NOTES ON THE QUESTIONS OF PROTECTIVE COLORATION, RECOGNITION MARKS, AND THE INFLUENCE OF ENVIRONMENT ON LIVING ORGANISMS Harmony of colour in nature—Theory of protective coloration—Sexual selection—Conspicuous colours not harmful—The influence of environment—The leucoryx—The Barbary sheep—The Sardinian moufflon—African butterflies—Coloration of the musk ox and caribou—Arctic hares and foxes—Coloration of mammals in the Yukon Territory—The chamois in winter—Examples of conspicuous coloration in African mammals—Colour not always protective—Carnivorous animals usually hunt by scent—Wild dogs and wolves—Wild dog and sable antelope—Sense of smell in herbivorous animals—Sight of antelopes—Experience with waterbuck—Dull sight of caribou—Demeanour of wild animals when alarmed—Small antelopes—Lions—Large antelopes—Difficulty of seeing wild animals sometimes exaggerated—Powers of sight of Bushmen—Colour not protective against animals which hunt by night and by scent—Animals in motion easy to see—Restlessness of wild animals—Lions attacking bullocks—Zebras the principal prey of lions since the disappearance of buffaloes—Appearance of zebras—Undoubtedly conspicuous animals in open country—Zebras by moonlight—Strong smell of zebras—Conspicuous antelopes in East Africa—Effect of the juxtaposition of black and white—Bold coloration of the sable antelope. Although there are certain striking exceptions to the general rule, yet, broadly speaking, it cannot be gainsaid that living organisms are usually coloured in such a way as to make them difficult of detection by the human eye amongst their natural surroundings. Every collecting entomologist knows how In the open treeless regions within the Arctic Circle, as well as on bare mountain ranges, nearly all the resident species of animals and birds turn white in winter, when their whole visible world is covered with an unbroken mantle of pure white snow, and become brown or grey during the short period of summer. In treeless deserts again within the tropics, where the rainfall is very scanty and the climate excessively hot and dry, with intense sunlight throughout the year, all resident living organisms, mammals, birds, reptiles, and insects, are found to be of a dull coloration which harmonises in the most wonderful way with the sandy or stony soil on which they live. It is also very often the case that animals which live in forests where the foliage is not too dense to allow the sun to penetrate are spotted or striped, whilst those which live in really thick jungle or amongst deep gloomy ravines are of a uniform dark coloration. Now a most interesting question arises as to the true causes which have brought about the extraordinary variations of colour to be seen in living organisms inhabiting different parts of the world. It is, I believe, the general opinion of modern naturalists that, putting aside cases where brilliant colours may have been produced amongst birds and insects by the action of the law of sexual selection, the coloration of all living organisms is protective, "serving," as that distinguished naturalist Mr. Alfred Russel Wallace puts it, when discussing the Many very striking facts can be adduced in support of this theory, and no doubt it is of advantage to most species of mammals, birds, reptiles, and insects to harmonise in colour with their surroundings; but there are many instances in nature, especially amongst birds and insects, where a very striking and conspicuous coloration does not appear to have been prejudicial to the life of a species. The highly decorative but very conspicuously coloured plumage to be seen in the males of many species of birds, especially during the breeding season, was considered by the immortal Darwin to be due to the influence of sexual selection, and whatever may be urged against the correctness of this theory, it is supported by a long array of indisputable facts. Great, however, as is the divergence between the plumage of the males and females in many species of birds, not only during the breeding season, but in a great number of cases at all times of year, and however gaudy and conspicuous the coloration of the former may be compared with that of the latter, such conspicuous coloration never appears to be prejudicial to the life of a species, though in some cases the brighter coloured male assists the female in incubation, and it would thus appear that in all such cases the sombre coloured plumage of the female was not absolutely necessary for purposes of protection against enemies. I therefore think that if it is admitted that bright and conspicuous colours have been evolved in living organisms through the action of the law of sexual selection, without detriment to the life of the species It is worthy of remark, I think, that in hot, dry deserts, where the climatic conditions are stable, and where the general colour of the landscape is therefore very much the same all the year round, all the resident species of mammals, birds, reptiles, and insects are what is called protectively coloured, that is to say, they are all of a dull brown or greyish coloration, To my mind the loss of stripes in the quagga was entirely due to the environment in which this species had lived for long ages; for on the karoos of the Cape Colony everything is of one dull brown colour, whether on hill or plain, and no shade is to be found anywhere, for the whole country is without trees. The air, too, is intensely hot and dry, and the rainfall scanty. In these semi-deserts of South-Western Africa, not only did the quaggas lose their black stripes, but the elands also lost the white stripes of their immediate ancestors, whilst the blesboks had already lost much of the white to be seen in the body colouring of the bonteboks, from which they are descended, and had become of a much duller colour generally. In East Africa, however, the plains are surrounded by well-wooded hills, which give some colour to the landscape, whilst the rainfall every year is heavy. If it is not the influence of their several environments which has brought about the differences between the well-striped elands and zebras of East Africa and their dull-coloured relatives that once lived in the karoos of the Cape Colony, the theory of protective coloration must be equally at fault, for in spite of the fact that in both countries both races of these animals have been hunted by lions from time immemorial on open plains, and under precisely similar conditions, they developed very different schemes of coloration. The Barbary sheep, again, which inhabits the dry The male moufflon of Sardinia, which lives in a temperate climate where the colours of its surroundings are much brighter and more diversified than is the case in the habitat of the Barbary sheep, is a much more conspicuously coloured animal than the latter, or than the females of its own kind. As the females and young of the Sardinian moufflon, which are of a uniform brown colour, are more difficult to see than the males in their somewhat conspicuous autumn and winter coats, the latter cannot be said to be protectively coloured. Either through the influence of sexual selection or that of an environment the general colour of which varies very greatly at different seasons of the year, the male of the Sardinian moufflon becomes during autumn and winter conspicuously coloured compared with the female, without detriment, however, to the well-being of the species. During my long sojourn in the interior of South Africa, I made large collections of butterflies. There was one species (Precis artaxia, Hewits) which always puzzled me. This handsome insect is only found in shady forests, is seldom seen flying until disturbed, and always sits on the ground amongst dead leaves. Though handsomely coloured During the rainy season in South Africa, the open glades in the forests bordering the rivers are gay with multitudes of brightly coloured butterflies of many different species, and after a night's rain butterflies of various kinds may often be seen settling in masses round pools of water along waggon roads. Most of these butterflies are conspicuously coloured, though they are in perfect harmony with the sunlit flowers which spring up at the time of year when they appear. I cannot, however, believe that the need for protection against birds or other enemies has had anything whatever to do with the determination of their various colours, as in all my experience (and I have been all my life a close The coloration of certain animals in the Arctic and sub-Arctic Regions is somewhat remarkable, as at certain seasons it is conspicuously out of harmony with its surroundings, and cannot therefore be protective. The musk ox retains its dark brown coat the whole year round, although it lives almost constantly amidst a snowy environment. Mr. Wallace tells us that the reason why the musk ox does not turn white is because it has no enemies to fear, and therefore has no need of a protective coloration. He says: "Then we have that thoroughly Arctic animal the musk sheep, which is brown and conspicuous; but this animal is gregarious, and its safety depends on its association in small herds. It is therefore of more importance for it to be able to recognise its kind at a distance than to be concealed from its enemies, against which it can well protect itself so long as it keeps together in a compact body." As, however, according to the experience of Arctic travellers, large numbers of young musk oxen are annually killed by wolves, this explanation of a case in which an animal is manifestly not protectively coloured does not seem altogether satisfactory. Mr. Wallace, it may be noted, calls special attention to the coloration of the giraffe, which he considers to be protective; yet nothing, I think, is more certain than that a far smaller percentage of giraffes are killed annually by lions in Africa than of musk oxen by wolves in Arctic America. If this is so, the musk ox has more need of protective coloration than the giraffe. The musk ox is, I think, the only one amongst the few truly Arctic mammals which does not turn white during the winter months, for, unlike the barren ground caribou, it does not migrate southwards in the autumn to the dark spruce forests, which change Speaking of other Arctic animals, Mr. Wallace believes that the Arctic fox of necessity turns white in winter in order to enable it to capture the white Arctic hares upon which it chiefly lives. Very little, however, is known as to the life-history of these two animals. But if the Arctic foxes hunt by scent, as they almost certainly do, during the constant darkness of the long Arctic winter, and the hares burrow beneath the snow, and are caught as a rule when completely hidden from sight below its surface, I think it is arguable that the influence of environment During winter in the Yukon Territory, moose turn very dark in colour on the under parts of the body, and at this season of the year leave the thick forests and live in the comparatively open valleys amongst willow and birch scrub, where they are said to stand out like haystacks amidst their snowy surroundings. The local race of caribou (Rangifer osborni), which live all the year round on the treeless mountain plateaus, are very dark in colour (with the exception of their necks), and, as I myself can testify, stand out very plainly when the open ground they frequent is covered with snow. Of the various races of wild sheep inhabiting the mountains of Alaska, the Yukon Territory, and Northern British Columbia, some are white all the year round, and therefore very conspicuous in summer when there is no snow It would thus appear that in the sub-Arctic Regions of North America the coloration of mammals does not obviously serve the purpose of concealing the herbivorous species from their enemies, or of enabling carnivorous animals to approach their prey unperceived. To come nearer home, we find that whereas in the Alpine regions of Europe the mountain hare turns white in winter, the chamois living in the same snow-covered ground becomes deep black. It is true that in winter chamois often leave the open mountains and live amongst the higher forests, where it may be said that their dark colour harmonises well with the dark foliage of the spruce trees; but I have hunted chamois in December in the mountains of Transylvania, when they were in full winter coat, and I certainly found that their dark coloration often made them conspicuous. Turning to Africa, we have many instances of what seen in the open and at short range cannot possibly be called anything but conspicuous coloration, such as the jet black and pure white striping of the East African form of Burchell's zebra; the deep glossy black body and neck, with snow-white belly and parti-coloured face, of the sable antelope; To me it seems that the influence of environment might very well be deemed sufficient of itself to cause all animals that have lived for long ages in treeless deserts under constant strong sunlight to assume the dull brown coloration which they undoubtedly possess; whilst Arctic conditions might be expected to cause the whitening of an animal's hair in the winter, or the play of the sun's light through the leaves and branches of trees and bushes to be responsible for a spotted or striped coat. In the case of a combination of black and white—the two most conspicuous colours in nature—such as may be seen in the adult cock ostrich or male sable antelope, why should it not be supposed that the law of sexual selection has come into play, as it probably has done in the production of the lion's mane and the exaggerated size of the horns in the male koodoo. Having spent many years of my life in the constant pursuit of African game, I have certainly been afforded opportunities such as have been enjoyed by but few civilised men of becoming intimately acquainted with the habits and life-history of many species of animals living in that continent, and all that I have learnt during my long experience as a hunter compels me to doubt the correctness of the now very generally accepted theories that all the wonderfully diversified colours of mammals—the stripes of the zebra, the blotched coat of the giraffe, the spots of the bushbuck, the white face and rump of the bontebok, to mention only a few—have been evolved either as a means of protection In all recent articles which I have read by well-known naturalists on these subjects, it appears to be assumed that both carnivorous and herbivorous animals trust entirely to their sense of sight, the former to find their prey, and the latter to detect and avoid the approach of their enemies. Yet nothing is more certain than that all carnivorous animals hunt almost entirely by scent, until they have closely approached their quarry, and usually by night, when all the animals on which they prey must look very much alike as far as colour is concerned. The wild dogs of Africa and the wolves of northern latitudes are not so completely nocturnal, it is true, as the large Felidae, but the former I know, and the latter I have every reason to believe, hunt, as a rule, by night and only occasionally in the daytime. In both these animals the sense of smell is enormously developed, and must be of far greater use to them in procuring food than the sense of sight, however acute that may be. In all my wanderings I have only seen African wild dogs chasing game in the daytime on four occasions. I once saw a single wild dog chasing a sable antelope in the daytime. This wild dog—which was, however, then too far away to enable me to see what it was—first ran past the sable antelope and behind it from where I was watching. It must then have been running on the trail, with its nose on the ground, and must have passed quite close to the animal it was pursuing without seeing it. Its nose, however, kept it on the antelope's tracks and soon brought it But not only do all carnivorous animals hunt by scent, and rely far more upon their olfactory organs than upon their keenness of sight to procure food, but, as all practical hunters very well know, the sense of smell is also very highly developed in all, or at any rate in most, of the animals on which the carnivora prey, and personally I am persuaded that all browsing and grazing animals in Africa trust as much to their noses as to their eyes both to avoid danger and to find members of their own species. The eyes of antelopes are quick to detect a moving object, but they are by no means quick to notice any unusual colour in a stationary object. I will relate an anecdote illustrating this point. Early in 1883, I reached the spot on the Hanyani river in Mashunaland where I intended to establish my hunting camp for the season. Whilst my Kafirs were chopping down trees to build the cattle enclosures, I climbed to the top of the ridge at the foot of which I was having my camp made. It was late in the afternoon, and I was sitting on a rock looking over the open country to the south, when I heard a slight noise, and turning my eyes saw a fine male waterbuck coming towards me up the ridge. I sat perfectly still, and it presently walked slowly past within three yards of me and then went on along the ridge, into the forest beyond. As it passed me I noticed its shining wet nose, and the way in which its nostrils kept constantly opening and shutting at every step. It was evidently listening to the noise that my Kafirs were making chopping down small trees at the foot Of course, if I had made the very slightest movement, this waterbuck would have seen me instantly; but had it possessed much sense of colour, the contrast between the red brown of my sunburnt arms and face and the light-coloured shirt I was wearing would have attracted its attention, as I was sitting on a stone, on the top of a ridge which was quite free from trees or bush. I have never had any other African antelopes pass so close to me as this without seeing me, but many have fed slowly past me, as I sat watching them, with a tree or a bush behind me but nothing between myself and them, at distances of from 20 to 50 yards. Both in Newfoundland and in the Yukon Territory of Canada, I have had caribou walk almost over me when sitting in front of them on their line of march on ground devoid of any cover whatever. In such cases, of course, the wind was blowing from these animals towards where I was sitting, and I remained absolutely motionless. As a rule, when wild animals notice something suspicious approaching, say a man on horseback, and cannot get the scent of it, they run off before it gets near them or circle round to try and get the wind of it. But the smaller African antelopes, steinbucks, duikers, oribis, and reedbucks will occasionally, while keeping their eyes fixed on the unfamiliar object, crouch slowly down, and then, with their necks stretched along the ground, lie watching. I have ridden past a few oribis, steinbucks, and reedbucks within a few yards, as they lay absolutely motionless on the ground watching me. To pull in one's horse with the intention of shooting such a crouching antelope was the instant signal for it to jump up and bound away. Lions too, when they see a human being and imagine It is perhaps worthy of remark that I have never known a case of one of the larger antelopes trying to escape observation by lying down. Gemsbucks, roan and sable antelopes, elands, koodoos, hartebeests, indeed all the large African antelopes, directly they see anything suspicious, face towards it, and stand looking at it, holding their heads high, and not in any way shielding their bodies and only exposing their faces to view, which, when marked with black and white, as in the case of the gemsbuck and roan antelope, are supposed, though quite erroneously, to render these animals invisible. I am inclined to think, but it is only my personal opinion, that the difficulty of seeing wild animals in their natural surroundings has been greatly exaggerated by travellers who were not hunters, and whose eyesight therefore, although of normal strength, had not been trained by practice to see animals quickly in every kind of environment. I am quite sure that to a South-African Bushman Now, if carnivorous animals had throughout the ages depended on their eyesight for their daily food as the Bushmen have done, which is what naturalists who believe in the value of protective coloration to large mammals must imagine to be the case, surely their eyesight would have become so perfected that no colour or combination of colours could have concealed any of the animals on which they habitually preyed from their view. As a matter of fact, however, carnivorous animals hunt as a rule by scent and not by sight, and usually at night when herbivorous animals are moving about feeding or going to drink. At such a time it appears to me that the value of a coloration that assimilated perfectly with an animal's natural surroundings during the daytime would be very small as a protection from the attacks of carnivora which hunted by night and by scent. Reverting again to the question of quickness of eyesight, I will say that, although a Boer or an English hunter can never hope to become as keen-sighted as a Bushman, his eyes will nevertheless improve so much in power after a few years spent in the constant pursuit of game, that the difficulty of distinguishing wild animals amongst their native haunts will be very much less than it was when he first commenced to hunt, or than it must always be to a traveller or sportsman who has not had a long experience of hunting. However difficult an animal may be to see as long as it is lying down or standing motionless, as soon as it moves it becomes very apparent to the human eye; and, as I have had ample experience that any movement made by a man is very quickly noticed by a lion, leopard, hyÆna, or wild dog, I am quite sure that all these carnivora, if lying watching for prey by daylight, would at once see any animal moving about feeding anywhere near them; and all herbivorous animals move about and feed early in the morning and late in the evening, the very times when carnivorous animals would be most likely to be looking for game by daylight. During the heat of the day carnivorous animals are very seldom seen, as at that time they sleep, and most herbivorous animals do the same. But even when resting, wild animals are seldom motionless. Elephants and rhinoceroses are constantly moving their ears, whilst giraffes, elands, buffaloes, zebras, and other animals seldom stand for many seconds together without swishing their tails. All these movements at once attract the attention of the trained human eye, and I am very sure would be equally apparent to the sight of a lion or a leopard, were these animals to hunt by sight and during the daytime. But, speaking generally, they do not do so, though doubtless should antelopes or other As a rule, lions do not commence to hunt before darkness has set in. They then seek their prey by scent, either smelling the animals directly or following their tracks. They understand as well as the most experienced human hunter the art of approaching game below the wind, when hunting singly; but when there are several lions hunting together, I believe that some of them will sometimes creep close up to a herd of game below the wind, whilst one or more of their number go round to the other side. The buffaloes, zebras, or antelopes at once get the scent of these latter, and run off right on to the lions lying waiting below the wind, which then get a good chance to seize and pull down one of the frightened animals. As lions have played this game with my cattle upon several occasions, I presume that they often act in the same way with wild animals. No matter how dark the night may be, a lion has no difficulty in seizing an ox, a horse, or a donkey exactly in the right way, and I have no I have had innumerable opportunities of looking at wild zebras, and when met with on open ground they certainly have always appeared to me to be very conspicuous animals, except just at dawn and late in the evening, when they are not so easy to see as animals of some uniform dark colour, such as hartebeests. In Southern Africa, between the Limpopo and the Zambesi rivers, Burchell's zebras used to be very plentiful in all the uninhabited parts of the country, and although they were often met with feeding or resting in districts covered with open forest or scattered bush, I found them always very partial to open ground, where they were as plainly visible as a troop of horses. In East Africa the local race of Burchell's zebra is remarkable for the whiteness of the ground colour of the body and the intense blackness of the superimposed stripes. These beautiful animals congregate in large herds on the bare open plains traversed by the Uganda Railway, and probably form the chief food of the lions living in that district. When in East Africa a few years ago, I took special note of the appearance of zebras at different distances on the open plains between Lakes Nakuru and Elmenteita. I found that in the bright African When in East Africa I came to the conclusion that not only the zebras, but also the impala antelopes—which are of a much richer and darker red than in South Africa—were conspicuously coloured, To my eyes, and in the bright sunlight of Africa, the juxtaposition of black and white markings, so often seen on the faces of African antelopes, has never seemed to produce an indistinct blur of colour except at a considerable distance. At any distance up to 300 yards the black and white face-markings of the gemsbuck, the roan, and the sable antelope always appeared to me to be distinctly visible, and they have often been the first parts of these animals to catch my eye. It is all very well to say that a male sable antelope, in spite of its bold colouring, is often very difficult to see. That is no doubt the case, but that only means that there is no colour in nature, and no possible combination of colours, which at a certain distance, if stationary, would not be found to harmonise well with some portions of, or objects in, an African landscape. Speaking generally, however, the coloration of a sable antelope bull makes him a very conspicuous object to a trained human eye, and also, one would suppose, to that of a carnivorous animal, were it watching for prey by daylight. |