare Albert. We had it to ourselves. My heroic qualities, however, are less than notable, so when independence day came we too cleared out of the area, managing to reach Uganda's capital before the Congo blew up behind us.
There were many occasions in that depopulated month when we could not put aside the sensation of being either the last two people in the world or the first. We shared the African sky, the yellow, unending savannah, the choked, narrow strips of forest along swirling streams, the hazy, gray-blue central African lakes with our hosts, the elephant, the buffalo, the hippo, the lion and topi and water-buck and kob. We were their only guests. A sense of extraordinary intimacy pervaded our arrangements. The kob must have water every day, and so he favors this area around Lakes Edward and Albert. For hour upon hour we watched herds grazing on some long yellow slope, impala-like family parties with a dozen or fifteen does and a master ram, or perhaps, again like the impala, all-male bachelor parties. And the does watched us, raising their long-necked, delicate, hornless heads out of the deep grass, ears raised like signs of V-for-Victory; as did the males, with their challenging eyes and S-curved, swept-back horns. My sense of intimacy came to include a sense of authority. On the way out of kob country I discussed them with the director of the Uganda National Parks. When many months later I returned to my writing table in Rome, the conclusion was inescapable that, so far as the kob was concerned, I knew everything.
It was three years before I had the opportunity to return to Uganda. Rome, as we left, broiled like a chicken on a mid-summer grill. We de-planed on the high, cool equator with the joy of escaped prisoners who have somehow eluded the hot seat. The air of Kampala, most adorable of African capitals, was that of a shaded garden, newly watered. Dizzily I embraced the Kampala panorama: the temples and churches topping its hills, the blossoms and bank buildings, Indian merchants, African politicians in black jackets and black silk ties, Baganda students, the university's green lawns. On the veranda of the Grand Hotel (it had been the Imperial on my last visit, but that, of course, had been before Uganda's independence) I met an old friend, an anthropologist, who could scarcely wait to get past proper greetings so that he might pull at his
beard and inform me that for over a year there had been an American scientist around who had been looking for me, who claimed he had a bone to pick with me, and who said that everything I had written in African Genesis about the Uganda kob was wrong.
I was outraged. Professionally I was outraged, and I quoted my observations and my authorities while my friend just shook his head and nursed his happy secrets. Personally I was outraged that friends could prove so faithless, and I condemned his soul to dust. But spiritually I was worse than outraged, for I had been back in Africa for only two hours and already I had been ambushed, my euphoria was gone, and there was nothing I could do about it. I inquired gloomily as to what it was that I had got so wrong about the Uganda kob. "Territorial behavior," said Merrick Posnansky joyfully. I demanded the name of the American scientist who knew so much. "Buechner," he said, with regret. "A pity. He had to go back to the United States two weeks ago." He beamed evilly. "But his Swiss assistant's in town."
That afternoon I met Walter Leuthold, the assistant from Zurich. He was pink-faced, young, amiable, apparently harmless. But when tea and his story were finished my vanity was finished too. The family parties that I had watched for so long were the most casual social relationships. Not in a thousand kob years would one of those imperious rams have sexual relations with a member of his seeming harem. Copulation, by general species agreement, is left exclusively to a dozen or fifteen males, out of every population of eight hundred or a thousand, who discharge their massive obligations on something called a stamping ground. It was this stamping ground that Buechner had discovered. That I had been fooled no more thoroughly than several generations of game wardens, hunters, naturalists, and explorers was a poor sort of salve for my injuries. Since I was leaving for western Uganda within a few days in any event, I put the stamping ground on my itinerary. I had to see it before I could believe it.
Helmut K. Buechner is professor of zoology at Washington State University. It had not truly been Buechner but Mrs. Buechner who had for the first time looked at the kob with eyes unglazed by preconception. At about the time when I was coming to my staid conclusions, Buechner was working on a project in the Semliki Flats, a broad remote
area partly in Uganda, partly in the Congo, separating Lakes Albert and Edward in the hot, flat bottom of the Rift Valley. There, one day, his wife came home and announced that something went on with the Uganda kob. Since the Semliki Flats contain some twelve thousand members of the species, it was not too bad a place to look into the matter. In consequence, ethology was presented with a study as elegant as the discovery itself was spectacular; an author was presented with one more hard lesson on taking nothing for granted; and this inquiry is presented with an example of territorial behavior which, while of a most special sort, still makes an excellent jumping-off place.
A stamping ground, the breeding arena of a single population of kob, looks like nothing so much as a series of putting greens conveniently arranged for the benefit of idle guests behind a luxurious resort hotel. With a little help I found one among the most southerly foothills of the Mountains of the Moon. Without help I stumbled on still another several hundred miles to the north, on the bank; of the Albert Nile. As Buechner's long research demonstrated, members of the two populations could have had no possible contact. The complicated sexual game as played at my two widely separated arenas -- and Buechner's arenas in the Semliki Flats as well -- followed precisely the same formal pattern. Yet it could not have been learned one from another.
We deal here with an open instinct in which final behavior is regulated by a genetically determined pattern filled out by social tradition and individual experience. The pattern, governing motivation and what might be called the rules of the game, is common to the species and is instinctual. It never varies. The location of the stampings ground is traditional within each population. Probably accident and environmental assets combined originally to determine the location. Just as champion cricketers in London converge on Lord's, so champion males in a kob population converge on their stamping ground, because generations before them have done it and it is simply the place to go. Finally, the individual competitor must learn his territories and his opponents, those whom he can beat and those he can't, out of his own experience whether glorious or painful. So it is, then, that the open instinct of the kob, with all its authority and startling complexity, absorbs the traditions of a society and the learning of the
individual to perfect its final, inviolable determination.
A stamping ground is not large, a quality of great appeal for animal observers such as myself who look with distaste on all forms of violent exercise. Each little putting green with its close-cropped grass is about fifty feet in diameter and is a territory occupied and defended by a single male. A closely bunched cluster of a dozen or fifteen or eighteen such territories in a main arena may occupy an area no more than two hundred yards across. Here the champion males out of a population of almost a thousand -- a kind of sexual Olympic team -- fight, display, and jockey for position. Here needy females come seeking consolation.
Certain necessities of the gladiators' daily life dictate an aesthetic appeal for the human observer. To begin with, a single kob is beautiful, and fifteen kob commanding their putting greens are fifteen times as beautiful. Also, the site must stand in the midst of wide, rich grasslands with ample forage, water, and preferably a salt lick not too far away. It is best if it stands on a slight rise with an open view for a mile or so around. The lion, a beast with a dislike for wasted effort as pragmatic as my own, is discouraged by fast prey in a position to see him first. That the fabled Ruwenzoris, with their ice and their clouds, loomed up behind my first stamping ground must be regarded as a pleasant accident, not as a sexual necessity in the life of the kob.
There is, however, a vital ecological necessity for such an institution as the kob's. It is an equatorial species found only in areas of Africa where seasonal changes of weather and pasture are slight. The place may be permanent since herds need not migrate with the rains and the grass. And there is a vital physiological factor lending character to the institution. The female adheres to no season of heat. Unlike the females of most mammal species, she will come into her sexual season whenever she weans her latest offspring, in whatever part of the year that may be, and will promptly seek the entertainments of the arena. But from the male's point of view, this female peculiarity commands a towering obligation. Gladiators must be on duty 365 days a year. Buechner studied fifteen stamping grounds in the Semliki Flats and was able to demonstrate that at least seven of them had been in continuous operation at their present locations for at least thirty years. Realizing that a stamping ground was not going out of business just
because he was around, Buechner built a wooden stand beside one for the convenience of himself, his cameras, his notebooks, and his guests.
When from car or grandstand one watches the sexual shenanigans of the Uganda kob, one watches evidence that an open instinct, once perfected, can provide a natural performance as rigorously regulated as any to be found in nature. The female, for one thing, is sexually unresponsive to any male who has not succeeded in gaining one of the putting greens. She may linger for protection near one of those males whom we saw in the deep yellow grasses; she is incapable of copulation with him. But he, likewise, is incapable of copulation with her. The male who has not gained a territory on the stamping ground is sexually unmotivated. Such males gather as a rule in all-male bachelor herds, as careless of the ladies as male patrons in a Portuguese cafe.
The sexual action of a population is therefore concentrated on this little assembly of putting greens before us.. But while it may seem that the action is simple -- that males fight for territories and females flirt with winners -- it is not simple, for there are still more regulations. Within the arena, for example, some properties have greater sexual values than others. In a normal city, real-estate values increase block by block to the city's core; so on the stamping ground sexual values increase from the suburban market of the periphery to the flashing excitements of Times Square. Young ambitious maturing males fight for a foothold on the periphery, to gain a property even in the suburbs; the peripheral males challenge, fight, wait for an opening to gain better locations in the main arena; and on a few central territories -- perhaps only three or four -- stand the champions of the moment, challenged by all, envied by all, desired by every female heart.
The female wants her affection, but she wants it at a good address. Whether or not our human sensibilities are offended or intrigued, it is a harsh truth that the doe is attracted and excited by the qualities of the property, not the qualities of the proprietor. George Bartholomew once puzzled over comparable behavior on the part of Alaska fur seals in rookeries on lonely St. Paul's Island. Cows might congregate in harems passing a hundred on one bull's territory, in twos or threes on those of others. Yet they exhibited the most disdainful indifference to the bull 
himself. What attracted them to this territory, repelled them from that? Bartholomew concluded that they were attracted by the presence of other cows, somewhat in the manner of the nylon rushes of World War II.
We need not puzzle over the selective value in the Uganda kob of the female's addiction to high-value property. Since it coincides with the male's sense of value, it results in a scheme of natural selection of a remarkable order. Only a super-kob lasts long on a central territory. If he leaves his property for water and forage, he will return to find it occupied and must fight to regain it. On his putting green he will be continually challenged by the ambitious. One afternoon I watched a champion resist five such challenges in an hour and a half, one a twenty-minute horn-locked pushing contest that left him scarcely able to stand. Yet the doe, despite her apparent fragility, may in full sweep of estrus demand copulation ten times in a day. In a busy season the proprietor of one of the central territories must somehow sandwich between invasion problems presented by his colleagues perhaps twenty emotional problems presented by his admirers.
The human male, encountering a stamping ground for the first time, cannot fail to identify himself with the contestants before him. And despite his most secret dreams of sex and riot, he will thank a merciful evolutionary destiny that made him a man and not a Uganda kob; it is all just a bit too much trouble. The human female, on the other hand, will have a response quite different. Identifying herself with the doe, she will be embarrassed for all femininity.
One faces a grassy area of African space. Omnipotent males stand their posts, occasionally challenging another with a tussle, more often retreating before an imperious wave of the head. Then business arrives. Four or five slender, long-legged does come gamboling along like high-school girls on their way to an ice-cream soda. They may graze for a bit beyond the arena's perimeter in a preliminary display of how little such nonsense concerns them. Then one will break away, enter, and pass through the peripheral territories without a glance for the suburban opportunists. Arriving at a major territory, she will begin diligently to crop the proprietor's grass. Suddenly one understands that the putting greens are so conspicuous not because the beleaguered proprietors have eaten them down
to their last caloric morsel but because the does, with all of western Uganda for a dining table, have a special appetite for their limited ration.
Our anthropomorphic sympathies for the male, however, are wasted. All that seems to matter to him as his grass is devoured is that somebody cares. He draws himself up, he puffs himself out. He puts his head far back, his nostrils to the sky, and with tiny, strutting steps the noble animal minces back and forth before his intended. One recalls: every posture and movement of his body is that of a drum-majorette leading a school band down the Main Street of a Middle Western town. Dignity vanishes. He holds his head high in this unseemly ritual so that he may exhibit to the grass-cropper his long, strong, sexually irresistible, gloriously buff-colored neck. She continues to crop his grass, but now the action takes a turn. The male, evidently convinced that by now her heart is aflame, approaches her, nose to tail, sniffs her genitals. She gives up the grass and sniffs his. Slowly they circle, sniffing. To the uninitiated observer the game seems won, judgment vindicated. He approaches her rear and with a last ritualistic gesture raises: his forelegs between her hind legs. Having done the proper, he mounts her. And she promptly goes back to eating grass.
The scientific voyeur watches in disbelief. She gives a flip of her slim haunches and he slides off. All around
the stamping ground similar charades by now are in progress where other does have arrived. Males are mincing back and forth, noses to heaven, throats displayed, while voracious little females eat up their grass. On still other putting greens lone males ignore the shows, offer no interference, continue their games of challenge-and-defend, or simply stare into space. On our putting green the male is making another try. He mounts her. She moves. He tries to keep up on his hind legs, a maneuver adding little to his splendor. He falls off. Once again, however, he makes his try. And now comes the startling climax. She crosses the boundary into his neighbor's putting green.
It was as if a wand many millions of years old, borrowed for the occasion from some dusty collection of animal witchcraft, waved across the scene. All changed. Now his neighbor was marching back and forth with mincing step, exhibiting his glorious, buff-colored throat, while she clipped the neighbor's putting green. And our male?
Our male took all in good grace. When you are a member of an animal elite, then you are not only a proprietor but you also observe proprieties. When you are a member of a sexually privileged club with a charter going back into dim reaches of antelope beginnings, then you observe club rules. He forgot her in a moment. Noble, statuesque, he stood alone in the middle of his putting green, looking off at the distant, cloud-cloaked, immortal Mountains of the Moon. When she passed over his territorial boundary, she passed out of his world.
Buechner's observations of the Uganda kob furnish us with one of the most recent and sophisticated studies of territorial behavior. And since I intend to compare them with science's earliest reflections on the territorial principle, it will be useful to note a few of his conclusions for our future recollection.
1. Males compete for real estate, never for females. The kob's territorial and sexual appetites are so profoundly intermeshed that fights generate sexual stimulation. The champion whom we watched in a twenty-minute defense of his property had an erection through most of the combat. Nevertheless, when the female arrives on a territory,
she becomes the sole if momentary property of the male whose grass she crops. No rival will interfere. A flourishing arena is a Breughel-like scene of scattered kob couples in various stages of intimate disposal amid a scattering of solitary males paying no regard whatsoever.
2. Despite the hazards of his profession, the proprietor almost always bests the challenger. Selection throughout the herd has brought only top specimens to the arena, so all are quite equally matched. Possession of a territory offers some mysterious advantage usually sufficient to guarantee victory for the defender. In his first fifteen months of intensive observation, Buechner saw the challenger win on only a dozen or so occasions. Champions fall, of course, but usually from exhaustion. They fail to return from foraging or, returning, fail to regain their posts.
3. So powerful is the proprietor's psychological advantage that dangerous fighting is minimized. Simple ear-lowering, horn-waggling, or other stern display is frequently enough to discourage challenge. Leuthold reported to me a male with a broken leg who in a triumph of psychological warfare held onto his property for eight long days.
4. Off the stamping ground the gladiators display no antagonism. Should a hungry-enough lion appear, the first to spot it gives a stiff-legged hopping signal alerting his fellows. All retire by customary paths to wait amicably until the lion goes away.
5. The inspiration of ownership seems necessary to stimulate sexual desire in both males and females. Away from the stamping ground copulation is only rarely attempted, and apparently never consummated.
6. In the vast population of the Semliki Flats, each group of herds holds allegiance to a definite, traditional ground. Buechner marked many males, and reobserved them on two thousand occasions. Only seven of these reobservations were as far as five kilometers from home grounds.
7. Leuthold has given special attention to an aberrant design for living in kob country. Most males who fail to achieve the main event, or achieving it fail to hold on, join the contented bachelor herds. But there are lonely exceptions. Such a male stakes out a territory of his own somewhere. Near this jutting outcrop or that spreading tree he will always be found. A party of does may join
him -- a dozen, fifteen, twenty -- for the companionship or the security of his august presence. There you will see him in the sleepy afternoon, as I had observed them in 1960, the does lying in the grass, the male standing alert beneath his swept-back horns, an apparently normal antelope family. In a week or so the party of does will drift on to farther pastures. And the male will remain beside this rock, beneath that tree.
"Attachment to a piece of ground," writes Buechner, "is stronger than to the female herd."
Such attachment to a piece of ground has been the subject of organized study in the natural sciences since 1920. In that year Eliot Howard published his memorable volume Territory in Bird Life and established the word and the concept in the language of science. But for many a century before Howard, observers had pondered, briefly or at length, on the notable attachment of a particular animal for a particular piece of earth.
Aristotle had puzzled over birds of prey: "A pair of eagles demands an extensive space for its maintenance, and consequently cannot allow other eagles to quarter themselves in close neighborhood." Pliny, in Rome, bothered too about eagles: "One pair of eagles needs a very considerable space of ground to forage over, in order to find enough food; for which reason they mark out by boundaries their respective allotments, and seek their prey in succession to one another." A thirteenth-century German emperor, Frederick II, like many another royal figure of the day, was a dedicated falconer, and recorded thoughts to be published some centuries later: "After fledgling falcons have learned to fly and hunt bird prey, the parent drives them away not only from the immediate neighborhood of the eyrie but from the entire nesting locality. Were the mother and her offspring to hunt in the same territory, their bird quarry would soon take fright and there soon would be not enough food to supply the needs of the whole family."
Aristotle, Pliny, and Frederick II, we should say today, all subscribed to the food theory of territory. They were not the last to ascribe attachment for a piece of ground to the economic motivation of securing a food supply -- a type of motivation which the Uganda kob has so conspicuously never heard of.
Other early bird-watchers considered the robin. Zenodotus
seems the earliest, in the third century B.C., when he stated flatly and inarguably, "One bush does not shelter two robins." In 1622 G. P. Olina gave his attention to the robin in his Uccelliera, an attention concerned mostly with the bird's bad disposition: "It has a peculiarity that it cannot abide a companion in the place where it lives and will attack with all its strength any who dispute this claim." Not till 1772 did anyone, to my knowledge, bring sex into it. Then Gilbert White wrote: "During the amorous season, such a jealousy prevails among the male birds that they can scarcely bear to be together in the same hedge or field . . . and it is to this spirit of jealousy that I chiefly attribute the equal dispersion of birds in the spring over the face of the country." White's famous contemporary, the Count de Buffon, took exception and opted for economics in the life of the nightingale: "Nightingales select certain tracts and oppose the encroachment of others on their territory. But the conduct is not occasioned by rivalship, as some have supposed; it is suggested by the solicitude for the maintenance of their young, and regulated by the extent of ground necessary to afford sufficient food."
The naturalists of the late eighteenth century were staking out territories of controversy which descendant scientists still quarrel over today. But not until 1868, when the German ornithologist Bernard Altum published Der Vogel und Sein Leben, did anyone take the time to construct a theory out of it. Altum established two main hypotheses that have stood the erosion of time and research: first, that male birds fight for the possession of land, not of females; and, second, that birds sing not for the joy of life but to warn off any intruders who may be contemplating intrusion on their private domains. These were revolutionary thoughts that today have been validated in the notebooks of a thousand researchers. At the time, however, Altum's thoughts vanished. They had been recorded in but a few pages of a study of the life of the bird; and, besides, no one seemed to be listening.
Another pioneer even more emphatically neglected was an Irish ornithologist named C. B. Moffat. He recorded his odd notions in 1903 in a paper called "The Spring Rivalry of Birds," published in the obscure Irish Naturalist. Only now are we beginning to know what he meant. Like Altum, he had observed that property-holding birds sing not so much to impress the female, nor even to express the sheer
joy of being rich, so much as to scare the appropriate daylights out of anybody with designs on their property. In ornithology, this today is scripture. But Moffat went further. With daring opposition to Darwin's best thoughts, he suggested that the male's bright coloration exists for the same reason. "Have we not here some ground afforded us for suspecting that the bright plumage may have been originally evoked as war paint? In other words, as a sort of warning coloration to rival males, rather than attractive coloration to dazzle the females?"
Not even in our time have we caught up with the challenge that Moffat laid down. Whom does the male have on his mind? The male or the female? For many chapters in Darwin's second great work, The Descent of Man, he develops the proposition called sexual selection -- that the focus of masculine life is the female, and that evolutionary dynamics rest on the male's success or failure at enchanting her. Moffat said no. What the male has on his mind is the male. The female will make her choice, well and good. What is eternally bothering the male is not female estimate, but how he is doing in the eyes of his fellows. Many a contemporary school of psychology would regard this as a homosexual tendency. Nature sighs.
Less articulately, Moffat explored in his little paper another grand notion (and a startling one to have originated in Ireland): that territory acts as a natural mechanism of birth control. The interpretation cannot be valid in such arena species as the kob, but it operates rigorously in species basing social arrangements on the breeding pair. Through the defense and antagonism of territorial proprietors a given area is divided between a consequent number of breeding pairs; surplus population is condemned to sexual nonexistence, and the reproductive population is limited to the land that will support it. This seems a rough way to go about birth control, but the Scottish ecologist and ornithologist V. C. Wynne-Edwards, in his monumental Animal Dispersion in Relation to Social Behavior, has demonstrated that it works.
When we have finished our inspection of arena species other than the kob, I shall come back to what I regard as the most revolutionary thought of this forgotten Irish ornithologist: that territory acts not so much in the interest of the individual as in restraint against the individual in the interest of the group, the population,. and the species.
It is enough now to wish that someone, someday, would record for us the life of C^ B. Moffat. He was evidently a member of the Dublin Naturalists Field Club. He inscribed his paper "Ballyhyland, Wexford." Beyond that I know nothing about him other than that, like the stout Cortez upon a peak in Darien, he gazed upon a blue, unexplored philosophical sea that we have yet to traverse.
Altum died, his ideas neglected. Moffat died, his ideas unknown. Others made an observation here, put forth a speculation there. An American ornithologist named Brewster was impressed by the way one bird will honor the next, bird's "rights." It was a profound impression, but it faded with the sunset. Another American named Herrick studied gull communities and speculated on the relation of territory to society. One more grand boulevard of inquiry was opened, then vanished. There is small likelihood that Eliot Howard, watching his warblers and his buntings among Worcestershire's hills, was conscious of Altum or Moffat or Brewster or Herrick. In the ruthless language of natural selection, the difference between Howard and his predecessors is that their ideas died without offspring, while his left progeny all over the map.
Henry Eliot Howard was a businessman who finished his career as director of Stewarts and Lloyds, one of Britain's two largest manufacturers of steel pipe. His place of business was near Birmingham, but his home was in the country, at a house called Clareland. He began all the days of his life by rising long before dawn, assembling his unprepossessing country costume, and hanging about his neck a pair of binoculars. Eliot Howard was a member of that extraordinary British breed, virtually a species in itself, the birdwatcher. At eight in the morning he would return from the fields and have breakfast. At eight thirty he departed for work, in the 1890's via bicycle and train, later on in a motor car. At teatime he would return. He had five children, a son and four daughters. He would play tennis or chess, discuss Plato. After dinner the family would gather in what the children called the smoking room, an interesting name for it since no one smoked except Father. Mother sat on one side of the fireplace, Father on the other, making his notes of the morning or simply inspecting space. Mother occasionally said to the children, "Shh! Father's thinking."
Father, as things turned out, was doing a fair job with his thinking. Through a series of volumes published in this
century's early decades he became the acknowledged authority within a limited field, that of the British warbler. Then in 1920 he produced the slim monument that will recall his name until men cease to ponder, a book called Territory in Bird Life. It was the first, and until this present volume, the only book devoted solely to the innate relationship between property and animate behavior.
He was a silent man, on the whole; handsome in a British way, with narrow angular face and slender mustache. He had several great friends in the sciences, Lloyd Morgan for example, and the young Julian Huxley. One finds among his old bits and pieces letters from another young scientist, Konrad Lorenz in Austria. Howard was an amateur, and yet he practiced a scientific discipline that any professional might envy. Never in his book does he allow his conclusions to stray beyond the species he knew so well, the world of birds. Not once does he permit himself speculation concerning territory in the life of men. Rarely did he share his preoccupations with his family, for it was not his way. And yet I have spoken with a very old woman who today lives in the north of Wales and who was nanny at Clareland when the children were young. And she recalls a startling night when the silent man came into the nursery and sat down and stared at her. To judge by the age of the children then, it must have been in the year 1904 or 1905. And he said to her, out of nothing, "Nanny! It's territory. That's what everything's all about. Territory. Territory." It gave her quite a turn.
Eliot Howard died at Christmastime in 1940. The Battle of Britain had engaged his countrymen in one of history's most memorable demonstrations of the territorial
imperative. Howard was an old man now, but to the end he went out every morning with his boots and his binoculars J and his unbeguiling costume. By this date, of course, many another Englishman was out in the fields and on the hilltops with binoculars, spotting not birds but bombers, and; keeping a wary eye out for Nazi parachutists. One morning I Howard came in to breakfast. He was meditating and seemed to have something most puzzling on his mind. At last he inquired mildly, "Who do you suppose they think I am?" Days later he was dead.
I dedicated African Genesis to the memory of Eugene Marais, South Africa's pioneer naturalist. If it were the task of a dramatist to invent a character the precise antithesis of Marais, then one could only end up with Eliot Howard. The two were contemporaries. When Marais at the turn of the century was watching baboons in the Transvaal's Waterberg, Howard was watching warblers beside the River Severn. Marais was lonely, tortured, a morphine addict. Howard led one of the calmest lives of the twentieth century. Marais was a lawyer, doctor, journalist, poet, teacher. Howard went to the office. Marais was a genius, a Van Gogh of the natural sciences, whose career was written in waste and passion and demons and who died of his own hand. Howard had a clear eye and a clear mind and he died in bed, and there will be no man, someday, who is not his inheritor.
According to James Fisher and Sir Julian Huxley, who have written an introduction to a new edition of Territory in Bird Life now fortunately available in both Britain and America, the book caused small commotion when it was published. (I believe, indeed, that its only printing had to be remaindered.) Ibis, the journal of British ornithology, rated it "an attractive and thoughtful little work." It is entirely normal that when an astonishingly new idea comes off somebody's mental assembly line, it will take a while before other people's assembly lines tool up sufficiently to deal with it. By the close of the 1920's such tooling up had been accomplished, at least in ornithology. It would still be a while, of course, before the new concept would be recognized as applicable to animals other than birds.
In a way the territorial principle was not that revolutionary, since Howard presented few conclusions unanticipated by someone else. Also, one might say that wha came off his intellectual assembly line arrived in unfinished
condition; nowhere did he define territory. His failure to attempt it may have been just as well, however, since no one down to this date has been able to offer a definition unshadowed by doubt and unpunctured by exception. Howard's accomplishment was to prove that an instinct called territory exists, and he did it through example after example drawn from the lives of those birds he knew so well. Reed buntings and guillemots pass through his pages: ravens, moor hens, pied wagtails, cuckoos, wood pigeons, whitethroats, sedge warblers, tree pipits, ruffs and rooks, nightingales and skylarks, all pass in a testament of love, giving each his sworn statement that the author's word is true.
To wander through Eliot Howard's prose is to walk through budding woodlands under April skies. The quiet of the man commands us. Our hearts are stilled. We hear birds sing. And when theory comes to us, we inhale it through our nostrils like a pungent recollection of last autumn's leaves.
In such modest fashion was the word "territory" lastingly introduced to the vocabulary of science. Others might in later years discover its significance in the lives of lizards and lions, crickets and men. Howard made no such claims. The bird's world was the world he knew, and he made no claim beyond. We, however, may today compare a few of Howard's conclusions, drawn from the Worcestershire countryside in 1920, with a few of Buechner's drawn forty years later in central Africa's Semliki Flats. By such a brief exercise in comparative ethology we shall begin to see territory both as a particular reality and as an underlying pattern in animal affairs. And we shall be wise to remember that we are comparing birds with antelopes, creatures as remotely related as are birds and ourselves.
I recorded seven of Buechner's conclusions to hold in our memories:
(1) That male kobs compete for territories, never for females.
It was Eliot Howard's principal conclusion that, contrary to Darwin's "law of battle" and to all of our most romantic tenets, male birds never compete for females.
(2) On a kob stamping ground, the territorial proprietor almost always wins.
Another of Howard's principal conclusions was that the invulnerability of the proprietor (in the quite different
situation of breeding pairs) is a chief guarantee for the security of the nest and of the young.
(3) The psychological advantage of the proprietor reduces the incidence and severity of actual fighting.
Like Altum before him, Howard concluded that bird song is a territorial display and that, while an invitation to hostilities, it is associated with invulnerability and so discourages challenge.
(4) Antagonism between male kobs is confined to the stamping ground. Elsewhere their relations are amiable.
Eliot Howard had a gift for incisive observation. His mind swept away all irrelevant or obscuring detail that might detract from the purity of a conclusion. Perhaps the quality was no different from that of an artillery officer who carefully and intuitively clears away all obstruction from the range of his fire.
Howard watched moor hens. The moor hen is a water bird living in and around marshy reed-fringed pools, feeding frequently in nearby fields. He (a moor hen can be a he) is an amiable enough citizen throughout most of the year, swimming or waddling about his damp little world. But then, about the middle of February, all changes. Pairs establish territories in what has been a peaceful pond, each territory including a bit of rushy shore. Intolerance rages, for the moor hen's new belligerence is not confined to the male. Pairs fight pairs, storming about the pond and its shores like little modern landing craft fighting by sea or land. Remarkable in territorial conflict, the antagonism is not confined to the species. The moor hen's intolerance for any, intruder is such that a pair will attack harmless strangers who have simply dropped by for a sip of water -- lapwings, thrushes, starlings, even a partridge covey if it comes too close to the pool. Yet in the fields where he goes to feed, or in any area beyond the reedy shore, the moor hen becomes again in an instant an amiable creature. Pairs feed beside pairs. Hostility vanishes. The thrush is ignored.
(5) Copulation occurs nowhere but on the stamping ground.
Another of Howard's fundamental conclusions was that in territorial species breeding pairs are limited in number to those who gain an exclusive property. A male with a mate and no territory is a natural impossibility. Even the puzzling, parasitic cuckoo must have her territory, even though she lays her eggs elsewhere in somebody else's nest.
(6) Populations of kob do not really mix, though to the eye they may seem to. Each breeding population retains an identity with its own stamping ground, making interbreeding unlikely. Genetic isolation is thus achieved.
Warblers are mostly migratory species, returning each spring from winter quarters to northern breeding grounds. It was difficult for Howard to prove, but he became convinced that the same birds return, season after season, to the same breeding grounds. If it were true, then the same genetic isolation would be achieved as in the Semliki Flats. (Post-Howard research of a most exact sort would reveal that the Laysan albatross, for example, which breed on Midway Island in the north Pacific, build their nests in the same spot from year to year. Although all nests have been demolished by the winter's storms, and there seem few landmarks, 50 percent will succeed in building within four feet of last year's site.)
(7) "Attachment to a piece of ground is stronger than to the female herd."
Again and again and again Howard stresses the attachment of the male bird to a piece of ground or leafy space, and the attraction for the female of a male so attached. But never to my knowledge did he observe anything to resemble the solitary territory of the bachelor kob. Perhaps for once preconception clouded his eye. Howard regarded territory as invariably a portion of the reproductive process, something that broader investigation would show is usually but not always true. Perhaps there were propertied bachelor birds whom he failed to notice. It seems more likely to me, however, that it is something that just does not happen at the evolutionary level of the bird. In any case, we must record one aspect of territorial behavior in western Uganda which Eliot Howard, forty years earlier, failed to recognize in the fields and the pools, the heaths and water meadows and woodlands of a manicured English county.
There is a moral, somewhere, in the story of the quiet English businessman: perhaps that it is not a downright necessity to die of drink, wear your hair to your shoulders, beat your wife, and starve in a garret in order to enter the magical forests of human adventure and bring back in your pockets much remarkable fruit.
The force called territory as it affects a world of living beings bears at least one resemblance to a force called electricity as it affects a world of apparatus: the substance of each is as elusive as the effects are spectacular. Most of us will go to our graves still unable to describe with any precision what actually happened when we touched the little flipper in the wall and the lights went on. Similarly, though we may speak of open instincts and innate behavior patterns, no biologist alive can today tell you just what are the genetical arrangements that command a territorial animal to behave in the manner of his species. Someday, perhaps, we shall know. In the meantime, just as one way to find out about electricity is to keep turning on lights, one way to find out about territory is to keep turning on species.
It is a far leap from a stamping ground in central Africa to an athletic field in Brooklyn, and it is a farther leap, so far as evolution is concerned, between a species of antelope and a species of predatory wasp. Simpson has calculated that the mammal and the insect could have had a common ancestor not less than 500 million years ago, and probably closer to a billion. Yet any comparison between the two species will reveal how, despite such disparate inheritance and disparate environments, their arena behavior follows closely the same rules and regulations.
There are hundreds of genera of wasps and thousands of species, and Tinbergen's digger wasp who preyed only on bees was quite normal in her closed-mindedness. Only a few species will choose victims from more than a single sort of insect, and many will hunt their special victims only in a special situation. One species preying only on flies became known as "the horseguard" from hovering constantly over the flanks of horses. A Bembix species also preys on flies but hunts only at twilight, when they have first settled for a night's rest. A species of digger wasp, with a disposition somewhere between that of Karl Marx and that of the Marquis de Sade, preys only on queen ants, never on workers, and takes none but winged ones at the moment of nuptial flight. One might judge, therefore, that so far as wasps go there is nothing too extraordinary about one called the cicada-killer. And yet in his addiction to territory -- so far as we know -- he is unique among wasps and remarkable among insects.
In general, territory is a vertebrate expression, and I am aware of no other insect species which has evolved that highly specialized territorial pattern, arena behavior. But apparently anything can happen. Only a few years ago a man named Norman Lin, walking beside a high wire fence enclosing a Brooklyn baseball field, found the equivalent of the kob's stamping grounds being operated by cicada-killer wasps. With fence posts at nine-foot intervals, it was not too difficult for Lin to lay out a precise grid for observation. With a few dyes and a spray gun it was not too difficult to mark his contestants.
Cicada-killers are ground-nesting wasps who live in colonies of several hundred burrows from which the female emerges when she is mature. The males arrange themselves in territories not unlike the kob's putting greens, tightly adjacent and small. The properties on Lin's ground ranged from four feet square to longer areas up to six by sixteen. On each property a male has a perching place, such as a pebble, which he always returns to after a chase. It is generally accepted that a male bird has his favorite twig, his accustomed fence post, to advertise the fact that he is home and is prepared to take on all intruders. The cicada-killer wasp, on a remote evolutionary track, has evolved his trait through similar selective advantage.
The attachment for his territory is as profound as in any species. Having chased off an intruder, he will return in
seconds to his perch. Lin experimented with marked wasps. He took one 1000 feet away, released it out of sight of the home grounds. It was back on its perch in twenty minutes. He took another almost half a mile away. It was back in fifteen. How did they get back? It is as fascinating a problem as any in science.
So far as defense is concerned, it is continual and inviolable. The wasp will repel an intruder by threat, or by a chase if threat is not enough, or by butting him in midair if chasing will not do, or, in the last resort, grappling with him, tumbling to earth and trying if possible to bite out his eyes. It is a rough game. Lin found that in five cases out of six the conflicts took place between adjacent proprietors, just as in colonies of sea gulls the real rows go on with the fellow next door. The loudly buzzing wasp, either because he is angrier by nature or more dim-witted by historical endowment, will defend his preserve against anything, a passing butterfly or bird. He will attack a pebble rolled across his border. Off the territorial ground, however, such conflicts never take place. Like the kob when the lion comes along, the wasp is a practical creature. When the afternoon grows unbearably hot, all retire for a siesta.
The complete exhibition of arena behavior is demonstrated only when the female comes along. Females are larger than males and easily recognized. He may attack a pebble, but he will never attack her. The female mates only once, and so as she comes flying across a territorial ground there is a question to be settled: is she not ready yet, is she ready, or is it too late? She answers it simply. If she is sexually unresponsive, she will always fly a zigzag course, and always slowly. Males may rise from their perches to investigate, and they may pursue her gently, but never across a territorial border. She will be escorted, as it were, by a succession of males, each crossing his territory with her. But if she is responsive, she always flies straight. And the first alert male whose property she crosses will rise and grasp her from behind, and they will fly off together in tandem to alight somewhere and copulate.
Lin has one story garnered from his jungle in Brooklyn which illustrates what instinctual bewilderment can come about when events take an unexpected turn. Normally a fevered couple vanish somewhere to perform their rites. One copulating pair, however, landed on another male's
territory. Since it was his place he promptly landed on her, attempted copulation, and discovered of course that she was occupied. Then a third male intruded on his territory. The proprietor chased him, grappled with him, drove him off, returned to within a few inches of his coupled guests, quite obviously lacking a clear inward directive as to what to do about them. Two more males came over his territory, hovering, and he rose and drove them off, again returned. Still attracted, the two intruders returned and by now the proprietor was committed to defending his guest along with his rights. Until affections had been exhausted and they departed, he continued to protect their privacy.
Perhaps Lin's bewildered wasp suggests why insects so seldom indulge in the delights of the arena: its demands are too trying. For a creature whose instincts tend to be closed with less room for the final programming of tradition and learning, the arena becomes a complicated place. Should this be so, then the wonder remains that cicada-killer wasps have mastered its intricacies as well as they have. Or perhaps, on the other hand, Lin's wasps seem unique only because we do not know enough about insects. Certainly it is the long-studied world of birds that furnishes us with our richest arena examples.
Ruffs and reeves constitute a single species related to the sandpiper, but because male and female look so very unlike, custom has held separate christenings. This quality which zoologists call sexual dimorphism, a marked dissimilarity between sexes, is more common than not in arena species. I mentioned how unlike are the sturdy, splendid male kobs and the slender, delicate does. Although it is not true in the cicada-killer wasp, the chances are that in species where the male has no part in raising babies or protecting a nest, but devotes maximum energy to impressing his fellows in the local arena, natural selection speeds up the acquisition of finery. The process has gone to an extreme in the ruff. For eight months of the year the cock, although a bit larger, is no more conspicuous than the hen, and neither has an appearance more notable than that of any other sandpiper. Then with the first breath of almighty spring something happens to the cock, and he starts assembling his costume for the arena.
The ruff takes his name from the most impressive of his adornments, a circular shield of feathers which he sprouts about his neck and which he can raise or lower as occasion
demands. Patterned in white, black, bay, chestnut, gray, blue, violet, even gold, no two cocks ever look exactly alike. Few living creatures acquire in a time so short an adornment combining such beauty and individuality, such pomposity and practicality. For the ruff's ruff is a shield so strong that it protects his chest from his rivals' bills.
Thus arrayed like a medieval knight in his loved one's colors, the ruff with the breath of awakening spring proceeds to the hilling ground. No parting tears, however.' signal his departure. Aside from a moment's classic attention which he will pay the hen in the arena, the cock live a cock's life in a cock's world, and throughout the yes has no more to do with the reeve than if she indeed wer of another species.
Along the Dutch coast the traditional hilling ground as permanent as Buechner's stamping ground -- lies adjacent to the traditional breeding ground of the reeve. Occupied as she is with nest-building and feminine sociality, the reeve ignores the arena except when the moment of desire arrives. In the meantime, from early April until June, the hilling ground is the ruffs ball park and social center. They are smallish birds, and their territories are smallish, and each has a little hill less than two feet in diameter kicked up by ruff feet over many a decade. On
this hill he displays his glories, defends his real estate, and insults his neighbors.
Back in 1920, the year of Territory in Bird Life, a Miss E. L. Turner visited several hilling grounds in Holland and reported her bewilderment in the journal called British Birds: "The ruff is either as motionless as if he were carved in stone, or else he is vibrating like a toy on wires. . . . They rush around with the regularity of a clockwork mouse. When several are fighting together they are an indistinguishable blur of feathers. . . . They filled me with amazement. Why do they behave in this ridiculous manner?"
The ridiculous activity about which Miss Turner complained cannot compare, I should say, with the ridiculous inactivity when a reeve at last appears. We are beginning to note, I hope, how in these sporting events of the natural arena, whatever the fierceness of the competition, there are always rules and conventional restraints. As part of his display before his competitors, the ruff has developed a dismaying capacity for holding his breath. By such means he distends himself to his anatomical maximum and spreads his ruff to its grandest proportions. Now, as the reeve enters the ground, all on their little hills bow low. Beaks almost to the ground, ruffs perpendicular like hanging shields, they hold their breath while she inspects the art show. None moves. That would be against the rules. They have reminded many an observer of a bed of flowers. She wanders here, she wanders there. At last she chooses, pecking the neck feathers of the ruff of her choice. They mate immediately. There will be no objection from the disappointed for the very good reason that they have all collapsed.
Ernst Mayr has defined arena behavior as a territorial pattern in which males defend mating stations unrelated to feeding or nesting. But V. C. Wynne-Edwards, in a description of the lek, the traditional dancing arena of the blackcock in Scotland, has recently introduced another point which we must not neglect: that these displays, directed by males toward one another, decide and maintain the social status of each. On an early page I regretted the necessity of fixing the attention of this inquiry on the territorial principle. As important as territory to social animals -- and we may find someday that it is more important -- is the compulsion to achieve status within one's society. Territory is essentially defensive, an inward mechanism aiding us to defend what
we have; status is essentially aggressive, an inward pressure to achieve dominance over our social partners. In the arena the two innate forces combine to bring about a single pattern. Through the holding of a territory, we defend what social status we have achieved; by challenging our neighbor, we attempt to better ourselves. In the kob, differential real-estate values prevailing on the stamping ground provide neat territorial ladder: this rung offers security for status so far achieved, while the next beckons us to rise. We shall find something very much like it in Wyoming's sage grouse, an example even better for our purposes than Wynne- Edwards' more famous blackcock.
Until the early 1940's there was so little known about the mating of sage grouse that a rumor thrived on our high western plains that the cock drops sperm and the hen pecks it up. Since the sage cock is another elaborate bird with an appearance very nearly as vain as a turkey's, the rumor was of a most derogatory sort. Then J. W. Scott, of the University of Wyoming, discovered the first known strutting ground, and from then on no further reflections could be cast on the cock's reputation.
A seasonal creature like the ruff, the sage cock has an arena of a seasonal nature. In winter the flocks mingle in peace, both sexes together. Then in March the cocks go their way, descending on the strutting ground, where for the next three months they will display, browbeat, scramble for geographical position, and sort themselves out into a hierarchy. Like the kob, a population has from 800 to 1000 members; unlike the kob, however, all males will participate in the carnival and so the arena must be enormous. The first ground Scott studied was half a mile long and 200 yards wide. Nevertheless, it had the same permanence of site as all natural arenas; one that he later studied had a road built through it, but the birds refused to abandon their hallowed area of sagebrush and space. As in other arena species, too, the contestants had that intent preoccupation with matters at hand which allowed Buechner to build a wooden platform beside his field of glory; Scott put up a comfortable tent, with windows, in the middle of his.
There was plenty to view. Even on the first day of the season, toward the end of March, with ice still a half-inch thick on ponds, 175 cocks reported for assembly. It was not hen weather. One showed up, either by accident or because
she was a mutant creature equipped with central heating. Her presence was unnecessary, since the male motive is to impress the next male. Claims were staked out. On a strutting ground every cock in the population gets a court, but, as Scott was to learn, only central courts have either social status or sex appeal.
The scramble continued virtually henless for some time. Normally the cocks scattered about the countryside, feeding most of the day, to assemble in the afternoon and to continue their exercises into the twilight. If a golden eagle soared into view, all fled, to resume their disputes when the eagle had passed. As a new moon came on, the performance carried on later and later, until it was possible for an hour-long riot to take place at three in the morning. But by this season the hens were arriving in numbers.
Action on a stamping ground in Uganda concerns a mannered elite who, like any elite, may seem preposterous but still retain an elegance. Action on a Wyoming strutting ground is an orgy organized in a rush hour. I seem to recall that in the early days of silent films Hollywood presented a few massive scenes of bacchanalian grandeur which might favorably be compared with the sexual entertainments of the sage grouse in Wyoming's open air. On one single occasion Scott counted 355 cocks and 141 hens. And yet it is an orgy organized to the last feather.
Orily five courts at the most have sexual value. Each is no more than eight feet by twelve, and onto these jam the entire population of ready hens. On each court is a master cock, the survivor of the status struggle. Nature has wisely recognized that the arithmetic of such a system, while flattering to the master cock, may be one which neither the species nor the master cock will survive. And so each court is equipped with his chief rival, a subcock, who will spell him off when the situation becomes trying. Each court is also equipped with two or three guard cocks, whose duties, whether taking tickets or subduing the mob, have never been quite clear to me.
The hen, however, will as a rule have no part of any but Number One, and for an hour or so, if he is the worse for wear, then his admirers jammed closely together will simply wait for his recovery. How physiologically directed is the sage hen to those behavioral laws applicable to her species may be judged by some of Scott's figures: On one strutting ground which provided only four mating courts for a normal
population, three-quarters of all matings were accomplished by the four master cocks. Over half the remainder were provided by the four strutting rivals, the subcocks. The guard cocks in moments when the traffic was heavy managed to get in on a few. And there is a very small indelicate remainder or reminder, the score of sage-grouse immorality chalked up in the anonymous sagebrush; the sage hen on occasion is only human.
For one who lives in Rome within walking distance of the Colosseum, of circuses, of amphitheaters -- the very term "arena" is a Roman word -- there is a temptation to ask, Is man at least in part an arena species? We have our prizes of property and status, our market places of male competition. We all of us shelter memories, comical or terrifying, poignant or absurd, of human behavior not unlike these animal exhibitions we have witnessed. And there are sound evolutionary reasons to inspire a moment's wonder, too.
Thomas Gilliard, of the American Museum of Natural History, has given many a fascinated year to contemplation of the bowerbird, of whom he writes: "A nineteenth-century naturalist once suggested that just as mammals are commonly divided into two groups, man and lower forms, all birds should be divided into two categories: bowerbirds and other birds." No creature in the animal world aside from man has gone quite so far in the creation of what must be called a culture, and Gilliard believes that arena behavior has been responsible for the bird's quite incredible accomplishments.
Arena behavior is rare. Of the world's million-odd animate species, not over a hundred so far known indulge in its patterned heroics. All authorities agree that the strenuous competition of the arena brings about a speed-up in the evolutionary process. Is it possible that arena selection has contributed not only to man's astonishing speed of development but also, as in his opposite number in the world of birds, to the evolution of culture as a substitute for sweeping horns, prestigious crests, and many-splendored tails?
The bowerbird is a zoological group with a few species in Australia but its greatest flowering in New Guinea, where Gilliard has done most of his work. Why New
Guinea should be the home of the world's nonhuman cultural champions, while on that lavish island the human species has restrained its own cultural accomplishments to such unremarkable activities as the collection like postal issues of other people's heads, must be written down, I presume, as an embarrassing evolutionary joke. But that New Guinea has presented us with the animal Acropolis none can dispute. Nor are the cultural wonders confined to birds who build playhouses.
Alfred Russel Wallace, Darwin's great contemporary and co-originator of the theory of natural selection, was one of the first European naturalists to explore the East Indies. While he never came on the bowerbird, he discovered the bird of paradise, and brought back two lively specimens properly to astonish English zoology. This most gorgeous of creatures, likewise an arena being, gives us more than a hint as to how natural selection started encouraging the aesthetic improvement of real estate.
In 1857, on a little island called Aru just off the New Guinea coast, Wallace encountered the species known as the great bird of paradise. He described its display: "The head and neck is of pure straw yellow above, and rich metallic green below. The long plumey tufts of golden orange feathers spring from the sides beneath each wing and when the bird is in repose are partly concealed by them. At the time of excitement, however, the wings are raised vertically over the back, the head is bent down and stretched out, and the long plumes are raised and expanded till they form two golden fans, striped with red at the base. The whole bird is then overshadowed by them, the crouching body, yellow head, and emerald green throat forming but the foundation and setting for the golden glory that waves above."
Now, the temptation to regard such a display as a purposeful performance is very nearly irresistible. But had Wallace proceeded farther into the lowland rain forests of New Guinea proper, he might have encountered another species, the magnificent bird of paradise, which at the height of a comparable display opens wide its mouth to reveal the pale-green lining. And what we must keep firmly in mind is that Diphyllodes magnificus has never seen the green lining of his own throat. He performs the display by innate command. We may interpret -- wrongly -- all the remainder of the bird's complex performance as one of pur-
pose motivated by a desire to show off his breathtaking finery. But when the pale-green lining of his mouth comes along, we must recognize that since the bird possesses no a mirror he cannot know what he is showing off. The final action, like the entire display, is a form of innate behavior as much a part of the bird as is the finery itself. The bird of paradise comes in one evolutionary bundle.
Having reminded ourselves of this, we may move on to the magnificent's cultural accomplishment. And I recognize that if the natural selection of the arena can bring about a bird who opens his mouth to display a green lining he has never seen, then it is not too great a wonder that he I has developed an innate concern for the arrangements of I his court. Nevertheless, I am pleased not to be reporting I an observation of my own. Let Austin Rand, of the Chicago Natural History Museum, be the Munchausen.
The trade winds deposit on some New Guinea lowlands a rainfall of ninety inches a year. The consequent rain forest is profoundly shadowed. Some species of the bird of paradise hold their courts in the treetops, others in the middle branches. But two, the magnificent and the Queen Carola's, have their courts either on the ground or very near to it in a dark tangle of vines and branches. And Rand has described how the magnificent will spend hour after patient hour in the branches high above, snipping away leaves, so that a shaft of light may penetrate the shadowed forest, reach his court, and illuminate to best advantage the| iridescence of his colors.
Among the cultural wonders of New Guinea the next evolutionary step beyond arrangement of the stage lighting, as Gilliard analyzes it, is construction of the stage itself. These display stages are what early travelers in New Guinea found along forest paths and assumed were little houses built by native children. They were the bowers, of course, that have given the bowerbird its name. A dozen species build the little decorated houses, some complete with roof, as stages for display. They are never nests, but like nests the appearance and construction varies between species. Within a kind it is always the same.
Archbold's bowerbird, for example, is a relatively primitive species in cultural achievement. He clears a small stage in his high mountain forest and carefully carpets it witt fern. Then he decorates it with snail shells, dead beetles lumps of charcoal, and other attractive bric-a-brac. He
waits, perched well above his sidewalk display. If a female comes by, he drops to the middle of his stage, crouches, makes begging sounds, crawls toward her. Should she be unimpressed, she will move on to another Archbold's bower-bird. The rejected one will rearrange his ornaments and resume his wait like a patient Arab.
At the other and more evolved extreme is the gardener bowerbird, frequently referred to as a maypole-builder. He leans sticks against a sapling in the shape of a teepee, adding more and more until the structure may be taller than a man. The most advanced species, the crestless gardener, will create internal chambers in his tower, then top it with a broad roof against the forest's heavy rains. In the cleared area around the tower he plants moss for his dancing stage. He will decorate his little lawn with shells, berries, and piles of cut flowers.
Lauterbach's bowerbird is another advanced member of the family, but he uses an entirely different construction scheme. He is classed as an avenue-builder. Instead of building a round house supported by a central sapling, he plants two sticks firmly for support and constructs a four-walled structure. Gilliard examined one of these bowers and found 1000 pale pebbles used to pave the stage, over 3000 sticks in the rigid, interlocking construction of the bower itself, and 1000 strands of grass used to line the interior walls. At least sixteen such parlors built in a spread-out area on the edge of New Guinea's grasslands were attended
each by a member of a single clan waiting for susceptible females.
Gilliard waited too, and on three occasions he had the luck to be present when a female came by. The male danced with excitement on his pebbled pavement. She entered his bower. Now enormously excited, the male picked up a bright red berry, and just as the ruff displays his intricate collar or the kob his glorious throat, the bowerbird displayed the berry for his intended's delight.
Now again, it is very difficult for the human mind, with all its preconceptions, to resist the conclusion that Lauterbach's bowerbird has done all this purposefully for the sake of getting the girl. But we must remember the magnificent bird of paradise who displays the green lining of his mouth without knowing he has one. This is innate behavior specific to the species. Lauterbach's bowerbird builds an elaborate house on the edge of the grasslands, paves it with pebbles, and displays his red berry because he has substituted culture for a green throat and a fancy crest. Like the male member of any arena species; he competes with other males, by inward compulsion, to achieve status. In his particular species to achieve that status he must gain and defend a piece of property and improve and display it in a particular fashion. As a key fits a lock, so his success in a demanding masculine competition turns something in the female heart. It attracts her. Those of the species who have failed in the basic competition, who are not one of the sixteen members of the club dancing before their sixteen houses, are somewhere off in the woods, sexually discarded. But even for this member of the exclusive club, the key may not turn all the way, the red berry may not do, her sexual desire will not be aroused, and she will go elsewhere. In that case he can do nothing, for to do anything now would be to violate club rules. As the kob looks off to the mountains, as the ruff collapses on his hill, he must go back to his bower.
Among all these species the satin bowerbird is the champion's champion, since he not only builds a house but paints it. I shall defer this ultimate animal, however, to a more appropriate place in the story: not because he is an Australian instead of a New Guinean, but because I believe my earlier question has been answered. Man is not, has never been, and can never be an arena species. We lack the biological morality.
There is probably a sound enough physiological reason why we cannot be creatures of the arena, whatever our playful indulgences, for it is unlikely that the human female, unassisted, can raise our long-maturing offspring to independence: the cock cannot be spared to lead a cock's life in a cock's flock. In no arena species which we have inspected or which has ever been studied do you find other than disposable males. But it is in the realm of comparative behavior that arena species reveal an evolutionary truth little known until today but salient to our inquiry into the territorial imperative.
For the century or so since Origin of Species you and I and our fathers have been taught that the animal's only motivational pole is self. It is what we have meant by the struggle for existence and the survival of the fittest. The animal struggles to survive: to eat, to avoid death, to mate, at the most to nourish and protect his offspring. He inhabits a world of self, and all of his instincts are organized to promote his success in that narrow corridor. We may call it Ego, or we may call it jungle law. What has seemed inconceivable is that evolution could encourage any physical or behavioral trait running contrary to the interest of the individual: such behavior could have no survival value.
It is this understanding of evolution which has brought such puzzlement, and sometimes despair, to human thought. No informed mind today denies that man is descended by slow but explicable process from the world of the fish and the frog, the bird and the lizard, the monkey and the ape. No mind, informed or otherwise, denies that man is capable of moral conduct, of choice between right and wrong, of action directed at goals taller and more luminous than self alone. How, then, did we get that way? If evolution with its struggle for existence can provide other animals with behavioral equipment necessary only for individual survival, then by what evolutionary bootstraps has man raised himself to the human condition?
No more impenetrable conundrum has faced human thought in the past century. We have come to a variety of answers. For excellent reasons, many of us have clung to divine intervention and the uniqueness of the human soul to explain the otherwise inexplicable. And, as I have already mentioned, there have been many of us who have denied that man has instincts to impede his conduct, or who have asserted man's new independence from the 
evolutionary process. Some have shrugged, skipped the problem, and simply described man as the moral or ethical animal, a description furnishing no explanation at all. And there have been those, of course, who have seized on our classical understanding of Darwinian evolution to justify, through its jungle law, man's every basest action..
I shall submit what I regard as the most pressing argument in these pages: that our earlier understanding of evolution is false. As I weigh the evidence gathered by the new biology in these remarkable years, I can discover no qualitative break between the moral nature of the animal and the moral nature of man. Evolution has been as ready to equip the animal with innate behavioral commands restraining the interests of the individual on behalf of larger or more immortal goals as it has been ready so to equip the human species. In the case of the arena species, indeed, evolution has gone further than with ourselves, and that is way I have begun our inquiry with a form of behavior so rare.
Whether we turn our regard to the formal antics of the kob's stamping ground, the sexual pandemonium of a sage-grouse strutting ground, or the quiet, preoccupied community of a bowerbird clan, we are watching the consequences of an order so moral -- or a moral order so foreign -- as to fall beyond human comprehension. Toward a single biological end, a high degree of selection of male genes within a population, individual interest is channeled or inhibited. The female, despite all sexual heat, is not free to mate with the first male she encounters, but must seek a proven member of the elite. The defeated male is not free to wander off and try his luck on another mating ground, but, tied by mysterious cords to his home community, must accept psychological castration and join the ranks of the discarded. Even the elite, the one in ten a or twenty or thirty or forty, must obey the sexual rules and regulations and offer no interference with the final workings of female choice. Were any portion of the innate code governing the sexual activity of an arena species to be disobeyed or disregarded, then the arena itself would fall into genetic nonsense. Yet no part of that code corresponds to our conventional view of animal instinct.
The key to that code, I believe, is territory, and that is why I have called this book The Territorial Imperative. I cannot claim that these observations of a few highly 
specialized species can do more than suggest that our century-long interpretation of evolution may be wrong. But I believe that as we go along we shall see unfolding a mass of evidence that through natural selection evolution is capable of fostering inborn traits, as Moffat anticipated, restraining the individual to the ultimate benefit of his species. I see no reason to regard this as other than a biological morality. Nor do I find reason to believe that territory is other than the chief mechanism of natural morality, something more than a mere behavior pattern, more than an open instinct, more than a superb defensive instrument -- in truth, a natural mediating device between the good of the one and the good of all.
So far as man is concerned, we have our territorial moralities, as I shall attempt to demonstrate. But they are not of an arena sort. Were we an arena species like the bowerbird, and our capacity for the rapid creation of cultures and assimilation of civilization a product of this form of evolutionary speed-up, then life would not be quite as it is. If a man failed to produce some minimum standard of culture or failed to behave according to some minimum standard of civilization, then he would find himself sexually impotent and sexually uncaring. Women, on the other hand, would find themselves incapable of intercourse with other than civilized men. And if not enough men succeeded in the competition to furnish numbers sufficient to service all our women, then we should be in a handsome pickle indeed. Most of our women, like surplus sage hens, would have no one to sleep with. And the population explosion, along with a good many other problems, would be removed from the list of contemporary concern.
I shall not deny that it might be an excellent idea for Homo sapiens to become an arena species, for then we should proceed like a genetic rocket toward our rendezvous with Utopia. But like so many comparably excellent ideas, it falls victim to a natural fact or two. We lack the particular biological morality. We lack that innate command which shapes our sexual impulses to our whole selective good. And so we must press on with our search for natural advantages more a portion of the human resource. And if we remain convinced that the line between man and the animal is that we are capable of moral action, whereas the
animal is not, then we must reflect on the mystery that a few arena species possess a sexual morality that man can by no means claim.