The moral nature of man has reached its present standard partly through the advancement of his reasoning powers and consequently of a just public opinion, but especially from his sympathies having been rendered more tender and widely diffused through the effects of habit, example, instruction and reflection. It is not improbable that after long practice virtuous tendencies may be inherited. With the more civilized races the conviction of the existence of an all-seeing Deity has had a potent influence on the advance of morality. Ultimately man does not accept the praise or blame of his fellows as his sole guide, though few escape this influence, but his habitual convictions, controlled by reason, afford him the safest rule. His conscience then becomes the supreme judge and monitor. Nevertheless, the first foundation or origin of the moral sense lies in the social instincts, including sympathy; and these instincts, no doubt, were primarily gained, as in the case of the lower animals, through natural selection.

The belief in God has often been advanced as not only the greatest but the most complete of all the distinctions between man and the lower animals. It is, however, impossible, as we have seen, to maintain that this belief is innate or instinctive in man. On the other hand, a belief in all-pervading spiritual agencies seems to be universal, and apparently follows from a considerable advance in man's reason and from a still greater advance in his faculties of imagination, curiosity and wonder. I am aware that the assumed instinctive belief in God has been used by many persons as an argument for His existence. But this is a rash judgment, as we should thus be compelled to believe in the existence of many cruel and malignant spirits, only a little more powerful than man; for the belief in them is far more general than in a beneficent Deity. The idea of a universal and beneficent Creator does not seem to arise in the mind of man until he has been elevated by long-continued culture.

He who believes in the advancement of man from some low organized form will naturally ask, How does this bear on the belief in the immortality of the soul? The barbarous races of man, as Sir J. Lubbock has shown, possess no clear belief of this kind; but arguments derived from the primeval beliefs of savages are, as we have just seen, of little or no avail. Few persons feel any anxiety from the impossibility of determining at what precise period in the development of the individual, from the first trace of a minute germinal vesicle, man becomes an immortal being; and there is no greater cause for anxiety because the period in the gradually ascending organic scale cannot possibly be determined.

I am aware that the conclusions arrived at in this work will be denounced by some as highly irreligious; but he who denounces them is bound to show why it is more irreligious to explain the origin of man as a distinct species by descent from some lower form, through the laws of variation and natural selection, than to explain the birth of the individual through the laws of ordinary reproduction. The birth both of the species and of the individual are equally parts of that grand sequence of events, which our minds refuse to accept as the result of blind chance. The understanding revolts at such a conclusion, whether or not we are able to believe that every slight variation of structure, the union of each pair in marriage, the dissemination of each seed, and other such events have all been ordained for some special purpose.

Sexual selection has been treated at great length in this work; for, as I have attempted to show, it has played an important part in the history of the organic world. I am aware that much remains doubtful, but I have endeavoured to give a fair view of the whole case. In the lower divisions of the animal kingdom sexual selection seems to have done nothing; such animals are often affixed for life to the same spot, or have the sexes combined in the same individual, or, what is still more important, their perceptive and intellectual faculties are not sufficiently advanced to allow of the feelings of love and jealousy, or of the exertion of choice. When, however, we come to the Arthropoda and Vertebrata, even to the lowest classes in these two great sub-kingdoms, sexual selection has effected much; and it deserves notice that we here find the intellectual faculties developed, but in two very distinct lines, to the highest standard, namely in the Hymenoptera [ants, bees, etc.], among the Arthropoda [many insects, spiders, etc.], and in the Mammalia, including man, among the Vertebrata.

In the most distinct classes of the animal kingdom—in mammals, birds, fishes, insects and even crustaceans—the differences between the sexes follow almost exactly the same rules. The males are almost always the wooers; and they alone are armed with special weapons for fighting with their rivals. They are generally stronger and larger than the females, and are endowed with the requisite qualities of courage and pugnacity. They are provided, either exclusively or in a much higher degree than the females, with organs for vocal or instrumental music, and with odoriferous glands. They are ornamented with infinitely diversified appendages and with the most brilliant or conspicuous colors, often arranged in elegant patterns, while the females are unadorned. When the sexes differ in more important structures it is the male which is provided with special sense-organs for discovering the female, with locomotive organs for reaching her, and often with prehensile organs for holding her. These various structures for charming or securing the female are often developed in the male during only part of the year; namely, the breeding season. They have in many cases been transferred in a greater or less degree to the females; and in the latter case they often appear in her as mere rudiments. They are lost or never gained by the males after emasculation. Generally they are not developed in the male during early youth, but appear a short time before the age for reproduction. Hence, in most cases the young of both sexes resemble each other; and the female somewhat resembles her young offspring throughout life. In almost every great class a few anomalous cases occur, where there has been an almost complete transposition of the characters proper to the two sexes; the females assuming characters which properly belong to the males. This surprisingly uniformity in the laws regulating the differences between the sexes in so many and such widely separated classes is intelligible if we admit the action throughout all the higher divisions of the animal kingdom of one common cause; namely, sexual selection.

Sexual selection depends on the success of certain individuals over others of the same sex, in relation to the propagation of the species; while natural selection depends on the success of both sexes, at all ages, in relation to the general conditions of life. The sexual struggle is of two kinds; in the one it is between the individuals of the same sex, generally the males, in order to drive away or kill their rivals, the females remaining passive; while in the other, the struggle is likewise between the individuals of the same sex, in order to excite or charm those of the opposite sex, generally the females, which no longer remain passive, but select the more agreeable partners. This latter kind of selection is closely analogous to that which man unintentionally, yet effectually, brings to bear on his domesticated productions, when he preserves during a long period the most pleasing or useful individuals, without any wish to modify the breed.

The laws of inheritance determine whether characters gained through sexual selection by either sex shall be transmitted to the same sex, or to both; as well as the age at which they shall be developed. It appears that variations arising late in life are commonly transmitted to one and the same sex. Variability is the necessary basis for the action of selection and is wholly independent of it. It follows from this that variations of the same general nature have often been taken advantage of and accumulated through sexual selection in relation to the propagation of the species, as well as through natural selection in relation to the general purposes of life. Hence secondary sexual characters, when equally transmitted to both sexes, can be distinguished from ordinary specific characters only by the light of analogy. The modifications acquired through sexual selection are often so strongly pronounced that the two sexes have frequently been ranked as distinct species, or even as distinct genera. Such strongly marked differences must be in some manner highly important; and we know that they have been acquired in some instances at the cost not only of inconvenience, but of exposure to actual danger.

The belief in the power of sexual selection rests chiefly on the following considerations: The characters which we have the best reasons for supposing to have been thus acquired are confined to one sex; and this alone renders it probable that in most cases they are connected with the act of reproduction. These characters in innumerable instances are fully developed only at maturity; and often during only a part of the year, which is always the breeding season. The males (passing over a few exceptional cases) are the more active in courtship; they are the best armed, and are rendered the most attractive in various ways. It is to be especially observed that the males display their attractions with elaborate care in the presence of the females; and that they rarely or never display them excepting during the season of love. It is incredible that all this should be purposeless. Lastly, we have distinct evidence with some quadrupeds and birds that the individuals of one sex are capable of feeling a strong antipathy or preference for certain individuals of the other sex.

Bearing in mind these facts and not forgetting the marked results of man's unconscious selection, it seems to me almost certain that if the individuals of one sex were during a long series of generations to prefer pairing with certain individuals of the other sex, characterized in some peculiar manner, the offspring would slowly but surely become modified in this same manner. I have not attempted to conceal that, excepting when the males are more numerous than the females, or when polygamy prevails, it is doubtful how the more attractive males succeed in leaving a larger number of offspring to inherit their superiority in ornaments or other charms than the less attractive males; but I have shown that this would probably follow from the females—especially the more vigorous ones, which would be the first to breed—preferring not only the more attractive but at the same time the more vigorous and victorious males.

Although we have some positive evidence that birds appreciate bright and beautiful objects, as with the bower-birds of Australia, and although they certainly appreciate the power of song, yet I fully admit that it is astonishing that the females of many birds and some mammals should be endowed with sufficient taste to appreciate ornaments, which we have reason to attribute to sexual selection; and this is even more astonishing in the case of reptiles, fish and insects. But we really know little about the minds of the lower animals. It cannot be supposed, for instance, that male birds of paradise or peacocks should take such pains in erecting, spreading and vibrating their beautiful plumes before the males for no purpose. We should remember the fact given on excellent authority in a former chapter that several peahens, when debarred from an admired male, remained widows during a whole season rather than pair with another bird.

Nevertheless, I know of no fact in natural history more wonderful than that the female Argus pheasant should appreciate the exquisite shading of the ball-and-socket ornaments and the elegant patterns on the wing feathers of the male. He who thinks that the male was created as he now exists must admit that the great plumes, which prevent the wings from being used for flight and which, as well as the primary feathers, are displayed in a manner quite peculiar to this one species during the act of courtship, and at no other time, were given to him as an ornament. If so, he must likewise admit that the female was created and endowed with the capacity of appreciating such ornaments. I differ only in the conviction that the male Argus pheasant acquired his beauty gradually, through the females having preferred during many generations the more highly ornamented males; the esthetic capacity of the females having been advanced through exercise or habit just as our own taste is gradually improved. In the male, through the fortunate chance of a few feathers not having been modified, we can distinctly see how simple spots with a little fulvous [tawny] shading on one side may have been developed by small steps into the wonderful ball-and-socket ornaments; and it is probable that they were actually thus developed.

Every one who admits the principle of evolution, and yet feels great difficulty in admitting that female mammals, birds, reptiles and fish, could have acquired the high taste implied by the beauty of the males, and which generally coincides with our own standard, should reflect that the nerve-cells of the brain in the highest as well as in the lowest members of the Vertebrate series, are derived from those of the common progenitor of the whole group. It thus becomes intelligible that the brain and mental faculties should be capable under similar conditions of nearly the same course of development, and consequently of performing nearly the same functions.

The reader who has taken the trouble to go through the several chapters devoted to sexual selection will be able to judge how far the conclusions at which I have arrived are supported by sufficient evidence. If he accepts these conclusions he may, I think, safely extend them to mankind; but it would be superfluous here to repeat what I have so lately said on the manner in which sexual selection apparently has acted on man, both on the male and female side, causing the two sexes of man to differ in body and mind, and the several races to differ from each other in various characters, as well as from their ancient and lowly organized progenitors.

He who admits the principle of sexual selection will be led to the remarkable conclusion that the cerebral system not only regulates most of the existing functions of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities. Courage, pugnacity, perseverance, strength and size of body, weapons of all kinds, musical organs, both vocal and instrumental, bright colours, stripes and marks, and ornamental appendages, have all been indirectly gained by the one sex or the other, through the influence of love and jealousy, through the appreciation of the beautiful in sound, colour or form, and through the exertion of a choice; and those powers of the mind manifestly depend on the development of the cerebral system.

Man scans with scrupulous care the character and pedigree of his horses, cattle and dogs before he matches them; but when he comes to his own marriage he rarely, or never takes any such care. He is impelled by nearly the same motives as the lower animals when left to their own free choice, though he is in so far superior to them that he highly values mental charms and virtues. On the other hand he is strongly attracted by mere wealth or rank. Yet he might by selection do something not only for the bodily constitution and frame of his offspring, but for their intellectual and moral qualities. Both sexes ought to refrain from marriage if they are in any marked degree inferior in body or mind; but such hopes are Utopian and will never be even partially realized until the laws of inheritance are thoroughly known. All do good service who aid toward this end. When the principles of breeding and inheritance are better understood, we shall not hear ignorant members of our legislature rejecting with scorn a plan for ascertaining whether or not consanguineous marriages are injurious to man.

The advancement of the welfare of mankind is a most intricate problem; all ought to refrain from marriage who cannot avoid abject poverty for their children; for poverty is not only a great evil, but tends to its own increase by leading to recklessness in marriage. On the other hand, as Mr. Galton has remarked, if the prudent avoid marriage, while the reckless marry, the inferior members tend to supplant the better members of society. Man, like every other animal, has no doubt advanced to his present high condition through a struggle for existence consequent on his rapid multiplication; and if he is to advance still higher, he must remain subject to a severe struggle. Otherwise he would sink into indolence, and the more gifted men would not be more successful in the battle of life than the less gifted. Hence our natural rate of increase, though leading to many and obvious evils, must not be greatly diminished by any means. There should be open competition for all men; and the most able should not be prevented by laws or customs from succeeding best and rearing the largest number of offspring. Important as the struggle for existence has been and even still is, yet as far as the highest part of man's nature is concerned there are other agencies more important. For the moral qualities are advanced, either directly or indirectly, much more through the effects of habit, the reasoning powers, instruction, religion, etc., than through natural selection; though to this latter agency the social instincts, which afforded the basis for the development of the moral sense, may be safely attributed.

The main conclusion arrived at in this work, namely, that man is descended from some lowly organized form, will, I regret to think, be highly distasteful to many. But there can hardly be a doubt that we are descended from barbarians. The astonishment I felt on first seeing a party of Fuegians on a wild and broken shore will never be forgotten by me, for the reflection at once rushed into my mind—such were our ancestors. These men were absolutely naked and bedaubed with paint, their long hair was tangled, their mouths frothed with excitement, and their expression was wild, startled and distrustful. They possessed hardly any arts, and like wild animals lived on what they could catch; they had no government, and were merciless to every one not of their own small tribe. He who has seen a savage in his native land will not feel much shame, if forced to acknowledge that the blood of some more humble creature flows in his veins. For my own part I would as soon be descended from that heroic little monkey who braved his dreaded enemy in order to save the life of his keeper; or from that old baboon, who, descending from the mountains, carried away in triumph his young comrade from a crowd of astonished dogs—as from a savage who delights to torture his enemies, offers up bloody sacrifices, practises infanticide without remorse, treats his wives like slaves, knows no decency, and is haunted by the grossest superstitions.

Man may be excused for feeling some pride at having risen, though not through his own exertions, to the very summit of the organic scale; and the fact of his having thus risen, instead of having been aboriginally placed there, may give him hope for a still higher destiny in the distant future. But we are not here concerned with hopes or fears, only with the truth as far as our reason permits us to discover it. I have given the evidence to the best of my ability, and we must acknowledge, as it seems to me, that man, with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his godlike intellect which has penetrated into the movements and constitution of the solar system—with all these exalted powers—Man still bears in his bodily frame the indelible stamp of his lowly origin.



MIMICRY AND OTHER PROTECTIVE RESEMBLANCES AMONG ANIMALS

ALFRED RUSSEL WALLACE

[Mr. Wallace, one of the greatest naturalists of the age, discovered the law of natural selection independently of Darwin, and about the same time. Among his works are "The Malay Archipelago," "Island Life," and "Darwinism." From "Natural Selection," which was published by Macmillan & Co., 1871, the following extracts are taken. The theme has received important development at the hands of Professor E. B. Poulton, in his "The Colours of Animals," International Scientific Series, 1890: and in F. E. Beddard's "Animal Colouration"; London, Swan Sonnenschein; N. Y., Macmillan, 1892.]

There is no more convincing proof of the truth of a comprehensive theory, than its power of absorbing and finding a place for new facts, and its capability of interpreting phenomena which had been previously looked upon as unaccountable anomalies. It is thus that the law of universal gravitation and the undulatory theory of light have become established and universally accepted by men of science. Fact after fact has been brought forward as being apparently inconsistent with them, and one after another these very facts have been shown to be the consequences of the laws they were at first supposed to disprove. A false theory will never stand this test. Advancing knowledge brings to light whole groups of facts which it cannot deal with, and its advocates steadily decrease in numbers, notwithstanding the ability and scientific skill with which it has been supported. The course of a true theory is very different, as may be well seen by the progress of opinion on the subject of natural selection. In less than eight years "The Origin of Species" has produced conviction in the minds of a majority of the most eminent living men of science. New facts, new problems, new difficulties as they arise are accepted, solved or removed by this theory; and its principles are illustrated by the progress and conclusions of every well established branch of human knowledge. It is the object of the present essay to show how it has recently been applied to connect together and explain a variety of curious facts which had long been considered as inexplicable anomalies.

Perhaps no principle has ever been announced so fertile in results as that which Mr. Darwin so earnestly impresses upon us, and which is indeed a necessary deduction from the theory of natural selection, namely—that none of the definite facts of organic nature, no special organ, no characteristic form or marking, no peculiarities of instinct or of habit, no relations between species or between groups of species—can exist, but which must now be or once have been useful to the individuals or races which possess them. This great principle gives us a clue which we can follow out in the study of many recondite phenomena, and leads us to seek a meaning and a purpose of some definite character in minutiae which we should be otherwise almost sure to pass over as insignificant or unimportant.

The adaptation of the external colouring of animals to their conditions of life has long been recognized, and has been imputed either to an originally created specific peculiarity, or to the direct action of climate, soil, or food. Where the former explanation has been accepted, it has completely checked inquiry, since we could never get any further than the fact of the adaptation. There was nothing more to be known about the matter. The second explanation was soon found to be quite inadequate to deal with all the varied phases of the phenomena, and to be contradicted by many well-known facts. For example, wild rabbits are always of gray or brown tints well suited for concealment among grass and fern. But when these rabbits are domesticated, without any change of climate or food, they vary into white or black, and these varieties may be multiplied to any extent, forming white or black races. Exactly the same thing has occurred with pigeons; and in the case of rats and mice, the white variety has not been shown to be at all dependent on alteration of climate, food or other external conditions. In many cases the wings of an insect not only assume the exact tint of the bark or leaf it is accustomed to rest on, but the form and veining of the leaf or the exact rugosity of the bark is imitated; and these detailed modifications cannot be reasonably imputed to climate or food, since in many cases the species does not feed on the substance it resembles, and when it does, no reasonable connection can be shown to exist between the supposed cause and the effect produced. It was reserved for the theory of natural selection to solve all these problems, and many others which were not at first supposed to be directly connected with them. To make these latter intelligible, it will be necessary to give a sketch of the whole series of phenomena which may be classed under the head of useful or protective resemblances.

Concealment, more or less complete, is useful to many animals, and absolutely essential to some. Those which have numerous enemies from which they cannot escape by rapidity of motion, find safety in concealment. Those which prey upon others must also be so constituted as not to alarm them by their presence or their approach, or they would soon die of hunger. Now, it is remarkable in how many cases nature gives this boon to the animal, by colouring it with such tints as may best serve to enable it to escape from its enemies or to entrap its prey. Desert animals as a rule are desert-coloured. The lion is a typical example of this, and must be almost invisible when crouched upon the sand or among desert rocks and stones. Antelopes are all more or less sandy-coloured. The camel is pre-eminently so. The Egyptian cat and the Pampas cat are sandy or earth-coloured. The Australian kangaroos are of the same tints, and the original colour of the wild horse is supposed to have been a sandy or clay-colour.

The desert birds are still more remarkably protected by their assimilative hues. The stone-chats, the larks, the quails, the goatsuckers and the grouse, which abound in the North African and Asiatic deserts, are all tinted and mottled so as to resemble with wonderful accuracy the average colour and aspect of the soil in the district they inhabit. The Rev. H. Tristram, in his account of the ornithology of North Africa in the first volume of the "Ibis," says: "In the desert, where neither trees, brushwood, nor even undulation of the surface afford the slightest protection to its foes, a modification of colour which shall be assimilated to that of the surrounding country is absolutely necessary. Hence without exception the upper plumage of every bird, whether lark, chat, sylvain, or sand-grouse, and also the fur of all the smaller mammals, and the skin of all the snakes and lizards, is of one uniform isabelline or sand colour." After the testimony of so able an observer it is unnecessary to adduce further examples of the protective colours of desert animals.

Almost equally striking are the cases of arctic animals possessing the white colour that best conceals them upon snowfields and icebergs. The polar bear is the only bear that is white, and it lives constantly among snow and ice. The arctic fox, the ermine and the alpine hare change to white in winter only, because in summer white would be more conspicuous than any other colour, and therefore a danger rather than a protection; but the American polar hare, inhabiting regions of almost perpetual snow, is white all the year round. Other animals inhabiting the same northern regions do not, however, change colour. The sable is a good example, for throughout the severity of a Siberian winter it retains its rich brown fur. But its habits are such that it does not need the protection of colour, for it is said to be able to subsist on fruits and berries in winter, and to be so active upon the trees as to catch small birds among the branches. So also the woodchuck of Canada has a dark-brown fur; but then it lives in burrows and frequents river banks, catching fish and small animals that live in or near the water.

Among birds, the ptarmigan is a fine example of protective colouring. Its summer plumage so exactly harmonizes with the lichen-coloured stones among which it delights to sit, that a person may walk through a flock of them without seeing a single bird; while in winter its white plumage is an almost equal protection. The snow-bunting, the jerfalcon, and the snowy owl are also white-coloured birds inhabiting the arctic regions, and there can be little doubt but that their colouring is to some extent protective.

Nocturnal animals supply us with equally good illustrations. Mice, rats, bats, and moles possess the least conspicuous of hues, and must be quite invisible at times when any light colour would be instantly seen. Owls and goatsuckers are of those dark mottled tints that will assimilate with bark and lichen, and thus protect them during the day, and at the same time be inconspicuous in the dusk.

It is only in the tropics, among forests which never lose their foliage, that we find whole groups of birds whose chief colour is green. The parrots are the most striking example, but we have also a group of green pigeons in the East; and the barbets, leaf-thrushes, bee-eaters, white-eyes, turacos, and several smaller groups, have so much green in their plumage as to tend greatly to conceal them among the foliage.

The conformity of tint which has been so far shown to exist between animals and their habitations is of somewhat general character; we will now consider the cases of more special adaptation. If the lion is enabled by his sandy colour readily to conceal himself by merely crouching down in the desert, how, it may be asked, do the elegant markings of the tiger, the jaguar, and the other large cats agree with this theory? We reply that these are generally cases of more or less special adaptation. The tiger is a jungle animal, and hides himself among tufts of grass or of bamboos, and in these positions the vertical stripes with which his body is adorned must so assimilate with the vertical stems of the bamboo, as to assist greatly in concealing him from his approaching prey. How remarkable it is that besides the lion and tiger, almost all the other large cats are arboreal in their habits, and almost all have ocellated or spotted skins, which must certainly tend to blend them with the background of foliage; while the one exception, the puma, has an ashy-brown uniform fur, and has the habit of clinging so closely to a limb of a tree while waiting for his prey to pass beneath as to be hardly distinguishable from the bark.

Among birds, the ptarmigan, already mentioned, must be considered a remarkable case of special adaptation. Another is a South American goatsucker (Caprimulgus rupestris) which rests in the bright sunshine on little bare rocky islets in the upper Rio Negro, where its unusually light colours so closely resemble those of the rock and sand, that it can scarcely be detected until trodden upon.

The Duke of Argyll, in his "Reign of Law," has pointed out the admirable adaptation of the colours of the woodcock to its protection. The various browns and yellows and pale ash-colour that occur on fallen leaves are all reproduced in its plumage, so that when according to its habit it rests upon the ground under trees, it is almost impossible to detect it. In snipes the colours are modified so as to be equally in harmony with the prevalent forms and colours of marshy vegetation. Mr. J. M. Lester, in a paper read before the Rugby School Natural History Society observes:—"The wood-dove, when perched amongst the branches of its favourite fir, is scarcely discernible; whereas, were it among some lighter foliage the blue and purple tints in its plumage would far sooner betray it. The robin redbreast, too, although it might be thought that the red on its breast made it much easier to be seen, is in reality not at all endangered by it, since it generally contrives to get among some russet or yellow fading leaves, where the red matches very well with the autumn tints, and the brown of the rest of the body with the bare branches."

Reptiles offer us many similar examples. The most arboreal lizards, the iguanas, are as green as the leaves they feed upon, and the slender whip-snakes are rendered almost invisible as they glide among the foliage by a similar colouration. How difficult it is sometimes to catch sight of the little green tree-frogs sitting on the leaves of a small plant enclosed in a glass case in the Zoological Gardens; yet how much better concealed they must be among the fresh green damp foliage of a marshy forest. There is a North American frog found on lichen-covered rocks and walls, which is so coloured as exactly to resemble them, and as long as it remains quiet would certainly escape detection. Some of the geckos which cling motionless on the trunks of trees in the tropics, are of such curiously marbled colours as to match exactly with the bark they rest upon.

In every part of the tropics there are tree snakes that twist among boughs and shrubs, or lie coiled up in the dense masses of foliage. These are of many distinct groups, and comprise both venomous and harmless genera; but almost all of them are of a beautiful green colour, sometimes more or less adorned with white or dusky bands and spots. There can be little doubt that this colour is doubly useful to them, since it will tend to conceal them from their enemies, and will lead their prey to approach them unconscious of danger. Dr. Gunthner informs me that there is only one genus of true arboreal snakes (Dipsas) whose colours are rarely green, but are of various shades of black, brown, and olive, and these are all nocturnal reptiles, and there can be little doubt conceal themselves during the day in holes, so that the green protective tint would be useless to them, and they accordingly retain the more usual reptilian hues.

Fishes present similar instances. Many flat fish, as, for example, the flounder and the skate, are exactly the colour of the gravel or sand on which they habitually rest. Among the marine flower gardens of an Eastern coral reef the fishes present every variety of gorgeous colour, while the river fish even of the tropics rarely if ever have gay or conspicuous markings. A very curious case of this kind of adaptation occurs in the sea-horse (Hippocampus) of Australia, some of which bear long foliaceous appendages resembling seaweed, and are of a brilliant red colour; and they are known to live among seaweed of the same hue, so that when at rest they must be quite invisible. There are now in the aquarium of the Zoological Society some slender green pipe-fish which fasten themselves to any object at the bottom by their prehensile tails, and float about with the current, looking exactly like some cylindrical algae.

It is, however, in the insect world that this principle of the adaptation of animals to their environment is most fully and strikingly developed. In order to understand how general this is, it is necessary to enter somewhat into details, as we shall thereby be better able to appreciate the significance of the still more remarkable phenomena we shall presently have to discuss. It seems to be in proportion to their sluggish motions or the absence of other means of defence, that insects possess the protective colouring. In the tropics there are thousands of species of insects which rest during the day clinging to the bark of dead or fallen trees; and the greater portion of these are delicately mottled with gray and brown tints, which though symmetrically disposed and infinitely varied, yet blend so completely with the usual colours of the bark that at two or three feet distance they are quite undistinguishable. In some cases a species is known to frequent only one species of tree. This is the case with the common South American long-horned beetle (Onychocerus scorpio) which, Mr. Bates informed me, is found only on a rough-barked tree, called Tapiriba, on the Amazon. It is very abundant, but so exactly does it resemble the bark in colour and rugosity, and so closely does it cling to the branches, that until it moves it is absolutely invisible! An allied species (O. concentricus) is found only at Para, on a distinct species of tree, the bark of which it resembles with equal accuracy. Both these insects are abundant, and we may fairly conclude that the protection they derive from this strange concealment is at least one of the causes that enable the race to flourish.

Many of the species of Cicindela, or tiger beetle, will illustrate this mode of protection. Our common Cicindela campestris frequents grassy banks and is of a beautiful green colour, while C. maritima, which is found only on sandy sea-shores, is of a pale bronzy yellow, so as to be almost invisible. A great number of the species found by myself in the Malay islands are similarly protected. The beautiful Cicindela gloriosa, of a very deep velvety green colour, was only taken upon wet mossy stones in the bed of a mountain stream, where it was with the greatest difficulty detected. A large brown species (C. heros) was found chiefly on dead leaves in forest paths; and one which was never seen except on the wet mud of salt marshes was of a glossy olive so exactly the colour of the mud as only to be distinguished when the sun shone, by its shadow! Where the sandy beach was coralline and nearly white, I found a very pale Cicindela; wherever it was volcanic and black, a dark species of the same genus was sure to be met with.

There are in the East small beetles of the family Buprestidae which generally rest on the midrib of a leaf, and the naturalist often hesitates before picking them off, so closely do they resemble pieces of bird's dung. Kirby and Spence mention the small beetle Onthophilus sulcatus as being like the seed of an umbelliferous plant; and another small weevil, which is much persecuted by predatory beetles of the genus Harpalus, is of the exact colour of loamy soil, and was found to be particularly abundant in loam pits. Mr. Bates mentions a small beetle (Chlamys pilula) which was undistinguishable by the eye from the dung of caterpillars, while some of the Cassidae, from their hemispherical forms and pearly gold-colour, resemble glittering dew-drops upon the leaves.

A number of our small brown and speckled weevils at the approach of any object roll off the leaf they are sitting on, at the same time drawing in their legs and antennae, which fit so perfectly into cavities for their reception that the insect becomes a mere oval brownish lump, which it is hopeless to look for among the similarly coloured little stones and earth pellets among which it lies motionless.

The distribution of colour in butterflies and moths respectively is very instructive from this point of view. The former have all their brilliant colouring on the upper surface of all four wings, while the under surface is almost always soberly coloured, and often very dark and obscure. The moths on the contrary have generally their chief colour on the hind wings only, the upper wings being of dull, sombre, and often imitative tints, and these generally conceal the hind wings when the insects are in repose. This arrangement of the colours is therefore eminently protective, because the butterfly always rests with his wings raised so as to conceal the dangerous brilliancy of his upper surface. It is probable that if we watched their habits sufficiently we should find the under surface of the wings of butterflies very frequently imitative and protective. Mr. T. W. Wood has pointed out that the little orange-tip butterfly often rests in the evening on the green and white flower heads of an umbelliferous plant, and that when observed in this position the beautiful green and white mottling of the under surface completely assimilates with the flower heads and renders the creature very difficult to be seen. It is probable that the rich dark colouring of the under side of our peacock, tortoiseshell, and red-admiral butterflies answers a similar purpose.

Two curious South American butterflies that always settle on the trunks of trees (Gynecia dirce and Callizona acesta) have the under surface curiously striped and mottled, and when viewed obliquely must closely assimilate with the appearance of the furrowed bark of many kinds of trees. But the most wonderful and undoubted case of protective resemblance in a butterfly which I have ever seen, is that of the common Indian Kallima inachis, and its Malayan ally, Kallima paralekta. The upper surface of these insects is very striking and showy, as they are of a large size, and are adorned with a broad band of rich orange on a deep bluish ground. The under side is very variable in colour, so that out of fifty specimens no two can be found exactly alike, but every one of them will be of some shade of ash or brown or ochre, such as are found among dead, dry or decaying leaves. The apex of the upper wings is produced into an acute point, a very common form in the leaves of tropical shrubs and trees, and the lower wings are also produced into a short, narrow tail. Between these two points runs a dark curved line exactly representing the midrib of a leaf, and from this radiate on each side a few oblique lines, which serve to indicate the lateral veins of a leaf. These marks are more clearly seen on the outer portion of the base of the wings, and on the inner side towards the middle and apex, and it is very curious to observe how the usual marginal and transverse striae of the group are here modified and strengthened so as to become adapted for an imitation of the venation of a leaf. We come now to a still more extraordinary part of the imitation, for we find representations of leaves in every stage of decay, variously blotched and mildewed and pierced with powdery black dots gathered into patches and spots, so closely resembling the various kinds of minute fungi that grow on dead leaves that is it impossible to avoid thinking at first sight that the butterflies themselves have been attacked by real fungi.

But this resemblance, close as it is, would be little use if the habits of the insect did not accord with it. If the butterfly sat upon leaves or upon flowers, or opened its wings so as to expose the upper surface, or exposed and moved its head and antennae as many other butterflies do, its disguise would be of little avail. We might be sure, however, from the analogy of many other cases, that the habits of the insect are such as still further to aid its deceptive garb; but we are not obliged to make any such supposition, since I myself had the good fortune to observe scores of Kallima paralekta, in Sumatra, and to capture many of them, and can vouch for the accuracy of the following details: These butterflies frequent dry forests and fly very swiftly. They were never seen to settle on a flower or a green leaf, but were many times lost sight of in a bush or tree of dead leaves. On such occasions they were generally searched for in vain, for while gazing intently at the very spot where one had disappeared, it would often suddenly dart out and again vanish twenty or fifty yards further on. On one or two occasions the insect was detected reposing, and it could then be seen how completely it assimilates itself to the surrounding leaves. It sits on a nearly upright twig, the wings fitting closely back to back, concealing the antennae and head, which are drawn up between their bases. The little tails of the hind wings touch the branch and form a perfect stalk to the leaf, which is supported in its place by the claws of the middle pair of feet, which are slender and inconspicuous. The irregular outline of the wings gives exactly the perspective effect of a shrivelled leaf. We thus have size, colour, form, markings, and habits, all combining together to produce a disguise which may be said to be absolutely perfect; and the protection which it affords is sufficiently indicated by the abundance of the individuals that possess it....

We will now endeavour to show how these wonderful resemblances have most probably been brought about. Returning to the higher animals, let us consider the remarkable fact of the rarity of white colouring in the mammalia or birds of the temperate or tropical zones in a state of nature. There is not a single white land-bird or quadruped in Europe, except the few arctic or alpine species to which white is a protective colour. Yet in many of these creatures there seems to be no inherent tendency to avoid white, for directly they are domesticated white varieties arise, and appear to thrive as well as others. We have white mice and rats, white cats, horses, dogs, and cattle, white poultry, pigeons, turkeys, and ducks, and white rabbits. Some of these animals have been domesticated for a long period, others only for a few centuries; but in almost every case in which an animal has been thoroughly domesticated, parti-coloured and white varieties are produced and become permanent.

It is also well known that animals in a state of nature produce white varieties occasionally. Blackbirds, starlings, and crows are occasionally seen white, as well as elephants, deer, tigers, hares, moles, and many other animals; but in no case is a permanent white race produced. Now there are no statistics to show that the normal-coloured parents produce white offspring oftener under domestication than in a state of nature, and we have no right to make such an assumption if the facts can be accounted for without it. But if the colours of animals do really, in the various instances already adduced, serve for their concealment and preservation, then white or any other conspicuous colour must be hurtful, and must in most cases shorten an animal's life. A white rabbit would be more surely the prey of hawk or buzzard, and the white mole, or field mouse, could not long escape from the vigilant owl. So, also, any deviation from those tints best adapted to conceal a carnivorous animal would render the pursuit of its prey much more difficult, would place it at a disadvantage among its fellows and in a time of scarcity would probably cause it to starve to death. On the other hand, if an animal spreads from a temperate into an arctic district, the conditions are changed. During a large portion of the year, and just when the struggle for existence is most severe, white is the prevailing tint of nature, and dark colours will be the most conspicuous. The white varieties will now have an advantage; they will escape from their enemies or will secure food, while their brown companions will be devoured or will starve; and "as like produces like" is the established rule in nature, the white race will become permanently established, and dark varieties, when they occasionally appear, will soon die out from their want of adaptation to their environment. In each case the fittest will survive, and a race will be eventually produced adapted to the conditions in which it lives.

We have here an illustration of the simple and effectual means by which animals are brought into harmony with the rest of nature. That slight amount of variability in every species, which we often look upon as something accidental or abnormal, or so insignificant as to be hardly worthy of notice, is yet the foundation of all those wonderful and harmonious resemblances which play such an important part in the economy of nature. Variation is generally very small in amount, but it is all that is required, because the change in the external conditions to which an animal is subject is generally very slow and intermittent. When these changes have taken place too rapidly, the result has often been the extinction of species; but the general rule is, that climatal and geological changes go on slowly, and the slight but continual variations in the colour, form and structure of all animals, has furnished individuals adapted to these changes, and who have become the progenitors of modified races. Rapid multiplication, incessant slight variation, and survival of the fittest—these are the laws which ever keep the organic world in harmony with the inorganic and with itself. These are the laws which we believe have produced all the cases of protective resemblance already adduced, as well as those still more curious examples we have yet to bring before our readers.

It must always be borne in mind that the more wonderful examples, in which there is not only a general but a special resemblance as in the walking leaf, the mossy phasma, and the leaf-winged butterfly—represent those few instances in which the process of modification has been going on during an immense series of generations. They all occur in the tropics, where the conditions of existence are the most favourable, and where climatic changes have for long periods been hardly perceptible. In most of them favourable variations both of colour, form, structure, and instinct or habit, must have occurred to produce the perfect adaptation we now behold. All these are known to vary, and favourable variations when not accompanied by others that are unfavourable, would certainly survive. At one time a little step might be made in this direction, at another time in that—a change of conditions might sometimes render useless that which it had taken ages to produce—great and sudden physical modifications might often produce the extinction of a race just as it was approaching perfection, and a hundred checks of which we can know nothing may have retarded the progress towards perfect adaptation; so that we can hardly wonder at there being so few cases in which a completely successful result has been attained as shown by the abundance and wide diffusion of the creatures so protected.

[Here are given many detailed examples of insects which gainfully mimic one another.]

We will now adduce a few cases in which beetles imitate other insects, and insects of other orders imitate beetles.

Charis melipona, a South American Longicorn of the family Necydalidae, has been so named from its resemblance to a small bee of the genus Melipona. It is one of the most remarkable cases of mimicry, since the beetle has the thorax and body densely hairy like the bee, and the legs are tufted in a manner most unusual in the order Coleoptera. Another Longicorn, Odontocera odyneroides, has the abdomen banded with yellow, and constricted at the base, and is altogether so exactly like a small common wasp of the genus Odynerus, that Mr. Bates informs us he was afraid to take it out of his net with his fingers for fear of being stung. Had Mr. Bates's taste for insects been less omnivorous than it was, the beetle's disguise might have saved it from his pin, as it had no doubt often done from the beak of hungry birds. A larger insect, Sphecomorpha chalybea, is exactly like one of the large metallic blue wasps, and like them has the abdomen connected with the thorax by a pedicle, rendering the deception most complete and striking. Many Eastern species of Longicorns of the genus Oberea, when on the wing exactly resemble Tenthredinidae, and many of the small species of Hesthesis run about on timber, and cannot be distinguished from ants. There is one genus of South American Longicorns that appears to mimic the shielded bugs of the genus Scutellera. The Gymnocerous capucinus is one of these, and is very like Pachyotris fabricii, one of the Scutelleridae. The beautiful Gymnocerous dulcissimus is also very like the same group of insects, though there is no known species that exactly corresponds to it; but this is not to be wondered at, as the tropical Hemiptera have been comparatively so little cared for by collectors.

The most remarkable case of an insect of another order mimicking a beetle is that of the Condylodera tricondyloides, one of the cricket family from the Philippine Islands, which is so exactly like a Tricondyla (one of the tiger beetles), that such an experienced entomologist as Professor Westwood placed it among them in his cabinet, and retained it there a long time before he discovered his mistake! Both insects run along the trunks of trees, and whereas Tricondylas are very plentiful, the insect that mimics it is, as in all other cases, very rare. Mr. Bates also informs us that he found at Santarem on the Amazon, a species of locust which mimicked one of the tiger beetles of the genus Odontocheila, and was found on the same trees which they frequented.

There are a considerable number of Diptera, or two-winged flies, that closely resemble wasps and bees, and no doubt derive much benefit from the wholesome dread which those insects excite. The Midas dives, and other species of large Brazilian flies, have dark wings and metallic blue elongate bodies, resembling the large stinging Sphegidae of the same country; and a very large fly of the genus Asilus has black-banded wings and the abdomen tipped with rich orange, so as exactly to resemble the fine bee Euglossa dimidiata, and both are found in the same parts of South America. We have also in our own country species of Bombylius which are almost exactly like bees. In these cases the end gained by the mimicry is no doubt freedom from attack, but it has sometimes an altogether different purpose. There are a number of parasitic flies whose larvae feed upon the larvae of bees, such as the British genus Volucella and many of the tropical Bombylii, and most of these are exactly like the particular species of bee they prey upon, so that they can enter their nests unsuspected to deposit their eggs. There are also bees that mimic bees. The cuckoo bees of the genus Nomada are parasitic on the Andrenidae, and they resemble either wasps or species of Andrena; and the parasitic humble-bees of the genus Apathus almost exactly resemble the species of humble-bees in whose nests they are reared. Mr. Bates informs us that he found numbers of these "cuckoo" bees and flies on the Amazon, which all wore the livery of working bees peculiar to the same country.

There is a genus of small spiders in the tropics which feed on ants, and they are exactly like ants themselves, which no doubt gives them more opportunity of seizing their prey; and Mr. Bates found on the Amazon a species of Mantis which exactly resembled the white ants which it fed upon, as well as several species of crickets (Saphura), which resembled in a wonderful manner different sand-wasps of large size, which are constantly on the search for crickets with which to provision their nests.

Perhaps the most wonderful case of all is the large caterpillar mentioned by Mr. Bates, which startled him by its close resemblance to a small snake. The first three segments behind the head were dilatable at the will of the insect, and had on each side a large black pupillated spot, which resembled the eye of the reptile. Moreover, it resembled a poisonous viper, not a harmless species of snake, as was proved by the imitation of keeled scales on the crown produced by the recumbent feet, as the caterpillar threw itself backward!

The attitudes of many of the tropical spiders are most extraordinary and deceptive, but little attention has been paid to them. They often mimic other insects, and some, Mr. Bates assures us, are exactly like flower buds, and take their station in the axils of leaves, where they remain motionless waiting for their prey.

I have now completed a brief, and necessarily very imperfect, survey of the various ways in which the external form and colouring of animals is adapted to be useful to them, either by concealing them from their enemies or from the creatures they prey upon. It has, I hope, been shown that the subject is one of much interest, both as regard a true comprehension of the place each animal fills in the economy of nature, and the means by which it is enabled to maintain that place; and also as teaching us how important a part is played by the minutest details in the structure of animals, and how complicated and delicate is the equilibrium of the organic world.

My exposition of the subject having been necessarily somewhat lengthy and full of details, it will be as well to recapitulate its main points.

There is a general harmony in nature between the colours of an animal and those of its habitation. Arctic animals are white, desert animals are sand-coloured; dwellers among leaves and grass are green; nocturnal animals are dusky. These colours are not universal, but are very general, and are seldom reversed. Going on a little further, we find birds, reptiles and insects, so tinted and mottled as exactly to match the rock, or bark, or leaf, or flower they are accustomed to rest upon—and thereby effectually concealed. Another step in advance, and we have insects which are formed as well as coloured so as exactly to resemble particular leaves, or sticks, or mossy twigs, or flowers; and in these cases very peculiar habits and instincts come into play to aid in the deception and render the concealment more complete. We now enter upon a new phase of the phenomena, and come to creatures whose colours neither conceal them nor make them like vegetable or mineral substances; on the contrary, they are conspicuous enough, but they completely resemble some other creature of a quite different group, while they differ much in outward appearance from those with which all essential parts of their organization show them to be really closely allied. They appear like actors or masqueraders dressed up and painted for amusement, or like swindlers endeavouring to pass themselves off for well-known and respectable members of society. What is the meaning of this strange travesty? Does nature descend to imposture or masquerade? We answer, she does not. Her principles are too severe. There is a use in every detail of her handiwork. The resemblance of one animal to another is of exactly the same essential nature as the resemblance to a leaf, or to bark, or to desert sand, and answers exactly the same purpose. In the one case the enemy will not attack the leaf or the bark, and so the disguise is a safeguard; in the other case it is found that for various reasons the creature resembled is passed over, and not attacked by the usual enemies of its order, and thus the creature that resembles it has an equally effectual safeguard. We are plainly shown that the disguise is of the same nature in the two cases, by the occurrence in the same group of one species resembling a vegetable substance, while another resembles a living animal of another group; and we know that the creatures resembled possess an immunity from attack, by their being always very abundant, by their being conspicuous and not concealing themselves, and by their having generally no visible means of escape from their enemies; while, at the same time, the particular quality that makes them disliked is often very clear, such as a nasty taste or an indigestible hardness. Further examination reveals the fact that, in several cases of both kinds of disguise, it is the female only that is thus disguised; and as it can be shown that the female needs protection much more than the male, and that her preservation for a much longer period is absolutely necessary for the continuance of the race, we have an additional indication that the resemblance is in all cases subservient to a great purpose—the preservation of the species.

In endeavouring to explain these phenomena as having been brought about by variation and natural selection, we start with the fact that white varieties frequently occur, and when protected from enemies show no incapacity for continued existence and increase. We know, further, that varieties of many other tints occasionally occur; and as "the survival of the fittest" must inevitably weed out those whose colours are prejudicial and preserve those whose colours are a safeguard, we require no other mode of accounting for the protective tints of arctic and desert animals. But this being granted, there is such a perfectly continuous and graduated series of examples of every kind of protective imitation, up to the most wonderful cases of what is termed "mimicry," that we can find no place at which to draw the line and say,—so far variation and natural selection will account for the phenomena, but for all the rest we require a more potent cause. The counter theories that have been proposed, that of the "special creation" of each imitative form, that of the action of similar "conditions of existence" for some of the cases, and of the laws of "hereditary descent and the reversion to ancestral forms" for others,—have all been shown to be beset with difficulties, and the two latter to be directly contradicted by some of the most constant and most remarkable of the facts to be accounted for.

The important part that protective "resemblance" has played in determining the colours and markings of many groups of animals will enable us to understand the meaning of one of the most striking facts in nature, the uniformity in the colours of the vegetable as compared with the wonderful diversity of the animal world. There appears no good reason why trees and shrubs should not have been adorned with as many varied hues and as strikingly designed patterns as birds and butterflies, since the gay colours of flowers show that there is no incapacity in vegetable tissues to exhibit them. But even flowers themselves present us with none of those wonderful designs, those complicated arrangements of stripes and dots and patches of colour, that harmonious blending of hues in lines and bands and shaded spots, which are so general a feature in insects. It is the opinion of Mr. Darwin that we owe much of the beauty of flowers to the necessity of attracting insects to aid in their fertilization, and that much of the development of colour in the animal world is due to "sexual selection," colour being universally attractive, and thus leading to its propagation and increase; but while fully admitting this, it will be evident from the facts and arguments here brought forward, that very much of the variety both of colour and markings among animals is due to the supreme importance of concealment, and thus the various tints of minerals and vegetables have been directly reproduced in the animal kingdom, and again and again modified as more special protection became necessary. We shall thus have two causes for the development of colour in the animal world and shall be better enabled to understand how, by their combined and separate action, the immense variety we now behold has been produced. Both causes, however, will come under the general law of "Utility," the advocacy of which, in its broadest sense, we owe almost entirely to Mr. Darwin. A more accurate knowledge of the varied phenomena connected with this subject may not improbably give us some information both as to the senses and the mental faculties of the lower animals. For it is evident that if colours which please us also attract them, and if the various disguises which have been here enumerated are equally deceptive to them as to ourselves, then both their powers of vision and their faculties of perception and emotion, must be essentially of the same nature as our own—a fact of high philosophical importance in the study of our own nature and our true relations to the lower animals.[4]

FOOTNOTES:

[4] The author continues this study in Chapter ix of "Darwinism": New York, Macmillan Co., 1889.—Ed.



THE EVOLUTION OF THE HORSE

THOMAS HENRY HUXLEY

[Professor Huxley as a naturalist, educator, and controversialist was one of the commanding figures of the nineteenth century. To physiology and morphology his researches added much of importance: as an expositor he stood unapproached. As the bold and witty champion of Darwinism he gave natural selection an acceptance much more early and wide than it would otherwise have enjoyed. In 1876 he delivered in America three lectures on Evolution: the third of the series is here given. All three are copyrighted and published by D. Appleton & Co., New York, in a volume which also contains a lecture on the study of biology. Since 1876 the arguments of Professor Huxley have been reinforced by the discovery of many fossils connecting not only the horse, but other quadrupeds, with species widely different and now extinct. The most comprehensive collection illustrating the descent of the horse is to be seen at the American Museum of Natural History, New York, where also the evolution of tapirs, camels, llamas, rhinoceroses, dinosaurs, great ground sloths and other animals are clearly to be traced—in most cases by remains discovered in America. A capital book on the theme broached by Professor Huxley is "Animals of the Past," by Frederic A. Lucas, Curator of the Division of Comparative Anatomy, United States National Museum, Washington, D. C., published by McClure, Phillips & Co., New York.

"The Life and Letters of Professor Huxley," edited by his son, Leonard Huxley, is a work of rare interest: it is published by D. Appleton & Co., New York.]

The occurrence of historical facts is said to be demonstrated, when the evidence that they happened is of such a character as to render the assumption that they did not happen in the highest degree improbable; and the question I now have to deal with is, whether evidence in favour of the evolution of animals of this degree of cogency is, or is not, obtainable from the record of the succession of living forms which is presented to us by fossil remains.

Those who have attended to the progress of palaeontology are aware that evidence of the character which I have defined has been produced in considerable and continually-increasing quantity during the last few years. Indeed, the amount and the satisfactory nature of that evidence are somewhat surprising, when we consider the conditions under which alone we can hope to obtain it.

It is obviously useless to seek for such evidence, except in localities in which the physical conditions have been such as to permit of the deposit of an unbroken, or but rarely interrupted, series of strata through a long period of time; in which the group of animals to be investigated has existed in such abundance as to furnish the requisite supply of remains; and in which, finally, the materials composing the strata are such as to insure the preservation of these remains in a tolerably perfect and undisturbed state.

It so happens that the case which, at present, most nearly fulfils all these conditions is that of the series of extinct animals which culminates in the horses; by which term I mean to denote not merely the domestic animals with which we are all so well acquainted, but their allies, the ass, zebra, quagga, and the like. In short, I use "horses" as the equivalent of the technical name Equidae, which is applied to the whole group of existing equine animals.

The horse is in many ways a remarkable animal; not least so in the fact that it presents us with an example of one of the most perfect pieces of machinery in the living world. In truth, among the works of human ingenuity it cannot be said that there is any locomotive so perfectly adapted to its purposes, doing so much work with so small a quantity of fuel, as this machine of nature's manufacture—the horse. And, as a necessary consequence of any sort of perfection, of mechanical perfection as of others, you find that the horse is a beautiful creature, one of the most beautiful of all land animals. Look at the perfect balance of its form, and the rhythm and force of its action. The locomotive machinery is, as you are aware, resident in its slender fore and hind limbs; they are flexible and elastic levers, capable of being moved by very powerful muscles; and, in order to supply the engines which work these levers with the force which they expend, the horse is provided with a very perfect apparatus for grinding its food and extracting therefrom the requisite fuel.

Without attempting to take you very far into the region of osteological detail, I must nevertheless trouble you with some statements respecting the anatomical structure of the horse; and, more especially, will it be needful to obtain a general conception of the structure of its fore and hind limbs, and of its teeth. But I shall only touch upon these points which are absolutely essential to our inquiry.

Let us turn in the first place to the fore-limb. In most quadrupeds, as in ourselves, the fore-arms contains distinct bones called the radius and the ulna. The corresponding region in the horse seem at first to possess but one bone. Careful observation, however, enables us to distinguish in this bone a part which clearly answers to the upper end of the ulna. This is closely united with the chief mass of the bone which represents the radius, and runs out into a slender shaft which may be traced for some distance downwards upon the back of the radius, and then in most cases thins out and vanishes. It takes still more trouble to make sure of what is nevertheless the fact, that a small part of the lower end of the bone of the horse's fore-arm, which is only distinct in a very young foal, is really the lower extremity of the ulna.

What is commonly called the knee of a horse is its wrist. The "cannon bone" answers to the middle bone of the five metacarpal bones, which support the palm of the hand in ourselves. The "pastern," "coronary," and "coffin" bones of veterinarians answer to the joints of our middle fingers, while the hoof is simply a greatly enlarged and thickened nail. But if what lies below the horse's "knee" thus corresponds to the middle finger in ourselves, what has become of the four other fingers or digits? We find in the places of the second and fourth digits only two slender splint-like bones, about two-thirds as long as the cannon bone, which gradually taper to their lower ends and bear no finger joints, or, as they are termed, phalanges. Sometimes, small bony or gristly nodules are to be found at the bases of these two metacarpal splints, and it is probable that these represent rudiments of the first and fifth toes. Thus, the part of the horse's skeleton, which corresponds with that of the human hand, contains one overgrown middle digit, and at least two imperfect lateral digits; and these answer, respectively, to the third, the second and the fourth fingers in man.

Corresponding modifications are found in the hind limb. In ourselves, and in most quadrupeds, the leg contains two distinct bones, a large bone, the tibia, and a smaller and more slender bone, the fibula. But, in the horse, the fibula seems, at first, to be reduced to its upper end; a short slender bone united with the tibia and ending in a point below, occupying its place. Examination of the lower end of a young foal's shin-bone, however, shows a distinct portion of osseous matter, which is the lower end of the fibula; so that the, apparently single, lower end of the shin-bone is really made up of the coalesced ends of the tibia and fibula, just as the, apparently single, lower end of the fore-arm bone is composed of the coalesced radius and ulna.

The heel of the horse is the part commonly known as the hock. The hinder cannon bone answers to the middle metatarsal bone of the human foot, the pastern, coronary, and coffin bones, to the middle toe bones; the hind hoof to the nail; as in the fore-foot. And, as in the fore-foot, there are merely two splints to represent the second and the fourth toes. Sometimes a rudiment of a fifth toe appears to be traceable.

The teeth of a horse are not less peculiar than its limbs. The living engine, like all others, must be well stoked if it is to do its work; and the horse, if it is to make good its wear and tear, and to exert the enormous amount of force required for its propulsion, must be well and rapidly fed. To this end good cutting instruments and powerful and lasting crushers are needful. Accordingly, the twelve cutting teeth of a horse are close-set and concentrated in the fore-part of its mouth, like so many adzes or chisels. The grinders or molars are large, and have an extremely complicated structure, being composed of a number of different substances of unequal hardness. The consequence of this is that they wear away at different rates; and, hence, the surface of each grinder is always as uneven as that of a good millstone.

I have said that the structure of the grinding teeth is very complicated, the harder and the softer parts being, as it were, interlaced with one another. The result of this is that, as the tooth wears, the crown presents a peculiar pattern, the nature of which is not very easily deciphered at first, but which it is important we should understand clearly. Each grinding tooth of the upper jaw has an outer wall so shaped that, on the worn crown, it exhibits the form of two crescents, one in front and one behind, with their concave sides turned outwards. From the inner side of the front crescent, a crescentic front ridge passes inwards and backwards, and its inner face enlarges into a strong longitudinal fold or pillar. From the front part of the hinder crescent, a back ridge takes a like direction, and also has its pillar.

The deep interspaces or valleys between these ridges and the outer wall are filled by bony substance, which is called cement, and coats the whole tooth.

The pattern of the worn face of each grinding tooth of the lower jaw is quite different. It appears to be formed of two crescent-shaped ridges, the convexities of which are turned outwards. The free extremity of each crescent has a pillar, and there is a large double pillar where the two crescents meet. The whole structure is, as it were, imbedded in cement, which fills up the valleys, as in the upper grinders.

If the grinding faces of an upper and of a lower molar of the same side are applied together, it will be seen that the opposed ridges are nowhere parallel, but that they frequently cross; and that thus, in the act of mastication, a hard surface in the one is constantly applied to a soft surface in the other, and vice versa. They thus constitute a grinding apparatus of great efficiency, and one which is repaired as fast as it wears, owing to the long-continued growth of the teeth.

Some other peculiarities of the dentition of the horse must be noticed, as they bear upon what I shall have to say by and by. Thus the crowns of the cutting teeth have a peculiar deep pit, which gives rise to the well-known "mark" of the horse. There is a large space between the outer incisors and the front grinders. In this space the adult male horse presents, near the incisors on each side, above and below, a canine or "tush," which is commonly absent in mares. In a young horse, moreover, there is not unfrequently to be seen, in front of the first grinder, a very small tooth, which soon falls out. If this small tooth be counted as one, it will be found that there are seven teeth behind the canine on each side; namely, the small tooth in question, and the six great grinders, among which, by an unusual peculiarity, the foremost tooth is rather larger than those which follow it.

I have now enumerated those characteristic structures of the horse which are of most importance for the purpose we have in view.

To any one who is acquainted with the morphology [comparative forms] of vertebrated animals, they show that the horse deviates widely from the general structure of mammals; and that the horse type is, in many respects, an extreme modification of the general mammalian plan. The least modified mammals, in fact, have the radius and ulna, the tibia and fibula, distinct and separate. They have five distinct and complete digits on each foot, and no one of these digits is very much larger than the rest. Moreover, in the least modified mammals the total number of the teeth is very generally forty-four, while in horses the usual number is forty, and in the absence of the canines it may be reduced to thirty-six; the incisor teeth are devoid of the fold seen in those of the horse: the grinders regularly diminish in size from the middle of the series to its front end; while their crowns are short, early attain their full length, and exhibit simple ridges or tubercles, in place of the complex foldings of the horse's grinders.

Hence the general principles of the hypothesis of evolution lead to the conclusion that the horse must have been derived from some quadruped which possessed five complete digits on each foot; which had the bones of the fore-arm and of the leg complete and separate; and which possessed forty-four teeth, among which the crowns of the incisors and grinders had a simple structure; while the latter gradually increased in size from before backwards, at any rate in the anterior part of the series, and had short crowns.

And if the horse has been thus evolved, and the remains of the different stages of its evolution have been preserved, they ought to present us with a series of forms in which the number of the digits becomes reduced; the bones of the fore-arm and leg gradually take on the equine condition; and the form and arrangement of the teeth successively approximate to those which obtain in existing horses.

Let us turn to the facts, and see how far they fulfil these requirements of the doctrine of evolution.

In Europe abundant remains of horses are found in the Quaternary and later Tertiary strata as far as the Pliocene formation. But these horses, which are so common in the cave-deposits and in the gravels of Europe, are in all essential respects like existing horses. And that is true of all the horses of the latter part of the Pliocene epoch. But in deposits which belong to the earlier Pliocene and later Miocene epochs, and which occur in Britain, in France, in Germany, in Greece, in India, we find animals which are extremely like horses—which, in fact, are so similar to horses that you may follow descriptions given in works upon the anatomy of the horse upon the skeletons of these animals—but which differ in some important particulars. For example, the structure of their fore and hind limbs is somewhat different. The bones which, in the horse, are represented by two splints, imperfect below, are as long as the middle metacarpal and metatarsal bones; and attached to the extremity of each is a digit with three joints of the same general character as those of the middle digit, only very much smaller. These small digits are so disposed that they could have had but very little functional importance, and they must have been rather of the nature of the dew-claws, such as are to be found in many ruminant animals. The Hipparion, as the extinct European three-toed horse is called, in fact, presents a foot similar to that of the American Protohippus (Fig. 9), except that in the Hipparion the smaller digits are situated farther back and are of smaller proportional size than in the Protohippus.

The ulna is slightly more distinct than in the horse; and the whole length of it, as a very slender shaft intimately united with the radius, is completely traceable. The fibula appears to be in the same condition as in the horse. The teeth of the Hipparion are essentially similar to those of the horse, but the pattern of the grinders is in some respects a little more complex, and there is a depression on the face of the skull in front of the orbit, which is not seen in existing horses.

In the earlier Miocene, and perhaps the later Eocene deposits of some parts of Europe, another extinct animal has been discovered, which Cuvier, who first described some fragments of it, considered to be a Palaeotherium. But as further discoveries threw new light on its structure, it was recognized as a distinct genus under the name of Anchitherium.

In its general characters, the skeleton of Anchitherium is very similar to that of the horse. In fact, Lartet and De Blainville called it Palaeotherium equinum or hippoides; and De Christol, in 1847, said that it differed from Hipparion in little more than the characters of its teeth, and gave it the name of Hipparitherium. Each foot possesses three complete toes; while the lateral toes are much larger in proportion to the middle toe than in Hipparion, and doubtless rested on the ground in ordinary locomotion.

The ulna is complete and quite distinct from that radius, though firmly united with the latter. The fibula seems also to have been complete. Its lower end, though intimately united with that of the tibia, is clearly marked off from the latter bone.

There are forty-four teeth. The incisors have no strong pit. The canines seem to have been well developed in both sexes. The first of the seven grinders, which, as I have said, is frequently absent, and when it does exist, is small in the horse, is a good-sized and permanent tooth, while the grinder which follows it is but little larger than the hinder ones. The crowns of the grinders are short, and though the fundamental pattern of the horse-tooth is discernible, the front and back ridges are less curved, the accessory pillars, are wanting, and the valleys, much shallower, are not filled up with cement.

Seven years ago, when I happened to be looking critically into the bearing of palaeontological facts upon the doctrine of evolution, it appeared to me that the Anchitherium, the Hipparion, and the modern horses, constitute a series in which the modifications of structure coincide with the order of chronological occurrence, in the manner in which they must coincide, if the modern horses really are the result of the gradual metamorphosis, in the course of the Tertiary epoch, of a less specialized ancestral form. And I found by correspondence with the late eminent French anatomist and palaeontologist, M. Lartet, that he had arrived at the same conclusion from the same data.

That the Anchitherium type had become metamorphosed into the Hipparion type, and the latter into the Equine type,[5] in the course of that period of time which is represented by the latter half of the Tertiary deposits, seemed to me to be the only explanation of the facts for which there was even a shadow of probability.

And, hence, I have ever since held that these facts afford evidence of the occurrence of evolution, which, in the sense already defined, may be termed demonstrative.

All who have occupied themselves with the structure of Anchitherium, from Cuvier onwards, have acknowledged its many points of likeness to a well-known genus of extinct Eocene mammals, Palaeotherium. Indeed, as we have seen, Cuvier regarded his remains of Anchitherium as those of a species of Palaeotherium. Hence, in attempting to trace the pedigree of the horse beyond the Miocene epoch and the Anchitheroid form, I naturally sought among the various species of Palaeotheroid animals for its nearest ally, and I was led to the conclusion that the Palaeotherium minus (Plagiolophus) represented the next step more nearly than any form then known.

I think that this opinion was fully justifiable; but the progress of investigation has thrown an unexpected light on the question, and has brought us much nearer than could have been anticipated to a knowledge of the true series of the progenitors of the horse.

You are all aware that, when your country was first discovered by Europeans, there were no traces of the existence of the horse on any part of the American Continent. The accounts of the conquest of Mexico dwell upon the astonishment of the natives of that country when they first became acquainted with that astounding phenomenon—a man seated upon a horse. Nevertheless, the investigations of American geologists have proved that the remains of horses occur in the most superficial deposits of both North and South America, just as they do in Europe. Therefore, for some reason or other—no feasible suggestion on that subject, so far as I know, has been made—the horse must have died out on this continent at some period preceding the discovery of America. Of late years there has been discovered in your Western Territories that marvellous accumulation of deposits, admirably adapted for the preservation of organic remains, to which I referred the other evening, and which furnishes us with a consecutive series of records of the fauna of the older half of the Tertiary epoch, for which we have no parallel in Europe. They have yielded fossils in an excellent state of conservation and in unexampled numbers and variety. The researches of Leidy and others have shown that forms allied to the Hipparion and the Anchitherium are to be found among these remains. But it is only recently that the admirably conceived and most thoroughly and patiently worked-out investigations of Professor Marsh have given us a just idea of the vast fossil wealth, and of the scientific importance, of these deposits. I have had the advantage of glancing over the collections in Yale Museum; and I can truly say, that so far as my knowledge extends, there is no collection from any one region and series of strata comparable, for extent, or for the care with which the remains have been got together, or for their scientific importance, to the series of fossils which he has deposited there. This vast collection has yielded evidence bearing upon the question of the pedigree of the horse of the most striking character. It tends to show that we must look to America, rather than to Europe, for the original seat of the equine series; and that the archaic forms and successive modifications of the horse's ancestry are far better preserved here than in Europe.

Professor Marsh's kindness has enabled me to put before you a diagram, every figure of which is an actual representation of some specimen which is to be seen at Yale at this present time (Fig. 9).

The succession of forms which he has brought together carries us from the top to the bottom of the Tertiaries. Firstly, there is the true horse. Next we have the American Pliocene form of the horse (Pliohippus); in the conformation of its limbs it presents some very slight deviations from the ordinary horse, and the crowns of the grinding teeth are shorter. Then comes the Protohippus, which represents the European Hipparion, having one large digit and two small ones on each foot, and the general characters of the fore-arm and leg to which I have referred. But it is more valuable than the European Hipparion for the reason that it is devoid of some of the peculiarities of that form—peculiarities which tend to show that the European Hipparion is rather a member of a collateral branch, than a form in the direct line of succession. Next, in the backward order in time, is the Miohippus, which corresponds pretty nearly with the Anchitherium of Europe. It presents three complete toes—one large median and two smaller lateral ones; and there is a rudiment of that digit, which answers to the little finger of the human hand.

The European record of the pedigree of the horse stops here; in the American Tertiaries, on the contrary, the series of ancestral equine forms is continued into the Eocene formations. An older Miocene form, termed Mesohippus, has three toes in front, with a large splint-like rudiment representing the little finger; and three toes behind. The radius and ulna, the tibia and the fibula, are distinct, and the short crowned molar teeth are anchitheroid in pattern.

But the most important discovery of all is the Orohippus, which comes from the Eocene formation, and which is the oldest member of the equine series, as yet known. Here we find four complete toes on the front-limb, three toes on the hind-limb, a well-developed ulna, a well-developed fibula, and short-crowned grinders of simple pattern.

Thus, thanks to these important researches, it has become evident that, so far as our present knowledge extends, the history of the horse-type is exactly and precisely that which could have been predicted from a knowledge of the principles of evolution. And the knowledge we now possess justifies us completely in the anticipation, that when the still lower Eocene deposits, and those which belong to the Cretaceous epoch, have yielded up their remains of ancestral equine animals, we shall find, first, a form with four complete toes and a rudiment of the innermost or first digit in front, with probably, a rudiment of the fifth digit in the hind foot;[6] while, in still older forms, the series of the digits will be more and more complete, until we come to the five-toed animals, in which, if the doctrine of evolution is well founded, the whole series must have taken its origin.

That is what I mean by demonstrative evidence of evolution. An inductive hypothesis is said to be demonstrated when the facts are shown to be in entire accordance with it. If that is not scientific proof, there are no merely inductive conclusions which can be said to be proved. And the doctrine of evolution, at the present time, rests upon exactly as secure a foundation as the Copernican theory of the motions of the heavenly bodies did at the time of its promulgation. Its logical basis is precisely of the same character—the coincidence of the observed facts with theoretical requirements.

The only way of escape, if it be a way of escape, from the conclusions which I have just indicated, is the supposition that all these different equine forms have been created separately at separate epochs of time; and, I repeat, that of such an hypothesis as this there neither is, nor can be, any scientific evidence; and, assuredly, so far as I know, there is none which is supported, or pretends to be supported, by evidence or authority of any other kind. I can but think that the time will come when such suggestions as these, such obvious attempts to escape the force of demonstration, will be put upon the same footing as the supposition made by some writers, who are, I believe, not completely extinct at present, that fossils are mere simulacra [images], are no indications of the former existence of the animals to which they seem to belong; but that they are either sports of Nature, or special creations, intended—as I heard suggested the other day—to test our faith.

In fact, the whole evidence is in favour of evolution, and there is none against it. And I say this, although perfectly well aware of the seeming difficulties which have been built up upon what appears to the uninformed to be a solid foundation. I meet constantly with the argument that the doctrine of evolution cannot be well founded because it requires the lapse of a very vast period of time; while the duration of life upon the earth, thus implied, is inconsistent with the conclusions arrived at by the astronomer and the physicist. I may venture to say that I am familiar with those conclusions, inasmuch as some years ago, when president of the Geological Society of London, I took the liberty of criticising them, and of showing in what respects, as it appeared to me, they lacked complete and thorough demonstration. But, putting that point aside, suppose that, as the astronomers, or some of them, and some physical philosophers tell us, it is impossible that life could have endured upon the earth for so long a period as is required by the doctrine of evolution—supposing that to be proved—I desire to be informed, what is the foundation for the statement that evolution does require so great a time? The biologist knows nothing whatever of the amount of time which may be required for the process of evolution. It is a matter of fact that the equine forms, which I have described to you, occur, in the order stated, in the Tertiary formations. But I have not the slightest means of guessing whether it took a million of years, or ten millions, or a hundred millions, or a thousand millions of years, to give rise to that series of changes. A biologist has no means of arriving at any conclusions as to the amount of time which may be needed for a certain quantity of organic change. He takes his time from the geologist. The geologist, considering the rate at which deposits are formed and the rate at which denudation goes on upon the surface of the earth, arrives at more or less justifiable conclusions as to the time which is required for the deposit of a certain thickness of rocks; and if he tells me that the Tertiary formations required 500,000,000 years for their deposit, I suppose he has good ground for what he says, and I take that as a measure of the duration of the evolution of the horse from the Orohippus up to its present condition. And, if he is right, undoubtedly evolution is a very slow process, and requires a great deal of time. But suppose now, that an astronomer or a physicist—for instance, my friend Sir William Thomson—tells me that my geological authority is quite wrong; and that he has weighty evidence to show that life could not possibly have existed upon the surface of the earth 500,000,000 years ago, because the earth would have then been too hot to allow of life, my reply is: "That is not my affair; settle that with the geologist, and when you have come to an agreement among yourselves I will adopt your conclusions." We take our time from the geologists and physicists, and it is monstrous that, having taken our time from the physical philosopher's clock, the physical philosopher should turn round upon us, and say we are too fast or too slow. What we desire to know is, is it a fact that evolution took place? As to the amount of time which evolution may have occupied, we are in the hands of the physicist and the astronomer, whose business it is to deal with those questions.



Fore Foot. Hind Foot. Fore-arm. Leg. Upper Molar. Lower Molar.

RECENT. EQUUS.

PLIOCENE. PLIOHIPPUS.

PROTOHIPPUS (Hipparion).

MIOCENE. MIOHIPPUS (Anchitherium).

MESOHIPPUS.

EOCENE. OROHIPPUS.

FOOTNOTES:

[5] I use the word "type" because it is highly probable that many of the forms of Anchitherium-like and Hipparion-like animals existed in the Miocene and Pliocene epochs, just as many species of the horse tribe exist now; and it is highly improbable that the particular species of Anchitherium or Hipparion, which happen to have been discovered, should be precisely those which have formed part of the direct line of the horse's pedigree.

[6] Since this lecture was delivered, Professor Marsh has discovered a new genus of equine mammals (Eohippus) from the lowest Eocene deposits of the West, which corresponds very nearly to this description.—American Journal of Science, November, 1876.