Contributions to the Theory of Natural Selection
Many birds can only exist by migrating, when their food becomes scarce, to regions possessing a milder, or at least a different climate, though, as these migrating birds are seldom excessively abundant, it is evident that the countries they visit are still deficient in a constant and abundant supply of wholesome food. Those whose organization does not permit them to migrate when their food becomes periodically scarce, can never attain a large population. This is probably the reasons why woodpeckers are scarce with us, while in the tropics they are among the most abundant of solitary birds. Thus the house sparrow is more abundant than the redbreast, because its food is more constant and plentiful,—seeds of grasses being preserved during the winter, and our farm-yards and stubble-fields furnishing an almost inexhaustible supply. Why, as a general rule, are aquatic, and especially sea birds, very numerous in individuals? Not because they are more prolific than others, generally the contrary; but because their food never fails, the sea-shores and river-banks daily swarming with a fresh supply of small mollusca and crustacea. Exactly the same laws will apply to mammals. Wild cats are prolific and have few enemies; why then are they never as abundant as rabbits? The only intelligible answer is, that their supply of food is more precarious. It appears evident, therefore, that so long as a country remains physically unchanged, the numbers of its animal population cannot materially increase. If one species does so, some others requiring the same kind of food must diminish in proportion. The numbers that die annually must be immense; and as the individual existence of each animal depends upon itself, those that die must be the weakest—the very young, the aged, and the diseased—while those that prolong their existence can only be the most perfect in health and vigour—those who are best able to obtain food regularly, and avoid their numerous enemies. It is, as we commenced by remarking, “a struggle for existence,” in which the weakest and least perfectly organized must always succumb.
The Abundance or Rarity of a Species dependent upon its more or less perfect Adaptation to the Conditions of Existence
It seems evident that what takes place among the individuals of a species must also occur among the several allied species of a group,—viz., that those which are best adapted to obtain a regular supply of food, and to defend themselves against the attacks of their enemies and the vicissitudes of the seasons, must necessarily obtain and preserve a superiority in population; while those species which from some defect of power or organization are the least capable of counteracting the vicissitudes of food-supply, &c., must diminish in numbers, and, in extreme cases, become altogether extinct. Between these extremes the species will present various degrees of capacity for ensuring the means of preserving life; and it is thus we account for the abundance or rarity of species. Our ignorance will generally prevent us from accurately tracing the effects to their causes; but could we become perfectly acquainted with the organization and habits of the various species of animals, and could we measure the capacity of each for performing the different acts necessary to its safety and existence under all the varying circumstances by which it is surrounded, we might be able even to calculate the proportionate abundance of individuals which is the necessary result.
If now we have succeeded in establishing these two points—1st, that the animal population of a country is generally stationary, being kept down by a periodical deficiency of food, and other checks; and, 2nd, that the comparative abundance or scarcity of the individuals of the several species is entirely due to their organization and resulting habits, which, rendering it more difficult to procure a regular supply of food and to provide for their personal safety in some cases than in others, can only be balanced by a difference in the population which have to exist in a given area—we shall be in a condition to proceed to the consideration of varieties, to which the preceding remarks have a direct and very important application.
Useful Variations will tend to Increase; useless or hurtful Variations to Diminish
Most or perhaps all the variations from the typical form of a species must have some definite effect, however slight, on the habits or capacities of the individuals. Even a change of colour might, by rendering them more or less distinguishable, affect their safety; a greater or less development of hair might modify their habits. More important changes, such as an increase in the power or dimensions of the limbs or any of the external organs, would more or less affect their mode of procuring food or the range of country which they could inhabit. It is also evident that most changes would affect, either favourably or adversely, the powers of prolonging existence. An antelope with shorter or weaker legs must necessarily suffer more from the attacks of the feline carnivora; the passenger pigeon with less powerful wings would sooner or later be affected in its powers of procuring a regular supply of food; and in both cases the result must necessarily be a diminution of the population of the modified species. If, on the other hand, any species should produce a variety having slightly increased powers of preserving existence, that variety must inevitably in time acquire a superiority in numbers. These results must follow as surely as old age, intemperance, or scarcity of food produce an increased mortality. In both cases there may be many individual exceptions; but on the average the rule will invariably be found to hold good. All varieties will therefore fall into two classes—those which under the same conditions would never reach the population of the parent species, and those which would in time obtain and keep a numerical superiority. Now, let some alteration of physical conditions occur in the district—a long period of drought, a destruction of vegetation by locusts, the irruption of some new carnivorous animal seeking “pastures new”—any change in fact tending to render existence more difficult to the species in question, and tasking its utmost powers to avoid complete extermination; it is evident that, of all the individuals composing the species, those forming the least numerous and most feebly organized variety would suffer first, and, were the pressure severe, must soon become extinct. The same causes continuing in action, the parent species would next suffer, would gradually diminish in numbers, and with a recurrence of similar unfavourable conditions might also become extinct. Tho superior variety would then alone remain, and on a return to favourable circumstances would rapidly increase in numbers and occupy the place of the extinct species and variety.
Superior Varieties will ultimately Extirpate the original Species
The variety would now have replaced the species, of which it would be a more perfectly developed and more highly organized form. It would be in all respects better adapted to secure its safety, and to prolong its individual existence and that of the race. Such a variety could not return to the original form; for that form is an inferior one, and could never compete with it for existence. Granted, therefore, a “tendency” to reproduce the original type of the species, still the variety must ever remain preponderant in numbers, and under adverse physical conditions again alone survive. But this new, improved, and populous race might itself, in course of time, give rise to new varieties, exhibiting several diverging modifications of form, any of which, tending to increase the facilities for preserving existence, must, by the same general law, in their turn become predominant. Here, then, we have progression and continued divergence deduced from the general laws which regulate the existence of animals in a state of nature, and from the undisputed fact that varieties do frequently occur. It is not, however, contended that this result would be invariable; a change of physical conditions in the district might at times materially modify it, rendering the race which had been the most capable of supporting existence under the former conditions now the least so, and even causing the extinction of the newer and, for a time, superior race, while the old or parent species and its first inferior varieties continued to flourish. Variations in unimportant parts might also occur, having no perceptible effect on the life-preserving powers; and the varieties so furnished might run a course parallel with the parent species, either giving rise to further variations or returning to the former type. All we argue for is, that certain varieties have a tendency to maintain their existence longer than the original species, and this tendency must make itself felt; for though the doctrine of chances or averages can never be trusted to on a limited scale, yet, if applied to high numbers, the results come nearer to what theory demands, and, as we approach to an infinity of examples, become strictly accurate. Now the scale on which nature works is so vast—the numbers of individuals and the periods of time with which she deals approach so near to infinity, than any cause, however slight, and however liable to be veiled and counteracted by accidental circumstances, must in the end produce its full legitimate results.
The Partial Reversion of Domesticated Varieties explained
Let us now turn to domesticated animals, and inquire how varieties produced among them are affected by the principles here enunciated. The essential difference in the condition of wild and domestic animals is this,—that among the former, their well-being and very existence depend upon the full exercise and healthy condition of all their senses and physical powers, whereas, among the latter, these are only partially exercised, and in some cases are absolutely unused. A wild animal has to search, and often to labour, for every mouthful of food—to exercise sight, hearing, and smell in seeking it, and in avoiding dangers, in procuring shelter from the inclemency of the seasons, and in providing for the subsistence and safety of its offspring. There is no muscle of its body that is not called into daily and hourly activity; there is no sense or faculty that is not strengthened by continual exercise. The domestic animal, on the other hand, has food provided for it, is sheltered, and often confined, to guard it against the vicissitudes of the seasons, is carefully secured from the attacks of its natural enemies, and seldom even rears its young without human assistance. Half of its senses and faculties become quite useless, and the other half are but occasionally called into feeble exercise, while even its muscular system is only irregularly brought into action.
Now when a variety of such an animal occurs, having increased power or capacity in any organ or sense, such increase is totally useless, is never called into action, and may even exist without the animal ever becoming aware of it. In the wild animal, on the contrary, all its faculties and powers being brought into full action for the necessities of existence, any increase becomes immediately available, is strengthened by exercise, and must even slightly modify the food, the habits, and the whole economy of the race. It creates as it were a new animal, one of superior powers, and which will necessarily increase in numbers and outlive those which are inferior to it.
Again, in the domesticated animal all variations have an equal chance of continuance; and those which would decidedly render a wild animal unable to compete with its fellows and continue its existence are no disadvantage whatever in a state of domesticity. Our quickly fattening pigs, short-legged sheep pouter pigeons, and poodle dogs could never have come into existence in a state of nature, because the very first step towards such inferior forms would have led to the rapid extinction of the race; still less could they now exist in competition with their wild allies. The great speed but slight endurance of the race horse, the unwieldy strength of the ploughman’s team, would both be useless in a state of nature. If turned wild on the pampas, such animals would probably soon become extinct, or under favourable circumstances might each gradually lose those extreme qualities which would never be called into action, and in a few generations revert to a common type, which must be that in which the various powers and faculties are so proportioned to each other as to be best adapted to procure food and secure safety,—that in which by the full exercise of every part of its organisation the animal can alone continue to live. Domestic varieties, when turned wild, must return to something near the type of the original wild stock, or become altogether extinct.6
We see, then, that no inferences as to the permanence of varieties in a state of nature can be deduced from the observations of those occurring among domestic animals. The two are so much opposed to each other in every circumstance of their existence, that what applies to the one is almost sure not to apply to the other. Domestic animals are abnormal, irregular, artificial; they are subject to variations which never occur and never can occur in a state of nature: their very existence depends altogether on human care; so far are many of them removed from that just proportion of faculties, that true balance of organisation, by means of which alone an animal left to its own resources can preserve its existence and continue its race.
Lamarck’s Hypothesis very different from that now advanced
The hypothesis of Lamarck—that progressive changes in species have been produced by the attempts of animals to increase the development of their own organs, and thus modify their structure and habits—has been repeatedly and easily refuted by all writers on the subject of varieties and species, and it seems to have been considered that when this was done the whole question has been finally settled; but the view here developed renders such hypothesis quite unnecessary, by showing that similar results must be produced by the action of principles constantly at work in nature. The powerful retractile talons of the falcon-and the cat-tribes have not been produced or increased by the volition of those animals; but among the different varieties which occurred in the earlier and less highly organized forms of these groups, those always survived longest which had the greatest facilities for seizing their prey. Neither did the giraffe acquire its long neck by desiring to reach the foliage of the more lofty shrubs, and constantly stretching its neck for the purpose, but because any varieties which occurred among its antitypes with a longer neck than usual at once secured a fresh range of pasture over the same ground as their shorter-necked companions, and on the first scarcity of food were thereby enabled to outlive them. Even the peculiar colours of many animals, more especially of insects, so closely resembling the soil or leaves or bark on which they habitually reside, are explained on the same principle; for though in the course of ages varieties of many tints may have occurred, yet those races having colours best adapted to concealment from their enemies would inevitably survive the longest. We have also here an acting cause to account for that balance so often observed in nature,—a deficiency in one set of organs always being compensated by an increased development of some others—powerful wings accompanying weak feet, or great velocity making up for the absence of defensive weapons; for it has been shown that all varieties in which an unbalanced deficiency occurred could not long continue their existence. The action of this principle is exactly like that of the centrifugal governor of the steam engine, which checks and corrects any irregularities almost before they become evident; and in like manner no unbalanced deficiency in the animal kingdom can ever reach any conspicuous magnitude, because it would make itself felt at the very first step, by rendering existence difficult and extinction almost sure soon to follow. An origin such as is here advocated will also agree with the peculiar character of the modifications of form and structure which obtain in organized beings—the many lines of divergence from a central type, the increasing efficiency and power of a particular organ through a succession of allied species, and the remarkable persistence of unimportant parts, such as colour, texture of plumage and hair, form of horns or crests, through a series of species differing considerably in more essential characters. It also furnishes us with a reason for that “more specialized structure” which Professor Owen states to be a characteristic of recent compared with extinct forms, and which would evidently be the result of the progressive modification of any organ applied to a special purpose in the animal economy.
Conclusion
We believe we have now shown that there is a tendency in nature to the continued progression of certain classes of varieties further and further from the original type—a progression to which there appears no reason to assign any definite limits—and that the same principle which produces this result in a state of nature will also explain why domestic varieties have a tendency, when they become wild, to revert to the original type. This progression, by minute steps, in various directions, but always checked and balanced by the necessary conditions, subject to which alone existence can be preserved, may, it is believed, be followed out so as to agree with all the phænomena presented by organized beings, their extinction and succession in past ages, and all the extraordinary modifications of form, instinct and habits which they exhibit.
III.
MIMICRY, AND OTHER PROTECTIVE RESEMBLANCES AMONG ANIMALS
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 phænomena 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 may have been supported. The great name of Edward Forbes did not prevent his theory of “Polarity in the distribution of Organic beings in Time” from dying a natural death; but the most striking illustration of the behaviour of a false theory is to be found in the “Circular and Quinarian System” of classification propounded by MacLeay, and developed by Swainson, with an amount of knowledge and ingenuity that have rarely been surpassed. This theory was eminently attractive, both from its symmetry and completeness, and from the interesting nature of the varied analogies and affinities which it brought to light and made use of. The series of Natural History volumes in “Lardner’s Cabinet Cyclopædia,” in which Mr. Swainson developed it in most departments of the animal kingdom, made it widely known; and in fact for a long time these were the best and almost the only popular text-books for the rising generation of naturalists. It was favourably received too by the older school, which was perhaps rather an indication of its unsoundness. A considerable number of well-known naturalists either spoke approvingly of it, or advocated similar principles, and for a good many years it was decidedly in the ascendent. With such a favourable introduction, and with such talented exponents, it must have become established if it had had any germ of truth in it; yet it quite died out in a few short years, its very existence is now a matter of history; and so rapid was its fall that its talented creator, Swainson, perhaps lived to be the last man who believed in it.
Such is the course of a false theory. That of a true one 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.
Importance of the Principle of Utility
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 the races which possess them. This great principle gives us a clue which we can follow out in the study of many recondite phænomena, and leads us to seek a meaning and a purpose of some definite character in minutiæ which we should be otherwise almost sure to pass over as insignificant or unimportant.
Popular Theories of Colour in Animals
The adaptation of the external colouring of animals to their conditions of life has long been recognised, 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 phænomena, and to be contradicted by many well-known facts. For example, wild rabbits are always of grey 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 to food, since in many cases the species does not feed on the substance it resembles, and when it does, no reasonable connexion 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 phænomena which may be classed under the head of useful or protective resemblances.
Importance of Concealment as Influencing Colour
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.