by Guy Murchie
The story is essentially the same for animals who hunt animals, even though they're not strong on culture, and it's hard to think of any wild animal species that has not either benefited by predation or suffered from its denial. This understanding of predation, however, was almost unheard of in the early part of this century, so it was only natural that a goodhearted pioneer of conservation like President Teddy Roosevelt would try to help the deer in Kaibab National Forest in northern Arizona by ordering the shooting of pumas and coyotes found preying on them. The rangers reported that there were about 4000 deer in the area in 1905 and the President was glad to hear that their numbers steadily increased in the next few years, as the pumas and coyotes were eliminated. But by 1920 when the deer population reached 60,000 and the forage had been grazed very short, serious doubts began to be heard. And in 1924 when the numbers got to 100,000, the deer were dying of starvation everywhere under disastrous conditions and Kaibab was turning into a desert. Even with the population down to 40,000 again by 1926, there were still ten times too many deer and it took another couple of decades before the dawn of the science of ecology and a new evaluation of pumas and coyotes eventually got things back to a reasonable balance.
An even more dramatic example of predation, and one better documented, is Isle Royale, the 45-mile-long wilderness sanctuary in Lake Superior which, being separated by fourteen miles of open water from the nearest land on the coast of Ontario, is relatively uninfluenced by mammalian migration. Before this century, as far as anyone knows, the inhabitants of the island included neither moose nor wolves, but around 1908 some moose, who are excellent swimmers, evidently swam out there to get away from wolves on the mainland, and by 1915 the island was home for an estimated 200 moose. From then on the unmolested moose multiplied steadily and numbered several thousand all through the 1920s, but inevitably ate up so much vegetation that by 1930 they were running out of it, getting scrawny and diseased and soon were starving in large numbers. The herd of about 800 moose that survived continued in pitiful, semistarving condition through the thirties and forties until the very cold winter of 1948-49 when a pack of about 16 timber wolves came out to Isle Royale on the ice and, finding easy pickings among the weakened moose, stayed there. Then, after a few years in which the wolf pack grew slightly to about 20 while the moose were reduced to about 600, the two species established what ecologists call a predator-prey equilibrium of around 30
moose per wolf. By this time the oldest and sickest of the moose had been culled and the browse had had time to recover enough so the moose herd was better fed and healthier than at any time in the half century it had lived on Isle Royale. In terms of what is called the food pyramid, .8 tons of wolves were living on 45 tons of moose who were living on 2900 tons of foliage. Or, you could say, the average wolf was eating his own weight in moose (or other game) every week plus, indirectly, more than sixty times that much in fodder spread over at least ten square miles of land.
The significant thing about this relationship, of course, is that the eater and eatee do not compete to finality as species, for they need each other. They are playing a continuous game which, like all games, has rules. And a prime rule for the predator is: he kills only what he needs to sustain him. He definitely cannot afford to exterminate the prey or even make the prey scarce, for that would be eliminating his own food supply. It would be suicidal. No, he is basically a conservationist. He has to be - something like a farmer who prunes his crop and reaps it but makes sure he can raise another harvest next year.
Being a conservationist, however, does not mean that a wolf or lion ever consciously moderates his hunting or holds back from killing with a thought for tomorrow or next month. He doesn't need to exercise such restraint because killing is just not that easy. In fact on Isle Royale the records of many years show that, whenever a moose at bay stands his ground, wolves give up in a few minutes. They know from experience how deadly accurate can be his kicks, so they shy away to look for easier prey elsewhere. And only one moose out of every eight who flees actually gets caught and killed by them, always a weak one: very young, very old or crippled in some way. All in all, I am told, only one wolf attack out of thirteen upon moose results in a kill. And in other parts of the world the "batting average" for lions, hyenas, leopards and other predators, including hawks, is about the same.
Predation has a curious balance about it and countless complexities. The animals on both sides know when the hunt is on - and off. A herd of gazelles will graze peacefully within fifty feet of lions, because they instinctively know the lions are well fed and sleepy. But as soon as they sense a certain tension - perhaps just the twitch of a tail or a whiff of adrenalin imperceptible to man - they will be off for the horizon. This is perfectly natural, like breathing, to animals - something prey take for granted and practically never concern themselves about except at those relatively rare moments when action is vital.
TERRITORIALITY
But if predation takes an animal's full attention only occasionally, other of his relationships can be much more continuously demanding. I mean his need to defend his territory (if he is a territorial animal) against rival groups of his own species, with each of whom he must hold a polarity relation of victor-vanquished, the end he gets depending on whether he is winning or losing ground. And there is also his need to hold or improve his individual rating (with its similar polarity) in the social order of his own group. These two relationships naturally often overlap and, on occasion, can be virtually synonymous.
The need for territory, a place of one's own, or for privacy for one's family or community is of course a very ancient and fundamental need. Some say it is an evolutionary force older than sex. And it affects nearly every creature from microbes to men, from the coral fish on the reef to the song sparrow in the meadow to the pet dog who barks at strangers from behind the fence his master built to define his own (and the dog's) property limits to those same strangers. About the only creatures nor territory-minded are ones who migrate continuously, such as the grazing animals who have to move to live. But their vegetal wanderings could in some cases be construed as a weakness and it may be the browsing gorilla's territorial vacillation that primarily has led to his tenuous position in evolution, one apparently nearing extinction. If indeed you think the territorial instinct is less compelling than, say, the mating instinct, just compare the number of men who have died to save their wives with the number who have died to save their lands. And consider whether it's as hard to get a woman as to get a home to put her in.
Having property of your own, though it requires a lot of attention and can lead to strife now and then, unquestionably does reduce conflict. It works - and is the way of least friction and simplest motivation. Almost everywhere in the world customs and laws support the private property system, even to a degree in communist countries - presumably out of practical necessity. And, as we have seen, animals and lesser creatures accept it as part of nature for the same reason. Most territories are simple, circular and more or less two-dimensional because they are created by surface-dwelling creatures like coyotes, turtles, and cockroaches, but birds and climbers extend them up into trees or cliffs, badgers and burrowers down into holes and swimmers into rivers and seas, They are apt to be formal, sometimes even luxurious in an animal kind of way. One occupied by a pair of African birds, for example, was found to contain: a nest, 2 sleeping roosts, a drinking place, a bathing pool, 3 feeding areas, a food store, 6 excretion dumps, 7 border stations for calling or singing, a rubbing post and a special boudoir for preening - altogether a lot more than the accommodations of any of the human inhabitants of a nearby village, including the chief.
Territorial shapes obviously must fit their environs. That of a city dog has to be latticed like the streets, a sewer rat's is linear like the sewer, a hippo's gourd-shaped with the narrow part on the riverbank (where competition is keen) and spreading inland. Although the boundaries often shift and overlap as well as being generally invisible to human eyes,
they are reasonably distinct to the creatures concerned: marked by the songs and displays of birds, the excreta and spoors of insects and mammals. They are thus semi-abstract biocrystal networks covering much of the habitable world, a different coarseness and texture for each kind of animal.
A pride of four or five lions in Africa tends to maintain a territory of approximately five square miles, so each lion uses on the average about one square mile of grazing land to provide him with food. The leopard needs three times as much. Stone-age man (relatively slow and feeble 10,000 years ago) needed eight times as much. And bigger animals, even those without a definite territorial system, suffer seriously without vast domains, so much so that they seem genetically restrained by geography on anything less than a heroic scale. Elephants in their natural state collectively require the sweep of a continent for their full evolution, and the tiger is unable to survive on any island smaller than about 2500 square miles, Bali (90 miles long) being the smallest island on Earth with an indigenous population of tigers. Curiously, smaller islands have the effect of dwarfing individual animals (in stature if not in population density) as in the case of Shetland ponies, the deer of the Florida keys or the little gray fox of Catalina. It works even down to the dimensions of an aquarium tank, for mature electric eels will grow more than an inch a day when suddenly put into larger space. Short-legged animals naturally need less land than longlegged ones, so the warthog is satisfied with a quarter of a square mile and the woodchuck with an acre. Most varied of all are the territories of birds, which range from the 40 square miles of a pair of golden eagles to one tenth of an acre for warbiers in time of plenty, while terns on volcanic Isla Raza in the Gulf of California, when brooding (not foraging), crowd together so densely they average a nest to every square foot!
An important reason for the territorial system is that, despite a few scouts and pioneers, the great majority of creatures (including man) tends to resist going out into the sparsely inhabited lands of opportunity, preferring instead the traffic and excitement of daily competition with close neighbors in a crowd. It's the lure of Broadway over the drear of the boondocks. The French ethologist Jean-Jacques Petter labeled it the noyau or society of inward antagonism, which, like an alloy, gains strength from the internal stress of disparate elements. It feeds on the paradoxical psychology of "I can't live with you - but I can't live without you" uttered by a bickering, loving couple. It is why Arabs delight in living among Arabs they can quarrel with more exultantly than with Jews or other foreigners. It builds on the stimulus of friction and is thrilling to citizens who like to outshine their neighbors, be heroes to their families, flaunt their possessions, yell names, argue, bet, sue. A study of juvenile street gangs in Chicago reported that a call for volunteers to steal a car usually brings out half the gang's members, while a summons to fight invariably brings them all. There is something about a brawl. It has almost the shock value of a fire in getting everyone's instant attention, especially when you know it is to decide issues as important as the border between you and your enemy. It combines the fascination of war with the masculine attraction of rivals for each other and the reconciling by tooth or treaty of opposing pressures into a boundary settlement both sides can live with.
If you find it hard to believe that the territorial competition of males produces measurable pressure, you should be interested in the experiment conducted by the U.S. Fish and Wildlife Service a few years ago to test it by killing all mated male birds in a 40-acre section of Maine woods. For, to everyone's surprise, almost every male vacancy they created was filled so quickly by a new and eager male that the hunters could not keep up with the influx of substitutes and, although they kept shooting all summer until male birds temporarily became scarce, the number that showed up the following spring was substantially the same as ever. Thus territorial pressure is maintained by flexible hidden reserves behind the frontier, this being, I assume, a key factor in its stability.
PECKING ORDER
If territoriality and its polar stresses create a vital stability, primarily between rival males, there is at least as great a polarity and perhaps an even more pervasive competition for rank within each social unit or family. This is the phenomenon widely known as the pecking order, because pioneer ethologist Schjelderup-Ebbe noticed that a flock of chickens maintains natural harmony between its members by keeping every bird aware of who may peck him if he doesn't look out, as well as whom he may peck if he can. It has some of the quality of a very stable, regular crystal, yet is a hierarchy of strength unwittingly balanced by relative intimidation spiced with occasional all-out fighting when the flock was formed. Although the proud top chicken outranks all the others and can peck anyone without fear of retaliation, and the shy bottom chicken is disdained or bullied by all, both are accepted members of the integrated flock, all the rest of whose members have at least one they may chase and at least one they must avoid. There is a natural discipline to it, as in a military organization, where the top general may command anyone below and every lowly private may be commanded by anyone above - and they all get along together in reasonable harmony.
It is not always that simple though, for the pecking order in one activity, say building a nest, can be different from that in another, say feeding. And the boss animal in hunting may well be forced to wait for others when it comes to taking a drink. Also population pressure can disrupt the order of rank, aging tyrants seldom retire gracefully and mating elevates a female to the level of her "husband." In fact the young female who "marries the boss" may suddenly find herself in a position to snub former female superiors and lord it over males who have been pushing her around for years, for now she naturally stands with her husband, and he with her, in any dispute. Interestingly, this kind of loyalty invariably seems to prevail between animal mates, for the pecking order is in practice definitely superseded by the marriage bond. However the "single standard" that enables a highranking male to raise the status of his humble bride does not work the other way, for a humble male just doesn't get infatuated by a bossy female and, as Konrad Lorenz declared in his report on jackdaws, "no male may marry a female who ranks above him."
Did you realize that even the gentlest adult humans have a comparable ranking order that can assert itself in action as innocuous as a glance? When two strangers look at each other for the first time, for instance, is it significant that one of them averts his eyes before the other? Yes, an experiment conducted by psychologist Brian Champness for the British Association for the Advancement of Science found that the more dominant person, whether a man or a woman, almost always looks away first. In fact it is a reliable signal that he is about to claim the floor. And test scores show that the signal is invariably accepted by the more submissive one and, from it, human rank can be established, even though, in some cases, the persons involved are not aware of it. Champness took great pains to check his findings with other kinds of tests and devised a scale in which {1} equals a complete hierarchy (everyone knows whom he dominates and who dominates him) and {0} equals no hierarchy (no one knows his place). On that scale a flock of chickens rated .9 and Champness's group of humans rated .8, but established their hierarchy in only half the time chickens usually take.
So, you see, all of these three types of conflict we have been discussing - predation, territoriality and pecking order - involve violence or threats in various degrees, but in each case what you might call the stable aspect of the well-balanced crystalline structure is promoted either by shortening the agony, lowering the batting average or turning the confrontation into a ritual through which the issue can almost always be settled without bloodshed. Ritual conclusions, in fact, are the rule for conflicts within, and often between, groups of animals and men, though the exceptions naturally attract more, attention. A typical encounter between two rival timber wolves in the same pack often starts off so furiously with bared fangs, snarls and lightning snaps at each other that any observer might think one of the two must surely get killed. But when the weaker w
olf eventually tires or becomes careless and the stronger takes advantage of him, something incredible happens. Instead of desperately defending himself, the losing wolf suddenly surrenders. He seems to know the game is up, so he does the exact opposite of what you might expect. No longer facing teeth with teeth, he turns his head away, exposing the vulnerable bend of his neck to his enemy. Now the victor with one bite could sever the loser's jugular vein and kill him, but he does not. Actually he cannot. A powerful inhibiting force stops him. It is mental, but more inviolable than a white flag or a red cross to a human, and it nicely resolves the polarity paradox within a species, avoiding a serious loss of life, which would have had no redeeming evolutionary benefit. Lorenz indeed won the Nobel Prize for elucidating this and other hard-to-understand aspects of animal behavior, many of which have to do with polarity.
POLARITY IN EVOLUTION
Polarity shows up frequently in evolution. We see it everywhere from the genetic mutation that is commonly regarded as destructive or "evil" except on the rare occasions when it turns out "good" to the complement known as sex that assures the mutual polar attraction of opposites. The latter of course produces the variation of offspring without which evolution could not select and evolve. And there are creatures who seem to resist evolution, such as a simple beach animal called the pre-Cambrian sandworm, who adapted to the young Earth's shore environment 600 million years ago so perfectly that he hasn't had a measurable need to evolve farther in all the eons since. In fact he attained an equilibrium with the planet perhaps more successfully than any other creature, including man, who, incidentally, has existed less than a hundredth as long and is already in such a precarious crisis that some prognosticators give him only an outside chance of survival for even one more millennium. But the sandworm's very success is paradoxical because he has a built-in polarity between stability and instability, between peaceful harmony and distressing strife. For there are many species of sandworms and the most successful ones, which stopped evolving, eventually stagnated in an evolutionary dead end, while some of their unsuccessful, poorly adapted cousins continued being unstable, restless and dissatisfied and, for that reason, went on trying out new forms and behaviors and eventually evolved into such unprecedented classes as reptiles, birds and mammals - demonstrating that failure to adjust in early evolution may be just what is needed for success later on, that stress and strife are ingredients of long-range harmony, that pain is vital to birth and creation.
