The Origin of Species by Means of Natural Selection · Charles Darwin
CHAPTER VII. MISCELLANEOUS OBJECTIONS TO THE THEORY OF NATURAL SELECTION.
All Mr. Mivart’s objections will be, or have been, considered in the present volume. The one new point which appears to have struck many readers is, “That natural selection is incompetent to account for the incipient stages of useful structures.” This subject is intimately connected with that of the gradation of the characters, often accompanied by a change of function, for instance, the conversion of a swim-bladder into lungs, points which were discussed in the last chapter under two headings. Nevertheless, I will here consider in some detail several of the cases advanced by Mr. Mivart, selecting those which are the most illustrative, as want of space prevents me from considering all. The giraffe, by its lofty stature, much elongated neck, fore legs, head and tongue, has its whole frame beautifully adapted for browsing on the higher branches of trees. It can thus obtain food beyond the reach of the other Ungulata or hoofed animals inhabiting the same country; and this must be a great advantage to it during dearths. The Niata cattle in South America show us how small a difference in structure may make, during such periods, a great difference in preserving an animal’s life. These cattle can browse as well as others on grass, but from the projection of the lower jaw they cannot, during the often recurrent droughts, browse on the twigs of trees, reeds, &c., to which food the common cattle and horses are then driven; so that at these times the Niatas perish, if not fed by their owners. Before coming to Mr. Mivart’s objections, it may be well to explain once again how natural selection will act in all ordinary cases. Man has modified some of his animals, without necessarily having attended to special points of structure, by simply preserving and breeding from the fleetest individuals, as with the race-horse and greyhound, or as with the game-cock, by breeding from the victorious birds.
So under nature with the nascent giraffe, the individuals which were the highest browsers and were able during dearths to reach even an inch or two above the others, will often have been preserved; for they will have roamed over the whole country in search of food. That the individuals of the same species often differ slightly in the relative lengths of all their parts may be seen in many works of natural history, in which careful measurements are given. These slight proportional differences, due to the laws of growth and variation, are not of the slightest use or importance to most species. But it will have been otherwise with the nascent giraffe, considering its probable habits of life; for those individuals which had some one part or several parts of their bodies rather more elongated than usual, would generally have survived. These will have intercrossed and left offspring, either inheriting the same bodily peculiarities, or with a tendency to vary again in the same manner; while the individuals less favoured in the same respects will have been the most liable to perish. We here see that there is no need to separate single pairs, as man does, when he methodically improves a breed: natural selection will preserve and thus separate all the superior individuals, allowing them freely to intercross, and will destroy all the inferior individuals. By this process long-continued, which exactly corresponds with what I have called unconscious selection by man, combined, no doubt, in a most important manner with the inherited effects of the increased use of parts, it seems to me almost certain that an ordinary hoofed quadruped might be converted into a giraffe.
To this conclusion Mr. Mivart brings forward two objections. One is that the increased size of the body would obviously require an increased supply of food, and he considers it as “very problematical whether the disadvantages thence arising would not, in times of scarcity, more than counterbalance the advantages.” But as the giraffe does actually exist in large numbers in Africa, and as some of the largest antelopes in the world, taller than an ox, abound there, why should we doubt that, as far as size is concerned, intermediate gradations could formerly have existed there, subjected as now to severe dearths. Assuredly the being able to reach, at each stage of increased size, to a supply of food, left untouched by the other hoofed quadrupeds of the country, would have been of some advantage to the nascent giraffe. Nor must we overlook the fact, that increased bulk would act as a protection against almost all beasts of prey excepting the lion; and against this animal, its tall neck—and the taller the better—would, as Mr. Chauncey Wright has remarked, serve as a watch-tower. It is from this cause, as Sir S. Baker remarks, that no animal is more difficult to stalk than the giraffe. This animal also uses its long neck as a means of offence or defence, by violently swinging its head armed with stump-like horns. The preservation of each species can rarely be determined by any one advantage, but by the union of all, great and small.
Mr. Mivart then asks (and this is his second objection), if natural selection be so potent, and if high browsing be so great an advantage, why has not any other hoofed quadruped acquired a long neck and lofty stature, besides the giraffe, and, in a lesser degree, the camel, guanaco and macrauchenia? Or, again, why has not any member of the group acquired a long proboscis? With respect to South Africa, which was formerly inhabited by numerous herds of the giraffe, the answer is not difficult, and can best be given by an illustration. In every meadow in England, in which trees grow, we see the lower branches trimmed or planed to an exact level by the browsing of the horses or cattle; and what advantage would it be, for instance, to sheep, if kept there, to acquire slightly longer necks? In every district some one kind of animal will almost certainly be able to browse higher than the others; and it is almost equally certain that this one kind alone could have its neck elongated for this purpose, through natural selection and the effects of increased use. In South Africa the competition for browsing on the higher branches of the acacias and other trees must be between giraffe and giraffe, and not with the other ungulate animals.
Why, in other quarters of the world, various animals belonging to this same order have not acquired either an elongated neck or a proboscis, cannot be distinctly answered; but it is as unreasonable to expect a distinct answer to such a question as why some event in the history of mankind did not occur in one country while it did in another. We are ignorant with respect to the conditions which determine the numbers and range of each species, and we cannot even conjecture what changes of structure would be favourable to its increase in some new country. We can, however, see in a general manner that various causes might have interfered with the development of a long neck or proboscis. To reach the foliage at a considerable height (without climbing, for which hoofed animals are singularly ill-constructed) implies greatly increased bulk of body; and we know that some areas support singularly few large quadrupeds, for instance South America, though it is so luxuriant, while South Africa abounds with them to an unparalleled degree. Why this should be so we do not know; nor why the later tertiary periods should have been much more favourable for their existence than the present time. Whatever the causes may have been, we can see that certain districts and times would have been much more favourable than others for the development of so large a quadruped as the giraffe.
In order that an animal should acquire some structure specially and largely developed, it is almost indispensable that several other parts should be modified and coadapted. Although every part of the body varies slightly, it does not follow that the necessary parts should always vary in the right direction and to the right degree. With the different species of our domesticated animals we know that the parts vary in a different manner and degree, and that some species are much more variable than others. Even if the fitting variations did arise, it does not follow that natural selection would be able to act on them and produce a structure which apparently would be beneficial to the species. For instance, if the number of individuals existing in a country is determined chiefly through destruction by beasts of prey—by external or internal parasites, &c.—as seems often to be the case, then natural selection will be able to do little, or will be greatly retarded, in modifying any particular structure for obtaining food. Lastly, natural selection is a slow process, and the same favourable conditions must long endure in order that any marked effect should thus be produced. Except by assigning such general and vague reasons, we cannot explain why, in many quarters of the world, hoofed quadrupeds have not acquired much elongated necks or other means for browsing on the higher branches of trees.
Objections of the same nature as the foregoing have been advanced by many writers. In each case various causes, besides the general ones just indicated, have probably interfered with the acquisition through natural selection of structures, which it is thought would be beneficial to certain species. One writer asks, why has not the ostrich acquired the power of flight? But a moment’s reflection will show what an enormous supply of food would be necessary to give to this bird of the desert force to move its huge body through the air. Oceanic islands are inhabited by bats and seals, but by no terrestrial mammals; yet as some of these bats are peculiar species, they must have long inhabited their present homes. Therefore Sir C. Lyell asks, and assigns certain reasons in answer, why have not seals and bats given birth on such islands to forms fitted to live on the land? But seals would necessarily be first converted into terrestrial carnivorous animals of considerable size, and bats into terrestrial insectivorous animals; for the former there would be no prey; for the bats ground-insects would serve as food, but these would already be largely preyed on by the reptiles or birds, which first colonise and abound on most oceanic islands.
Gradations of structure, with each stage beneficial to a changing species, will be favoured only under certain peculiar conditions. A strictly terrestrial animal, by occasionally hunting for food in shallow water, then in streams or lakes, might at last be converted into an animal so thoroughly aquatic as to brave the open ocean. But seals would not find on oceanic islands the conditions favourable to their gradual reconversion into a terrestrial form. Bats, as formerly shown, probably acquired their wings by at first gliding through the air from tree to tree, like the so-called flying-squirrels, for the sake of escaping from their enemies, or for avoiding falls; but when the power of true flight had once been acquired, it would never be reconverted back, at least for the above purposes, into the less efficient power of gliding through the air. Bats, might, indeed, like many birds, have had their wings greatly reduced in size, or completely lost, through disuse; but in this case it would be necessary that they should first have acquired the power of running quickly on the ground, by the aid of their hind legs alone, so as to compete with birds or other ground animals; and for such a change a bat seems singularly ill-fitted. These conjectural remarks have been made merely to show that a transition of structure, with each step beneficial, is a highly complex affair; and that there is nothing strange in a transition not having occurred in any particular case.
Lastly, more than one writer has asked why have some animals had their mental powers more highly developed than others, as such development would be advantageous to all? Why have not apes acquired the intellectual powers of man? Various causes could be assigned; but as they are conjectural, and their relative probability cannot be weighed, it would be useless to give them. A definite answer to the latter question ought not to be expected, seeing that no one can solve the simpler problem, why, of two races of savages, one has risen higher in the scale of civilisation than the other; and this apparently implies increased brain power. We will return to Mr. Mivart’s other objections. Insects often resemble for the sake of protection various objects, such as green or decayed leaves, dead twigs, bits of lichen, flowers, spines, excrement of birds, and living insects; but to this latter point I shall hereafter recur. The resemblance is often wonderfully close, and is not confined to colour, but extends to form, and even to the manner in which the insects hold themselves. The caterpillars which project motionless like dead twigs from the bushes on which they feed, offer an excellent instance of a resemblance of this kind. The cases of the imitation of such objects as the excrement of birds, are rare and exceptional. On this head, Mr. Mivart remarks, “As, according to Mr. Darwin’s theory, there is a constant tendency to indefinite variation, and as the minute incipient variations will be in _all directions_, they must tend to neutralize each other, and at first to form such unstable modifications that it is difficult, if not impossible, to see how such indefinite oscillations of infinitesimal beginnings can ever build up a sufficiently appreciable resemblance to a leaf, bamboo, or other object, for natural selection to seize upon and perpetuate.”
But in all the foregoing cases the insects in their original state no doubt presented some rude and accidental resemblance to an object commonly found in the stations frequented by them. Nor is this at all improbable, considering the almost infinite number of surrounding objects and the diversity in form and colour of the hosts of insects which exist. As some rude resemblance is necessary for the first start, we can understand how it is that the larger and higher animals do not (with the exception, as far as I know, of one fish) resemble for the sake of protection special objects, but only the surface which commonly surrounds them, and this chiefly in colour. Assuming that an insect originally happened to resemble in some degree a dead twig or a decayed leaf, and that it varied slightly in many ways, then all the variations which rendered the insect at all more like any such object, and thus favoured its escape, would be preserved, while other variations would be neglected and ultimately lost; or, if they rendered the insect at all less like the imitated object, they would be eliminated. There would indeed be force in Mr. Mivart’s objection, if we were to attempt to account for the above resemblances, independently of natural selection, through mere fluctuating variability; but as the case stands there is none.
Nor can I see any force in Mr. Mivart’s difficulty with respect to “the last touches of perfection in the mimicry;” as in the case given by Mr. Wallace, of a walking-stick insect (Ceroxylus laceratus), which resembles “a stick grown over by a creeping moss or jungermannia.” So close was this resemblance, that a native Dyak maintained that the foliaceous excrescences were really moss. Insects are preyed on by birds and other enemies whose sight is probably sharper than ours, and every grade in resemblance which aided an insect to escape notice or detection, would tend towards its preservation; and the more perfect the resemblance so much the better for the insect. Considering the nature of the differences between the species in the group which includes the above Ceroxylus, there is nothing improbable in this insect having varied in the irregularities on its surface, and in these having become more or less green-coloured; for in every group the characters which differ in the several species are the most apt to vary, while the generic characters, or those common to all the species, are the most constant.
The Greenland whale is one of the most wonderful animals in the world, and the baleen, or whalebone, one of its greatest peculiarities. The baleen consists of a row, on each side of the upper jaw, of about 300 plates or laminæ, which stand close together transversely to the longer axis of the mouth. Within the main row there are some subsidiary rows. The extremities and inner margins of all the plates are frayed into stiff bristles, which clothe the whole gigantic palate, and serve to strain or sift the water, and thus to secure the minute prey on which these great animals subsist. The middle and longest lamina in the Greenland whale is ten, twelve, or even fifteen feet in length; but in the different species of Cetaceans there are gradations in length; the middle lamina being in one species, according to Scoresby, four feet, in another three, in another eighteen inches, and in the Balænoptera rostrata only about nine inches in length. The quality of the whalebone also differs in the different species.
With respect to the baleen, Mr. Mivart remarks that if it “had once attained such a size and development as to be at all useful, then its preservation and augmentation within serviceable limits would be promoted by natural selection alone. But how to obtain the beginning of such useful development?” In answer, it may be asked, why should not the early progenitors of the whales with baleen have possessed a mouth constructed something like the lamellated beak of a duck? Ducks, like whales, subsist by sifting the mud and water; and the family has sometimes been called _Criblatores_, or sifters. I hope that I may not be misconstrued into saying that the progenitors of whales did actually possess mouths lamellated like the beak of a duck. I wish only to show that this is not incredible, and that the immense plates of baleen in the Greenland whale might have been developed from such lamellæ by finely graduated steps, each of service to its possessor.
The beak of a shoveller-duck (Spatula clypeata) is a more beautiful and complex structure than the mouth of a whale. The upper mandible is furnished on each side (in the specimen examined by me) with a row or comb formed of 188 thin, elastic lamellæ, obliquely bevelled so as to be pointed, and placed transversely to the longer axis of the mouth. They arise from the palate, and are attached by flexible membrane to the sides of the mandible. Those standing towards the middle are the longest, being about one-third of an inch in length, and they project fourteen one-hundredths of an inch beneath the edge. At their bases there is a short subsidiary row of obliquely transverse lamellæ. In these several respects they resemble the plates of baleen in the mouth of a whale. But towards the extremity of the beak they differ much, as they project inward, instead of straight downward. The entire head of the shoveller, though incomparably less bulky, is about one-eighteenth of the length of the head of a moderately large Balænoptera rostrata, in which species the baleen is only nine inches long; so that if we were to make the head of the shoveller as long as that of the Balænoptera, the lamellæ would be six inches in length, that is, two-thirds of the length of the baleen in this species of whale. The lower mandible of the shoveller-duck is furnished with lamellæ of equal length with these above, but finer; and in being thus furnished it differs conspicuously from the lower jaw of a whale, which is destitute of baleen. On the other hand, the extremities of these lower lamellæ are frayed into fine bristly points, so that they thus curiously resemble the plates of baleen. In the genus Prion, a member of the distinct family of the Petrels, the upper mandible alone is furnished with lamellæ, which are well developed and project beneath the margin; so that the beak of this bird resembles in this respect the mouth of a whale.
From the highly developed structure of the shoveller’s beak we may proceed (as I have learned from information and specimens sent to me by Mr. Salvin), without any great break, as far as fitness for sifting is concerned, through the beak of the Merganetta armata, and in some respects through that of the Aix sponsa, to the beak of the common duck. In this latter species the lamellæ are much coarser than in the shoveller, and are firmly attached to the sides of the mandible; they are only about fifty in number on each side, and do not project at all beneath the margin. They are square-topped, and are edged with translucent, hardish tissue, as if for crushing food. The edges of the lower mandible are crossed by numerous fine ridges, which project very little. Although the beak is thus very inferior as a sifter to that of a shoveller, yet this bird, as every one knows, constantly uses it for this purpose. There are other species, as I hear from Mr. Salvin, in which the lamellæ are considerably less developed than in the common duck; but I do not know whether they use their beaks for sifting the water.
Turning to another group of the same family. In the Egyptian goose (Chenalopex) the beak closely resembles that of the common duck; but the lamellæ are not so numerous, nor so distinct from each other, nor do they project so much inward; yet this goose, as I am informed by Mr. E. Bartlett, “uses its bill like a duck by throwing the water out at the corners.” Its chief food, however, is grass, which it crops like the common goose. In this latter bird the lamellæ of the upper mandible are much coarser than in the common duck, almost confluent, about twenty-seven in number on each side, and terminating upward in teeth-like knobs. The palate is also covered with hard rounded knobs. The edges of the lower mandible are serrated with teeth much more prominent, coarser and sharper than in the duck. The common goose does not sift the water, but uses its beak exclusively for tearing or cutting herbage, for which purpose it is so well fitted that it can crop grass closer than almost any other animal. There are other species of geese, as I hear from Mr. Bartlett, in which the lamellæ are less developed than in the common goose.
We thus see that a member of the duck family, with a beak constructed like that of a common goose and adapted solely for grazing, or even a member with a beak having less well-developed lamellæ, might be converted by small changes into a species like the Egyptian goose—this into one like the common duck—and, lastly, into one like the shoveller, provided with a beak almost exclusively adapted for sifting the water; for this bird could hardly use any part of its beak, except the hooked tip, for seizing or tearing solid food. The beak of a goose, as I may add, might also be converted by small changes into one provided with prominent, recurved teeth, like those of the Merganser (a member of the same family), serving for the widely different purpose of securing live fish.
Returning to the whales. The Hyperoodon bidens is destitute of true teeth in an efficient condition, but its palate is roughened, according to Lacepede, with small unequal, hard points of horn. There is, therefore, nothing improbable in supposing that some early Cetacean form was provided with similar points of horn on the palate, but rather more regularly placed, and which, like the knobs on the beak of the goose, aided it in seizing or tearing its food. If so, it will hardly be denied that the points might have been converted through variation and natural selection into lamellæ as well-developed as those of the Egyptian goose, in which case they would have been used both for seizing objects and for sifting the water; then into lamellæ like those of the domestic duck; and so onward, until they became as well constructed as those of the shoveller, in which case they would have served exclusively as a sifting apparatus. From this stage, in which the lamellæ would be two-thirds of the length of the plates of baleen in the Balænoptera rostrata, gradations, which may be observed in still-existing Cetaceans, lead us onward to the enormous plates of baleen in the Greenland whale. Nor is there the least reason to doubt that each step in this scale might have been as serviceable to certain ancient Cetaceans, with the functions of the parts slowly changing during the progress of development, as are the gradations in the beaks of the different existing members of the duck-family. We should bear in mind that each species of duck is subjected to a severe struggle for existence, and that the structure of every part of its frame must be well adapted to its conditions of life.
Text from Project Gutenberg, public domain in the United States.