FURTHER NOTES ON THE QUESTIONS OF PROTECTIVE COLORATION, RECOGNITION MARKS, AND THE INFLUENCE OF ENVIRONMENT ON LIVING ORGANISMS Occasional resemblance of African mammals to natural objects—Hartebeests—Elephants—Giraffes—Coloration of the Somali giraffe—Giraffes not in need of a protective coloration—Koodoos and sable antelopes—Acute sense of hearing in the moose—Possible explanation of large size of ears in the African tragelaphine antelopes—Coloration of bushbucks, situtungas, and inyalas—Leopards the only enemies of the smaller bush-haunting antelopes—Recognition marks—Must render animals conspicuous to friend and foe alike—Ranges of allied species of antelopes seldom overlap—Hybridisation sometimes takes place—Wonderful coloration of the bontebok—Coloration distinctly conspicuous and therefore not protective—Recognition marks unnecessary—Coloration of the blesbok—The blesbok merely a duller coloured bontebok—Difference in the habitat of the two species—The coloration of both species may be due to the influence of their respective environments—The weak point in the theory of protective coloration when applied to large mammals—Hares and foxes in the Arctic and sub-Arctic Regions—The efficacy of colour protection at once destroyed by movement—Buffaloes and lions—General conclusions regarding the theory of protective coloration as applied to large mammals. Certain observations have been made and theories propounded on the occasional resemblance of African mammals to natural objects, which have never seemed to me to have much significance, although they are often referred to as valuable observations by writers on natural history. Thus it has been said that hartebeests, which are Well, hartebeests are red in colour wherever they are found all over Africa. Ant-heaps are only red when they are built of red soil. In parts of the Bechwanaland Protectorate, where the Cape hartebeest used to be common, the ant-heaps are a glaring white. In East Africa, in different portions of which territory hartebeests of three species are very numerous, all of which are bright red in colour, red ant-heaps are certainly not a conspicuous feature in all parts of the country, and there were, if my memory serves me, very few ant-heaps of any size on the plains where I met with either Coke's, Neumann's, or Jackson's hartebeests. I was once hunting in 1885 with a Boer friend As to the theory that the long neck and the peculiarly formed head of the giraffe have been evolved in order to protect this remarkable animal against its carnivorous foes, by giving it the appearance of a dead or decayed tree, I personally consider such an idea to be so fantastic and extravagant as to be unworthy of serious consideration. In the course of my own hunting experience, I have shot a great many giraffes to obtain a supply of food for my native followers, and under the guidance of Bushmen have followed on the tracks of many herds of these animals until I at length sighted them. As regards the coloration of the species of giraffe inhabiting South and South-Western Africa, it assimilates very well with its surroundings, when amongst trees and bush; but as giraffes spend a great deal of their time passing through open stretches of country on their way from one feeding-ground to another, they are often very conspicuous animals. With respect to the Somali giraffe (Giraffa reticulata), a photograph taken by the photographer who accompanied one of Lord Delamere's expeditions, showing some of these animals feeding amongst mimosa trees, gives the impression of a most marvellous harmonisation of colour and arrangement of marking with their surroundings. But I cannot help thinking that the facts of the case have been very much exaggerated in this photograph, I must say that I rather distrust the camera as a true interpreter of nature, as I have seen so many photographs of the nests of small birds in bushes in which it was very difficult even for a trained eye to find the nest at all, although in all probability it would have been comparatively easy to detect these nests in the actual bushes in which they were placed. Speaking of the Somali giraffe, Colonel J. J. Harrison, in a footnote to a photograph of one of these animals shot by himself right out in open country, which appeared in the Bystander for January 30, 1907, says: "These handsome coloured giraffes are very striking when seen standing in the sun. Of a rich bright chestnut colour, with pure white rings, they stand out splendidly as compared with the dull grey colouring of the more southern giraffe." However, it appears to me that to whatever extent the coloration of the various races of giraffes harmonises with their surroundings, that result must have been brought about by the influence of their environment rather than by the need of protective coloration, for I cannot believe that the struggle for life against the attacks of carnivorous animals can have been sufficiently severe to have influenced the Altogether, the theory that the colour of the giraffe has been evolved by the necessity for concealment and protection from the attacks of carnivorous animals does not seem to me to be at all well supported by the life-history of that animal as seen by a practical hunter; but the fact that the coloration of this remarkable animal assimilates very well with the dull and monotonous shades of the trees and bushes in the parched and waterless districts it usually frequents, is a strong argument in favour of there being a law which, working through the ages, tends to bring the colours of all organic beings into harmony with their surroundings, irrespective of any special benefit they may receive in the way of protection from enemies by such harmonious coloration. Turning to the striped and spotted forest antelopes inhabiting various parts of Africa, I think The range of the koodoo to the south of the Zambesi extends farther to the south and west than that of the sable antelope, but I think I am justified in saying that up to the time of the deplorable visitation of rinderpest in 1896, wherever, between the Limpopo and the Zambesi, sable antelopes were to be met with, there koodoos were also to be found, and outside of districts infested by the "tse-tse" fly, excepting amongst rocky hills, I have never met with the latter animals in any country where I was not able to gallop after them on horseback. Living as they do in surroundings so very similar to those frequented by sable antelopes, I have never been able to understand why koodoos should have such much larger ears than the former animals. I have never been struck with the acute sense of hearing in koodoos as I have been in the case of the moose of North America, and I should scarcely think that this sense would often save them from the noiseless approach of such animals as lions or leopards, to which they very frequently fall a prey, judging by the number of the remains of koodoo bulls which I have found that had been killed by the former animals. I have often wondered whether the large size of the ears observable in the African tragelaphine ante Antelopes inhabiting open plains are very gregarious, and in the daytime would always be able to find their mates by sight. I have never heard them making anything but low grunting noises. As it is often assumed by naturalists that all bush-haunting species of antelopes have very large ears, it is perhaps worth noticing that in the little blue buck and the red bush duiker of South-East Africa, which both live in dense jungle near the coast, the ears are very small; whilst in the steinbuck, on the other hand, which is always found in very open country and never in thick bush, the ears are very large—both long and broad. The coloration observable in the different races of bushbucks inhabiting different localities, as well as in the situtunga and inyala antelopes, is, I think, very interesting and suggestive. It may, I think, be taken for granted that all the races of African bushbucks have been derived from an ancestral form which was both striped and spotted; but in the bushbucks found near the coast of the Cape Colony and Natal, the adult males are deep dark brown in colour, often absolutely devoid of any white spots or stripes on face or body, whilst the adult females are yellowish red, with only a few Now the only animal that preys habitually on bushbucks, inyalas, and situtungas is the leopard, and as leopards hunt by night and by scent, I cannot believe that the very different outward appearance of the various races of bushbucks inhabiting different parts of Africa is to be accounted for by the theory of protective coloration. The males and females of the Cape bushbuck and of the inyala antelope are very different one from another in the colour of their coats, but this does not seem to be prejudicial to either sex, though there is absolutely no difference in their habits or their habitat. In all the different races of bushbucks, however, with which I am acquainted, the males are much darker in colour than the females, so that it is not so very surprising that in the case of the inyala and the Cape bushbuck the males should have been the first to lose their stripes and spots in a sombre environment. In the case of the Cape bushbuck the adult females have already lost all the stripes and most of the spots of the ancestral form. The female inyala is, however, one of the most distinctly striped and spotted representatives of the tragelaphine group. I cannot see that facts support the opinion that the uniform dull brown coloration of both sexes of the southern race of situtunga has been brought about for the purpose of protection from carnivorous enemies. During the daytime these animals live in the midst of beds of reeds growing in water where they cannot be approached except by wading; but at night they are often killed by leopards, and perhaps sometimes by lions, whilst feeding just outside the reed beds, on open ground which has perhaps been recently swept by a veld fire, and where young reeds and grass are just sprouting. I gather from the writings of Mr. A. R. Wallace and other well-known naturalists that, whereas the coloration of all animals is supposed to be due to the need of protection from carnivorous beasts, many species have developed in addition what are known as recognition marks, to enable them to distinguish members of their own species from nearly allied forms, or to help them to quickly recognise and rejoin the members of the herd or family from which they may have been separated. That many large mammals belonging to different genera, and living in widely separated parts of the globe, are marked with conspicuous patches of white on the rump, neck, or face, or throw up bushy tails when running, showing a large white under surface, is an indisputable fact, though it is not possible to say that the possession of such a conspicuous coloration is absolutely necessary to the well-being of any particular species, because there will nearly always be other species living in the same country, and subject to the attacks of the same predatory animals, in which these so-called recognition marks are absent. However, on the supposition that carnivorous animals hunt by sight, it seems to me that no animal can be said to be protectively coloured which is marked in any way so conspicuously as to be recognisable by others of its own Personally, I cannot see why large antelopes which live in herds on open plains should require special recognition marks, as in such localities the bulk of an animal's whole body would be plainly visible at a great distance no matter what its colour might be. If an antelope became separated from its fellows by night, all so-called recognition marks would be invisible at a very short distance. It must be remembered, however, that every species of animal has a peculiar and very distinctive smell of its own, and my own observations would lead me to believe that most wild animals recognise one another, as a rule, more by scent than by sight. It seems difficult to believe that there can be any truth in the theory suggested by Mr. Wallace, that recognition marks have been developed in certain species of large mammals because they are necessary to enable nearly allied species of animals to know their own kind at a glance, and so prevent interbreeding; for the ranges of very nearly allied forms of one genus, such as the various species of hartebeests and oryxes, or the bontebok and the blesbok, very seldom overlap, and so each species keeps true of necessity and without the help of special recognition marks. Where the ranges of two nearly allied species do overlap interbreeding probably will take place. There seems little doubt that the species of hartebeest known as Neumann's hartebeest has interbred with Jackson's hartebeest in certain districts where the ranges of the two species meet. In the neighbourhood of Lake Nakuru, in British East Africa, I shot, in February 1903, a hartebeest This animal (an adult male) was shot by my friend Cornelis van Rooyen in Western Matabeleland, where the ranges of the two species just overlap. In coloration it was like a tsessebe, but had the comparatively bushy tail of the hartebeest, whilst its skull and horns (which are, I am glad to say, in the collection of the Natural History Museum at South Kensington) are exactly intermediate between those of the two parent species. This skull has been very unsatisfactorily labelled "supposed hybrid between B. Caama and D. lunatus." But as, when I presented it to the Natural History Museum, I gave at the same time a full description of the animal to which it had belonged, which I got from the man who actually shot it, there is no supposition in the matter. If the skull and horns in question are not those of a hybrid between the South African hartebeest and the tsessebe, then they must belong to an animal still unknown to science. There is, I think, no large mammal in the whole world whose coat shows a greater richness of bloom and a more abrupt contrast of colours than the bontebok, so called by the old Dutch colonists of the Cape because of its many coloured hide, for bont means spotted, or blotched, or variegated. The whole neck, the chest, the sides and under parts of the head, and the sides of the body of this remarkable antelope are of a rich dark brown, and the central part of the back is of a beautiful purple Now, a practical acquaintance with the very limited extent of country in which the bontebok has been evolved, and where the survivors of the race still live, makes it quite impossible for me to believe that the extraordinarily brilliant colouring of this species of antelope can have been gradually developed in order to make it inconspicuous and therefore difficult of detection by carnivorous animals, nor can I believe that it has been evolved for the purpose of mutual recognition between individuals of the species; for although the snow-white blaze down the face or the white rump patch might very well subserve such a purpose, I see no necessity, looking to the habitat and the habits of the bontebok, for special recognition marks. Now, before proceeding further, I think I ought to say a word as to the points of resemblance and the differences between the bontebok and its near ally the blesbok. In the latter, the wonderful contrasts of colour to be seen in the former are considerably toned down; but the difference between the two species is merely superficial. The general body colour of the blesbok is dark brown, but not so dark as on the neck and sides of the bontebok, and the delicate purply lilac colour of the back in the latter species is altogether wanting in the former. In the blesbok, too, the colour of the rump just above the tail, which in the bontebok is snow-white, is brown, In a word, the differences between the bontebok and the blesbok are confined to the intensity of the colours on various portions of their hides, the former being much more brilliantly coloured than the latter. Owing to the fact that the early Dutch settlers at the Cape first met with the antelopes which they called bonteboks on the plains near Cape Agulhas, and subsequently at first gave the same name to the nearly allied species which was discovered about one hundred years later in the neighbourhood of the Orange river, although these latter were undoubtedly blesboks and not bonteboks, a great confusion arose between these two nearly allied species, which I think that I was the first to clear up, in the article on the bontebok which I contributed to the Great and Small Game of Africa, published by Rowland Ward, Limited, in 1899. I cannot go into all the arguments I then used, but there can be no doubt that the animals which Captain (afterwards Sir Cornwallis) Harris first met with on the bontebok flats near the Orange river, in the Colesburg division of the Cape Colony, were blesboks and not bonteboks, and that all the millions of antelopes of the same species which he subsequently saw to the north of the Orange river and thought to be bonteboks were also all blesboks, and that he never saw a bontebok at all until after his return to the Cape, when he made a special journey to Cape Agulhas to secure specimens of I think myself that the correct determination of the true distribution of these two nearly allied species of antelopes is of the utmost importance to the question as to the influence of environment on the coloration of animals. I imagine that the white-faced bontebok was evolved from the same ancestral form as the topi and the tiang of East and Northern Africa, for the new-born bontebok as well as the blesbok has a blackish brown face, and I believe—however fantastic this belief may appear to be—that the wonderfully rich and varied coloration of this remarkable antelope has been brought about purely through the influence of its exceptional environment. The plains where these animals live lie along the shore of a deep blue sea, the ground beneath their feet is at certain seasons of the year carpeted with wild flowers, which grow in such profusion that they give a distinct colour to the landscape, whilst above them rises a range of mountains of a considerable altitude, the upper parts of which are often covered with a mantle of pure white snow. I cannot imagine how any one who has seen bonteboks on the plains they inhabit can believe that their white rumps, faces, bellies, and legs, contrasting as they do so vividly with the dark rich brown of their sides and necks, can afford them any protection against their carnivorous foes; nor, although a white rump or face is a conspicuous mark, can I see the necessity of recognition marks for animals which live on open plains where the vegetation is short, and where an animal's whole body can be seen at a long distance. In the blesbok, which also lives on open plains, Not only has the rich and beautifully variegated body colouring of the bontebok become an almost uniform dark brown in the blesbok, but the snow-white disc on the rump of the former animal has turned to a pale brown in the latter, whilst the area of white on the face and legs of the bontebok has already been considerably contracted in the blesbok. Personally, I look upon the blesbok as a faded bontebok; faded because it moved northwards out Could the opening up of Africa by the destructive civilised races have been delayed for a few hundred or a few thousand years, the blesbok would no doubt have lost the white blaze down the face as completely as it has lost the white disc over the tail, which is so conspicuous a feature in the coloration of its immediate ancestor, the bontebok. To those who believe that every spot or stripe or patch of colour on every animal is a beautiful illustration of the truth of the theory of protective coloration, this may seem a very fanciful idea. Yet I feel convinced that the influence of environment has played a greater part than is generally believed in the evolution of colour in living organisms. The weak point in the theory of protective coloration when applied to large mammals is the fact that all carnivorous animals are nocturnal and seek their prey habitually by night and by scent, and only occasionally by daylight and by sight. I submit that the beautiful case in the Central Hall of the Natural History Museum at South Kensington—showing an Arctic fox, in its white winter coat, approaching a Polar hare, also in winter dress, and an ermine (stoat) hunting for ptarmigan (evidently by sight)—gives an entirely false view of the struggle for life as carried on by animals inhabiting the Arctic Regions, for it conveys the idea of the carnivorous animals of those snow-covered wastes hunting for their prey in a bright light and by eyesight alone. But the truth is that the Arctic winter, during the long continuance of which all living resident creatures, with the exception of the musk ox, become There is one other point regarding the protection afforded by colour to large mammals against carnivorous foes which I think has not been sufficiently considered by naturalists, and that is, that no matter how well the colour of an animal may harmonise with its surroundings as long as it remains perfectly still, as soon as it moves "it jumps to the eyes," as the French say, no matter what its colour may be. What is called protective coloration to be effective must be motionless. Movement, even very slight movement, at once destroys its efficacy. But no herbivorous animals can remain constantly motionless. They lie down and rest certainly during the heat of the day, which is, however, just the time when all carnivorous animals are sleeping. At night and in the early mornings and late evenings they move about feeding, and it is at such times that carnivorous animals hunt for their prey. In the dark these latter are undoubtedly I have never yet heard any explanation given of the black, and therefore most conspicuous, coloration of the Cape buffalo. If any animals needed protective coloration buffaloes certainly did, for in the interior of South Africa they formed the favourite food of the lion, and enormous numbers of them must have been annually killed by these powerful carnivora, which seemed to live with and follow the larger herds in all their wanderings. It certainly seems very strange to me that giraffes, which are very seldom killed by lions or other carnivora, should have found it necessary to evolve a colour which harmonises with their surroundings, as a protection against such foes, whilst buffaloes, which in many districts used once to form the principal food for the lions living in the same countries, have retained throughout the ages a coloration which is everywhere except in deep shade singularly conspicuous. Altogether, a very long experience of the larger mammals inhabiting Africa and some other parts of the world has convinced me that neither the need of protection against carnivorous foes nor the theory of recognition marks can satisfactorily explain all the wonderful diversity of colour to be seen in the coats of wild animals. |