Theory of Natural Selection. I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views, as some of the previous discussions may thus be made clearer; but it would be useless to discuss all of them, as many have been made by writers who have not taken the trouble to understand the subject. Thus a distinguished German naturalist has asserted that the weakest part of my theory is, that I consider all organic beings as imperfect: what I have really said is, that all are not as perfect as they might have been in relation to their conditions; and this is shown to be the case by so many native forms in many quarters of the world having yielded their places to intruding foreigners. Nor can organic beings, even if they were at any one time perfectly adapted to their conditions of life, have remained so, when their conditions changed, unless they themselves likewise changed; and no one will dispute that the physical conditions of each country, as well as the numbers and kinds of its inhabitants, have undergone many mutations.
A critic has lately insisted, with some parade of mathematical accuracy, that longevity is a great advantage to all species, so that he who believes in natural selection must arrange his genealogical tree in such a manner that all the descendants have longer lives than their progenitors! Cannot our critic conceive that a biennial plant or one of the lower animals might range into a cold climate and perish there every winter; and yet, owing to advantages gained through natural selection, survive from year to year by means of its seeds or ova? Mr. E. Ray Lankester has recently discussed this subject, and he concludes, as far as its extreme complexity allows him to form a judgment, that longevity is generally related to the standard of each species in the scale of organisation, as well as to the amount of expenditure in reproduction and in general activity. And these conditions have, it is probable, been largely determined through natural selection.
It has been argued that, as none of the animals and plants of Egypt, of which we know anything, have changed during the last three or four thousand years, so probably have none in any part of the world. But, as Mr. G. H. Lewes has remarked, this line of argument proves too much, for the ancient domestic races figured on the Egyptian monuments, or embalmed, are closely similar or even identical with those now living; yet all naturalists admit that such races have been produced through the modification of their original types. The many animals which have remained unchanged since the commencement of the glacial period, would have been an incomparably stronger case, for these have been exposed to great changes of climate and have migrated over great distances; whereas, in Egypt, during the last several thousand years, the conditions of life, as far as we know, have remained absolutely uniform. The fact of little or no modification having been effected since the glacial period would have been of some avail against those who believe in an innate and necessary law of development, but is powerless against the doctrine of natural selection or the survival of the fittest, which implies that when variations or individual differences of a beneficial nature happen to arise, these will be preserved; but this will be effected only under certain favourable circumstances.
The celebrated palæontologist, Bronn, at the close of his German translation of this work, asks, how, on the principle of natural selection, can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions, they might live together; and if we lay on one side polymorphic species, in which the variability seems to be of a peculiar nature, and all mere temporary variations, such as size, aborted, etc., the more permanent varieties are generally found, as far as I can discover, inhabiting distinct stations, - such as high land or low land, dry or moist districts. Moreover, in the case of animals which wander much about and cross freely, their varieties seem to be generally confined to distinct regions.
Bronn also insists that distinct species never differ from each other in single characters, but in many parts; and he asks, how it always comes that many parts of the organisation should have been modified at the same time through variation and natural selection . But there is no necessity for supposing that all the parts of any being have been simultaneously modified. The most striking modifications, excellently adapted for some purpose, might, as was formerly remarked, be acquired by successive variations, if slight, first in one part and then in another; and as they would be transmitted all together, they would appear to us as if they had been simultaneously developed. The best answer, however, to the above objection is afforded by those domestic races which have been modified, chiefly through man's power of selection, for some special purpose. Look at the race and dray horse, or at the greyhound and mastiff. Their whole frames and even their mental characteristics have been modified; but if we could trace each step in the history of their transformation, - and the latter steps can be traced, - we should not see great and simultaneous changes, but first one part and then another slightly modified and improved. Even when selection has been applied by man to some one character alone, - of which our cultivated plants offer the best instances, - it will invariably be found that although this one part, whether it be the flower, fruit, or leaves, has been greatly changed, almost all the other parts have been slightly modified. This may be attributed partly to the principle of correlated growth, and partly to so-called spontaneous variation.
A much more serious objection has been urged by Bronn, and recently by Broca, namely, that many characters appear to be of no service whatever to their possessors, and therefore cannot have been influenced through natural selection. Bronn adduces the length of the ears and tails in the different species of hares and mice, - the complex folds of enamel in the teeth of many animals, and a multitude of analogous cases. With respect to plants, this subject has been discussed by Nageli in an admirable essay. He admits that natural selection has effected much, but he insists that the families of plants differ chiefly from each other in morphological characters, which appear to be quite unimportant for the welfare of the species. He consequently believes in an innate tendency towards progressive and more perfect development. He specifies the arrangement of the cells in the tissues, and of the leaves on the axis, as cases in which natural selection could not have acted. To these may be added the numerical divisions in the parts of the flower, the position of the ovules, the shape of the seed, when not of any use for dissemination, etc.
There is much force in the above objection. Nevertheless, we ought, in the first place, to be extremely cautious in pretending to decide what structures now are, or have formerly been, use to each species. In the second place, it should always be borne in mind that when part is modified, so will be other parts, through certain dimly seen causes, such as an increased or diminished flow of nutriment to a part, mutual pressure, an early developed part affecting one subsequently developed, and so forth, - as well as through other causes which lead to the many mysterious cases of correlation, which we do not in the least understand. These agencies may be all grouped together, for the sake of brevity, under the expression of the laws of growth. In the third place, we have to allow for the direct and definite action of changed conditions of life, and for so-called spontaneous variations, in which the nature of the conditions apparently plays a quite subordinate part. Bud variations, such as the appearance of a moss-rose on a common rose, or of a nectarine on a peach tree offer good instances of spontaneous variations; but even in these cases, if we bear in mind the power of a minute drop of poison in producing complex galls, we ought not to feel too sure that the above variations are not the effect of some local change in the nature of the sap, due to some change in the conditions. There must be some efficient cause for each slight individual difference, as well as for more strongly marked variations which occasionally arise; and if the unknown cause were to act persistently, it is almost certain that all the individuals of the species would be similarly modified.
In the earlier editions of this work I under-rated, as it now seems probable, the frequency and importance of modifications due to spontaneous variability. But it is impossible to attribute to this cause the innumerable structures which are so well adapted to the habits of life of each species. I can no more believe in this than that the well-adapted form of a race-horse or greyhound, which before the principle of selection by man was well understood, excited so much surprise in the minds of the older naturalists, can thus be explained.
It may be worth while to illustrate some of the foregoing remarks. With respect to the assumed inutility of various parts and organs, it is hardly necessary to observe that even in the higher and best-known animals many structures exist, which are so highly developed that no one doubts that they are of importance, yet their use has not been, or has only recently been, ascertained. As Bronn gives the length of the ears and tail in the several species of mice as instances, though trifling ones, of differences in structure which can be of no special use, I may mention that, according to Dr. Schöbl, the external ears of the common mouse are supplied in an extraordinary manner with nerves, so that they no doubt serve as tactile organs; hence the length of the ears can hardly be quite unimportant. We shall, also, presently see that the tail is a highly useful prehensile organ to some of the species; and its use would be much influenced by its length.
With respect to plants, to which on account of Nägeli's essay I shall confine myself in the following remarks, it will be admitted that the flowers of orchids present a multitude of curious structures, which a few years ago would have been considered as mere morphological differences without any special function; but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects, and have probably been gained through natural selection. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils, and their arrangement, could have been of any service, but now we know this to be the case.
In certain whole groups of plants the ovules stand erect, and in others they are suspended; and within the same ovarium of some few plants, one ovule holds the former and a second ovule the latter position. These positions seem at first purely morphological, or of no physiological signification; but Dr. Hooker informs me that within the same ovarium, the upper ovules alone in some cases, and in other cases the lower ones alone are fertilised; and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium. If so, the position of the ovules, even when one is erect and the other suspended within the same ovarium, would follow from the selection of any slight deviations in position which favoured their fertilisation, and the production of seed.
Several plants belonging to distinct orders habitually produce flowers of two kinds, - the one open of the ordinary structure, the other closed and imperfect. These two kinds of flowers sometimes differ wonderfully in structure, yet may be seen to graduate into each other on the same plant. The ordinary and open flowers can be intercrossed; and the benefits which certainly are derived from this process are thus secured. The closed and imperfect flowers are, however, manifestly of high importance, as they yield with the utmost safety a large stock of seed, with the expenditure of wonderfully little pollen. The two kinds of flowers often differ much, as just stated, in structure. The petals in the imperfect flowers almost always consist of mere rudiments, and the pollen-grains are reduced in diameter. In Ononis columnæ five of the alternate stamens are rudimentary; and in some species of Viola three stamens are in this state, two retaining their proper function, but being of very small size. In six out of thirty of the closed flowers in an Indian violet (name unknown, for the plants have never produced with me perfect flowers), the sepals are reduced from the normal number of five to three. In one section of the Malpighiaceæ the closed flowers, according to A. de Jussieu, are still further modified, for the five stamens which stand opposite to the sepals are all aborted, sixth stamen standing opposite to a petal being alone developed; and this stamen is not present in the ordinary flowers of these species; the style is aborted; and the ovaria are reduced from three to two. Now although natural selection may well have had the power to prevent some of the flowers from expanding, and to reduce the amount of pollen, when rendered by the closure of the flowers superfluous, yet hardly any of the above special modifications can have been thus determined, but must have followed from the laws of growth, including the functional inactivity of parts, during the progress of the reduction of the pollen and the closure of the flowers.
It is so necessary to appreciate the important effects of the laws of growth, that I will give some additional cases of another kind, namely of differences in the same part or organ, due to differences in relative position on the same plant. In the Spanish chestnut, and in certain fir-trees, the angles of divergence of the leaves differ, according to Schacht, in the nearly horizontal and in the upright branches. In the common rue and some other plants, one flower, usually the central or terminal one, opens first, and has five sepals and petals, and five divisions to the ovarium; whilst all the other flowers on the plant are tetramerous. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous, whilst the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. In many Compositæ and Umbelliferæ (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre; and this seems often connected with the abortion of the reproductive organs. It is a more curious fact, previously referred to, that the achenes or seeds of the circumference and centre sometimes differ greatly in form, colour, and other characters. In Carthamus and some other Compositæ the central achenes alone are furnished with a pappus; and in Hyoseris the same head yields achenes of three different forms. In certain Umbelliferæ the exterior seeds, according to Tausch, are orthospermous, and the central one clospermous, and this is a character which was considered by De Candolle to be in other species of the highest systematic importance. Prof. Braun mentions a Fumariaceous genus, in which the flowers in the lower part of the spike bear oval, ribbed, one-seeded nutlets; and in the upper part of the spike, lanceolate, two-valved, and two-seeded siliques. In these several cases, with the exception of that of the well developed ray-florets, which are of service in making the flowers conspicuous to insects, natural selection cannot, as far as we can judge, have come into play, or only in a quite subordinate manner. All these modifications follow from the relative position and inter-action of the parts; and it can hardly be doubted that if all the flowers and leaves on the same plant had been subjected to the same external and internal condition, as are the flowers and leaves in certain positions, all would have been modified in the same manner.
In numerous other cases we find modifications of structure, which are considered by botanists to be generally of a highly important nature, affecting only some of the flowers on the same plant, or occurring on distinct plants, which grow close together under the same conditions. As these variations seem of no special use to the plants, they cannot have been influenced by natural selection. Of their cause we are quite ignorant; we cannot even attribute them, as in the last class of cases, to any proximate agency, such as relative position. I will give only a few instances. It is so common to observe on the same plant, flowers indifferently tetramerous, pentamerous, etc., that I need not give examples; but as numerical variations are comparatively rare when the parts are few, I may mention that, according to De Candolle, the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies), or three sepals with six petals. The manner in which the petals are folded in the bud is in most groups a very constant morphological character; but Professor Asa Gray states that with some species of Mimulus, the æstivation is almost as frequently that of the Rhinanthideæ as of the Antirrhinideæ, to which latter tribe the genus belongs. Auguste de Saint-Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutacese with a single ovary, but in some species flowers may be found on the same plant, and even in the same panicle, with either one or two ovaries. In Helianthemum the capsule has been described as unilocular or trilocular; and in H. mutabile, Une lame, plus ou moins large, s'etend entre le pericarpe et le placenta. In the flowers of Saponaria officinalis, Dr. Masters has observed instances of both marginal and free central placentation. Lastly, Saint-Hilaire found towards the southern extreme of the range of Gomphia oleæformis two forms which he did not at first doubt were distinct species, but he subsequently saw them growing on the same bush; and he then adds, Voilà donc dans un même individu des loges et un style qui se rattachent tantôt à un axe verticale et tantôt à un gynobase.
We thus see that with plants many morphological changes may be attributed to the laws of growth and the inter-action of parts, independently of natural selection. But with respect to Nägeli's doctrine of an innate tendency towards perfection or progressive development, can it be said in the case of these strongly pronounced variations, that the plants have been caught in the act of progressing towards a higher state of development? On the contrary, I should infer from the mere fact of the parts in question differing or varying greatly on the same plant, that such modifications were of extremely small importance to the plants themselves, of whatever importance they may generally be to us for our classifications. The acquisition of a useless part can hardly be said to raise an organism in the natural scale; and in the case of the imperfect, closed flowers above described, if any new principle has to be invoked, it must be one of retrogression rather than of progression; and so it must be with many parasitic and degraded animals. We are ignorant of the exciting cause of the above specified modifications; but if the unknown cause were to act almost uniformly for a length of time, we may infer that the result would be almost uniform; and in this case all the individuals of the species would be modified in the same manner.
From the fact of the above characters being unimportant for the welfare of the species, any slight variations which occurred in them would not have been accumulated and augmented through natural selection. A structure which has been developed through long-continued selection, when it ceases to be of service to a species, generally becomes variable, as we see with rudimentary organs; for it will no longer be regulated by this same power of selection. But when, from the nature of the organism and of the conditions, modifications have been induced which are unimportant for the welfare of the species, they may be, and apparently often have been, transmitted in nearly the same state to numerous, otherwise modified, descendants. It cannot have been of much importance to the greater number of mammals, birds, or reptiles, whether they were clothed with hair, feathers, or scales; yet hair has been transmitted to almost all mammals, feathers to all birds, and scales to all true reptiles. A structure, whatever it may be, which is common to many allied forms, is ranked by us as of high systematic importance, and consequently is often assumed to be of high vital importance to the species. Thus, as I am inclined to believe, differences, which we consider as important - such as the arrangement of the leaves, the divisions of the flower or of the ovarium, the position of the ovules, etc. - first appeared in many cases as fluctuating variations, which sooner or later became constant through the nature of the organism and of the surrounding conditions, as well as through the intercrossing of distinct individuals, but not through natural selection; for as these morphological characters do not affect the welfare of the species, any slight deviations in them could not have been governed or accumulated through this latter agency. It is a strange result which we thus arrive at, namely that characters of slight vital importance to the species, are the most important to the systematist; but, as we shall hereafter see when we treat of the genetic principle of classification, this is by no means so paradoxical as it may at first appear.
Although we have no good evidence of the existence in organic beings of an innate tendency towards progressive development, yet this necessarily follows, as I have attempted to show in the fourth chapter, through the continued action of natural selection. For the best definition which has ever been given of a high standard of organisation, is the degree to which the parts have been specialised or differentiated; and natural selection tends towards this end, inasmuch as the parts are thus enabled to perform their functions more efficiently.
A distinguished zoologist, Mr. St. George Mivart, has recently collected all the objections which have ever been advanced by myself and others against the theory of natural selection, as propounded by Mr. Wallace and myself, and has illustrated them with admirable art and force. When thus marshalled, they make a formidable array; and as it forms no part of Mr. Mivart's plan to give the various facts and considerations opposed to his conclusions, no slight effort of reason and memory is left to the reader, who may wish to weigh the evidence on both sides. When discussing special cases, Mr. Mivart passes over the effects of the increased use and disuse of parts, which I have always maintained to be highly important, and have treated in my Variation under Domestication at greater length than, as I believe, any other writer. He likewise often assumes that I attribute nothing to variation, independently of natural selection, whereas in the work just referred to I have collected a greater number of well-established cases than can be found in any other work known to me. My judgment may not be trustworthy, but after reading with care Mr. Mivart's book, and comparing each section with what I have said on the same head, I never before felt so strongly convinced of the general truth of the conclusions here arrived at, subject, of course, in so intricate a subject, to much partial error.
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 characters, often accompanied by a change of function, - for instance, the conversion of a swimbladder 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 S. 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, etc., 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; whilst 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 S. 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 his 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 S. 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 S. 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, whilst 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 S. America, though it is so luxuriant; whilst S. 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 so 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 co-adapted. 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, etc., - 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 man 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 an? 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 neutralise 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, whilst 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, whilst 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 whale-bone, 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 whale-bone 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 elypedta) 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 .14 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 inwards, instead of straight downwards. 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 Balanoptera, 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 those 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 learnt 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 50 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 the 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 ducks; but the lamellæ are not so numerous, nor so distinct from each other, nor do they project so much inwards; 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 27 in number on each side, and terminating upwards 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 the 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 Lacepède, 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 onwards, 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 Balaenoptera rostrata, gradations, which may be observed in still-existing cetaceans, lead us onwards 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.
The Pleuronectidæ, or flat-fish, are remarkable for their asymmetrical bodies. They rest on one side, - in the greater number of species on the left, but in some on the right side; and occasionally reversed adult specimens occur. The lower, or resting-surface, resembles at first sight the ventral surface of an ordinary fish: it is of a white colour, less developed in many ways than the upper side, with the lateral fins often of smaller size. But the eyes offer the most remarkable peculiarity; for they are both placed on the upper side of the head. During early youth, however, they stand opposite to each other, and the whole body is then symmetrical, with both sides equally coloured. Soon the eye proper to the lower side begins to glide slowly round the head to the upper side; but does not pass right through the skull, as was formerly thought to be the case. It is obvious that unless the lower eye did thus travel round, it could not be used by the fish whilst lying in its habitual position on one side. The lower eye would, also, have been liable to be abraded by the sandy bottom. That the Pleuronectidæ are admirably adapted by their flattened and asymmetrical structure for their habits of life, is manifest from several species, such as soles, flounders, etc., being extremely common. The chief advantages thus gained seem to be protection from their enemies, and facility for feeding on the ground. The different members, however, of the family present, as Schiödte remarks, a long series of forms exhibiting a gradual transition from Hippoglossus pinguis, which does not in any considerable degree alter the shape in which it leaves the ovum, to the soles, which are entirely thrown to one side.
Mr. Mivart has taken up this case, and remarks that a sudden spontaneous transformation in the position of the eyes is hardly conceivable, in which I quite agree with him. He then adds: If the transit was gradual, then how such transit of one eye a minute fraction of the journey towards the other side of the head could benefit the individual is, indeed, far from clear. It seems, even, that such an incipient transformation must rather have been injurious. But he might have found an answer to this objection in the excellent observations published in 1867 by Malm. The Pleuronectidæ whilst very young and still symmetrical, with their eyes standing on opposite sides of the head, cannot long retain a vertical position, owing to the excessive depth of their bodies, the small size of their lateral fins, and to their being destitute of a swimbladder. Hence soon growing tired, they fall to the bottom on one side. Whilst thus at rest they often twist, as Malm observed, the lower eye upwards, to see above them; and they do this so vigorously that the eye is pressed hard against the upper part of the orbit. The forehead between the eyes consequently becomes, as could be plainly seen, temporarily contracted in breadth. On one occasion Malm saw a young fish raise and depress the lower eye through an angular distance of about seventy degrees.
We should remember that the skull at this early age is cartilaginous and flexible, so that it readily yields to muscular action. It is also known with the higher animals, even after early youth, that the skull yields and is altered in shape, if the skin or muscles be permanently contracted through disease or some accident. With long-eared rabbits, if one ear lops forwards and downwards, its weight drags forward all the bones of the skull on the same side, of which I have given a figure. Malm states that the newly-hatched young of perches, salmon, and several other symmetrical fishes, have the habit of occasionally resting on one side at the bottom; and he has observed that they often then strain their lower eyes so as to look upwards; and their skulls are thus rendered rather crooked. These fishes, however, are soon able to hold themselves in a vertical position, and no permanent effect is thus produced. With the Pleuronectidæ, on the other hand, the older they grow the more habitually they rest on one side, owing to the increasing flatness of their bodies, and a permanent effect is thus produced on the form of the head, and on the position of the eyes. Judging from analogy, the tendency to distortion would no doubt be increased through the principle of inheritance. Schiödte believes, in opposition to some other naturalists, that the Pleuronectidæ are not quite symmetrical even in the embryo; and if this be so, we could understand how it is that certain species, whilst young, habitually fall over and rest on the left side, and other species on the right side. Malm adds, in confirmation of the above view, that the adult Trachypterus arcticus, which is not a member of the Pleuronectidæ, rests on its left side at the bottom, and swims diagonally through the water; and in this fish, the two sides of the head are said to be somewhat dissimilar. Our great authority on fishes, Dr. Günther, concludes his abstract of Malm's paper, by remarking that the author gives a very simple explanation of the abnormal condition of the pleuronectoids.
We thus see that the first stages of the transit of the eye from one side of the head to the other, which Mr. Mivart considers would be injurious, may be attributed to the habit, no doubt beneficial to the individual and to the species, of endeavouring to look upwards with both eyes, whilst resting on one side at the bottom. We may also attribute to the inherited effects of use the fact of the mouth in several kinds of flat-fish being bent towards the lower surface, with the jaw bones stronger and more effective on this, the eyeless side of the head, than on the other, for the sake, as Dr. Traquair supposes, of feeding with ease on the ground. Disuse, on the other hand, will account for the less developed condition of the whole inferior half of the body, including the lateral fins; though Yarrel thinks that the reduced size of these fins is advantageous to the fish, as there is so much less room for their action, than with the larger fins above. Perhaps the lesser number of teeth in the proportion of four to seven in the upper halves of the two jaws of the plaice, to twenty-five to thirty in the lower halves, may likewise be accounted for by disuse. From the colourless state of the ventral surface of most fishes and of many other animals, we may reasonably suppose that the absence of colour in flat-fish on the side, whether it be the right or left, which is undermost, is due to the exclusion of light. But it cannot be supposed that the peculiar speckled appearance of the upper side of the sole, so like the sandy bed of the sea, or the power in some species, as recently shown by Pouchet, of changing their colour in accordance with the surrounding surface, or the presence of bony tubercles on the upper side of the turbot, are due to the action of the light. Here natural selection has probably come into play, as well as in adapting the general shape of the body of these fishes, and many other peculiarities, to their habits of life. We should keep in mind, as I have before insisted, that the inherited effects of the increased use of parts, and perhaps of their disuse, will be strengthened by natural selection. For all spontaneous variations in the right direction will thus be preserved; as will those individuals which inherit in the highest degree the effects of the increased and beneficial use of any part. How much to attribute in each particular case to the effects of use, and how much to natural selection, it seems impossible to decide.
I may give another instance of a structure which apparently owes its origin exclusively to use or habit. The extremity of the tail in some American monkeys has been converted into a wonderfully perfect prehensile organ, and serves as a fifth hand. A reviewer who agrees with Mr. Mivart in every detail, remarks on this structure: It is impossible to believe that in any number of ages the first slight incipient tendency to grasp could preserve the lives of the individuals possessing it, or favour their chance of having and of rearing offspring. But there is no necessity for any such belief. Habit, and this almost implies that some benefit great or small is thus derived, would in all probability suffice for the work. Brehm saw the young of an African monkey (Cercopithecus) clinging to the under surface of their mother by their hands, and at the same time they hooked their little tails round that of their mother. Professor Henslow kept in confinement some harvest mice (Mus messorius) which do not possess a structurally prehensile tail; but he frequently observed that they curled their tails round the branches of a bush placed in the cage, and thus aided themselves in climbing. I have received an analogous account from Dr. Günther, who has seen a mouse thus suspend itself. If the harvest mouse had been more strictly arboreal, it would perhaps have had its tail rendered structurally prehensile, as is the case with some members of the same order. Why Cereopithecus, considering its habits whilst young, has not become thus provided, it would be difficult to say. It is, however, possible that the long tail of this monkey may be of more service to it as a balancing organ in making its prodigious leaps, than as a prehensile organ.
The mammary glands are common to the whole class of mammals, and are indispensable for their existence; they must, therefore, have been developed at an extremely remote period, and we can know nothing positively about their manner of development. Mr. Mivart asks: Is it conceivable that the young of any animal was ever saved from destruction by accidentally sucking a drop of scarcely nutritious fluid from an accidentally hypertrophied cutaneous gland of its mother? And even if one was so, what chance was there of the perpetuation of such a variation? But the case is not here put fairly. It is admitted by most evolutionists that mammals are descended from a marsupial form; and if so, the mammary glands will have been at first developed within the marsupial sack. In the case of the fish (Hippocampus) the eggs are hatched, and the young are reared for a time, within a sack of this nature; and an American naturalist, Mr. Lockwood, believes from what he has seen of the development of the young, that they are nourished by a secretion from the cutaneous glands of the sack. Now with the early progenitors of mammals, almost before they deserved to be thus designated, is it not at least possible that the young might have been similarly nourished? And in this case, the individuals which secreted a fluid, in some degree or manner the most nutritious, so as to partake of the nature of milk, would in the long run have reared a larger number of well-nourished offspring, than would the individuals which secreted a poorer fluid; and thus the cutaneous glands, which are the homologues of the mammary glands, would have been improved or rendered more effective. It accords with the widely extended principle of specialisation, that the glands over a certain space of the sack should have become more highly developed than the remainder; and they would then have formed a breast, but at first without a nipple as we see in the Ornithorhynchus, at the base of the mammalian series. Through what agency the glands over a certain space became more highly specialised than the others, I will not pretend to decide, whether in part through compensation of growth, the effects of use, or of natural selection.
The development of the mammary glands would have been of no service, and could not have been effected through natural selection, unless the young at the same time were able to partake of the secretion. There is no greater difficulty in understanding how young mammals have instinctively learnt to suck the breast, than in understanding how unhatched chickens have learnt to break the egg-shell by tapping against it with their specially adapted beaks; or how a few hours after leaving the shell they have learnt to pick up grains of food. In such cases the most probable solution seems to be, that the habit was at first acquired by practice at a more advanced age, and afterwards transmitted to the offspring at an earlier age. But the young kangaroo is said not to suck, only to cling to the nipple of its mother, who has the power of injecting milk into the mouth of her helpless, half-formed offspring. On this head, Mr. Mivart remarks: Did no special provision exist, the young one must infallibly be choked by the intrusion of the milk into the windpipe. But there is a special provision. The larynx is so elongated that it rises up into the posterior end of the nasal passage, and is thus enabled to give free entrance to the air for the lungs, while the milk passes harmlessly on each side of this elongated larynx, and so safely attains the gullet behind it. Mr. Mivart then asks how did natural selection remove in the adult kangaroo (and in most other mammals, on the assumption that they are descended from a marsupial form), this at least perfectly innocent and harmless structure? It may be suggested in answer that the voice, which is certainly of high importance to many animals, could hardly have been used with full force as long as the larynx entered the nasal passage; and Professor Flower has suggested to me that this structure would have greatly interfered with an animal swallowing solid food.
We will now turn for a short space to the lower divisions of the animal kingdom. The Echinodermata (star-fishes, sea-urchins, etc.) are furnished with remarkable organs, called pedicellariæ, which consist, when well developed, of a tridactyle forceps - that is, of one formed of three serrated arms, neatly fitting together and placed on the summit of a flexible stem, moved by muscles. These forceps can firmly seize hold of any object; and Alexander Agassiz has seen an Echinus or sea-urchin rapidly passing particles of excrement from forceps to forceps down certain lines of its body, in order that its shell should not be fouled. But there is no doubt that besides removing dirt of all kinds, they subserve other functions; and one of these apparently is defence.
With respect to these organs, Mr. Mivart, as on so many previous occasions, asks: What would be the utility of the first rudimentary beginnings of such structures, and how could such incipient buddings have ever preserved the life of a single Echinus? He adds, Not even the sudden development of the snapping action could have been beneficial without the freely moveable stalk, nor could the latter have been efficient without the snapping jaws, yet no minute merely indefinite variations could simultaneously evolve these complex co-ordinations of structure; to deny this seems to do no less than to affirm a startling paradox. Paradoxical as this may appear to Mr. Mivart, tridactyle forcepses, immovably fixed at the base, but capable of a snapping action, certainly exist on some starfishes; and this is intelligible if they serve, at least in part, as a means of defence. Mr. Agassiz, to whose great kindness I am indebted for much information on the subject, informs me that there are other star-fishes, in which one of the three arms of the forceps is reduced to a support for the other two; and again, other genera in which the third arm is completely lost. In Echinoneus, the shell is described by M. Perrier as bearing two kinds of pedicellariæ, one resembling those of Echinus, and the other those of Spatangus; and such cases are always interesting as affording the means of apparently sudden transitions, through the abortion of one of the two states of an organ.
With respect to the steps by which these curious organs have been evolved, Mr. Agassiz infers from his own researches and those of Müller, that both in star-fishes and sea-urchins the pedicellariæ must undoubtedly be looked at as modified spines. This may be inferred from their manner of development in the individual, as well as from a long and perfect series of gradations in different species and genera, from simple granules to ordinary spines, to perfect tridactyle pedicellariæ. The gradations extend even to the manner in which ordinary spines and pedicellariæ with their supporting calcareous rods are articulated to the shell. In certain genera of star-fishes, the very combinations needed to show that the pedicellariæ are only modified branching spines may be found. Thus we have fixed spines, with three equidistant, serrated, moveable branches, articulated to near their bases; and higher up, on the same spine, three other moveable branches. Now when the latter arise from the summit of a spine they form in fact a rude tridactyle pedicellaria, and such may be seen on the same spine together with the three lower branches. In this case the identity in nature between the arms of the pedicellariæ and the moveable branches of a spine, is unmistakable. It is generally admitted that the ordinary spines serve as a protection; and if so, there can be no reason to doubt that those furnished with serrated and moveable branches likewise serve for the same purpose; and they would thus serve still more effectively as soon as by meeting together they acted as a prehensile or snapping apparatus. Thus every gradation, from an ordinary fixed spine to a fixed pedicellaria, would be of service.
In certain genera of star-fishes these organs, instead of being fixed or borne on an immoveable support, are placed on the summit of a flexible and muscular, though short, stem; and in this case they probably subserve some additional function besides defence. In the sea-urchins the steps can be followed by which a fixed spine becomes articulated to the shell, and is thus rendered moveable. I wish I had space here to give a fuller abstract of Mr. Agassiz's interesting observations on the development of the pedicellariæ. All possible gradations, as he adds, may likewise be found between the pedicellariae of the star-fishes and the hooks of the ophiurians, another group of Echinodermata; and again between the pedicellariae of sea-urchins and the anchors of the Holothuriæ, also belonging to the same great class.
Certain compound animals, or zoöphytes as they have been termed, namely the Polyzoa, are provided with curious organs called avicularia. These differ much in structure in the different species. In their most perfect condition, they curiously resemble the head and beak of a vulture in miniature, seated on a neck and capable of movement, as is likewise the lower jaw or mandible. In one species observed by me all the avicularia on the same branch often moved simultaneously backwards and forwards, with the lower jaw widely open, through an angle of about 90°, in the course of five seconds; and their movement caused the whole polyzoary to tremble. When the jaws are touched with a needle they seize it so firmly that the branch can thus be shaken.
Mr. Mivart adduces this case, chiefly on account of the supposed difficulty of organs, namely the avicularia of the Polyzoa and the pedicellariæ of the Echinodermata, which he considers as essentially similar, having been developed through natural selection in widely distinct divisions of the animal kingdom. But, as far as structure is concerned, I can see no similarity between tridactyle pedicellariae and avicularia. The latter resemble somewhat more closely the chelæ or pincers of crustaceans; and Mr. Mivart might have adduced with equal appropriateness this resemblance as a special difficulty; or even their resemblance to the head and beak of a bird. The avicularia are believed by Mr. Busk, Dr. Smitt, and Dr. Nitsche - naturalists who have carefully studied this group - to be homologous with the zooids and their cells which compose the zoöphyte; the moveable lip or lid of the cell corresponding with the lower and moveable mandible of the avicularium. Mr. Busk, however, does not know of any gradations now existing between a zooid and an avicularium. It is therefore impossible to conjecture by what serviceable gradations the one could have been converted into the other: but it by no means follows from this that such gradations have not existed.
As the chelæ of crustaceans resemble in some degree the avicularia of Polyzoa, both serving as pincers, it may be worth while to show that with the former a long series of serviceable gradations still exists. In the first and simplest stage, the terminal segment of a limb shuts down either on the square summit of the broad penultimate segment, or against one whole side; and is thus enabled to catch hold of an object; but the limb still serves as an organ of locomotion. We next find one corner of the broad penultimate segment slightly prominent, sometimes furnished with irregular teeth; and against these the terminal segment shuts down. By an increase in the size of this projection, with its shape, as well as that of the terminal segment, slightly modified and improved, the pincers are rendered more and more perfect, until we have at last an instrument as efficient as the chelæ of a lobster; and all these gradations can be actually traced.
Besides the avicularia, the Polyzoa possess curious organs called vibracula. These generally consist of long bristles, capable of movement and easily excited. In one species examined by me the vibracula were slightly curved and serrated along the outer margin; and all of them on the same polyzoary often moved simultaneously; so that, acting like long oars, they swept a branch rapidly across the object-glass of my microscope. When a branch was placed on its face, the vibracula became entangled, and they made violent efforts to free themselves. They are supposed to serve as a defence, and may be seen, as Mr. Busk remarks, to sweep slowly and carefully over the surface of the polyzoary, removing what might be noxious to the delicate inhabitants of the cells when their tentacula are protruded. The avicularia, like the vibracula, probably serve for defence, but they also catch and kill small living animals, which it is believed are afterwards swept by the currents within reach of the tentacula of the zooids. Some species are provided with avicularia and vibracula; some with avicularia alone, and a few with vibracula alone.
It is not easy to imagine two objects more widely different in appearance than a bristle or vibraculum, and an avicularium like the head of a bird; yet they are almost certainly homologous and have been developed from the same common source, namely a zooid with its cell. Hence we can understand how it is that these organs graduate in some cases, as I am informed by Mr. Busk, into each other. Thus with the avicularia of several species of Lepralia, the moveable mandible is so much produced and is so like a bristle, that the presence of the upper or fixed beak alone serves to determine even its avicularian nature. The vibracula may have been directly developed from the lips of the cells, without having passed through the avicularian stage; but it seems more probable that they have passed through this stage, as during the early stages of the transformation, the other parts of the cell with the included zooid could hardly have disappeared at once. In many cases the vibracula have a grooved support at the base, which seems to represent the fixed beak; though this support in some species is quite absent. This view of the development of the vibracula, if trustworthy, is interesting; for supposing that all the species provided with avicularia had become extinct, no one with the most vivid imagination would ever have thought that the vibracula had originally existed as part of an organ, resembling a bird's head or an irregular box or hood. It is interesting to see two such widely different organs developed from a common origin; and as the moveable lip of the cell serves as a protection to the zooid, there is no difficulty in believing that all the gradations, by which the lip became converted first into the lower mandible of an avicularium and then into an elongated bristle, likewise served as a protection in different ways and under different circumstances.
In the vegetable kingdom Mr. Mivart only alludes to two cases, namely the structure of the flowers of orchids, and the movements of climbing plants. With respect to the former, he says, The explanation of their origin is deemed thoroughly unsatisfactory - utterly insufficient to explain the incipient, infinitesimal beginnings of structures which are of utility only when they are considerably developed. As I have fully treated this subject in another work, I will here give only a few details on one alone of the most striking peculiarities of the flowers of orchids, namely their pollinia. A pollinium when highly developed consists of a mass of pollen-grains, affixed to an elastic footstalk or caudicle, and this to a little mass of extremely viscid matter. The pollinia are by this means transported by insects from one flower to the stigma of another. In some orchids there is no caudicle to the pollen-masses, and the grains are merely tied together by fine threads; but as these are not confined to orchids, they need not here be considered; yet I may mention that at the base of the orchidaceous series, in Cypripedium, we can see how the threads were probably first developed. In other orchids the threads cohere at one end of the pollen-masses; and this forms the first or nascent trace of a caudicle. That this is the origin of the caudicle, even when of considerable length and highly developed, we have good evidence in the aborted pollen-grains which can sometimes be detected embedded within the central and solid parts.
With respect to the second chief peculiarity, namely the little mass of viscid matter attached to the end of the caudicle, a long series of gradations can be specified, each of plain service to the plant. In most flowers belonging to other orders the stigma secretes a little viscid matter. Now in certain orchids similar viscid matter is secreted, but in much larger quantities by one alone of the three stigmas; and this stigma, perhaps in consequence of the copious secretion, is rendered sterile. When an insect visits a flower of this kind, it rubs off some of the viscid matter and thus at the same time drags away some of the pollen-grains. From this simple condition, which differs but little from that of a multitude of common flowers, there are endless gradations, - to species in which the pollen-mass terminates in a very short, free caudicle, - to others in which the caudicle becomes firmly attached to the viscid matter, with the sterile stigma itself much modified. In this latter case we have a pollinium in its most highly developed and perfect condition. He who will carefully examine the flowers of orchids for himself will not deny the existence of the above series of gradations - from a mass of pollen-grains merely tied together by threads, with the stigma differing but little from that of an ordinary flower, to a highly complex pollinium, admirably adapted for transportal by insects; nor will he deny that all the gradations in the several species are admirably adapted in relation to the general structure of each flower for its fertilisation by different insects. In this, and in almost every other case, the enquiry may be pushed further backwards; and it may be asked how did the stigma of an ordinary flower become viscid, but as we do not know the full history of any one group of beings, it is as useless to ask, as it is hopeless to attempt answering, such questions.
We will now turn to climbing plants. These can be arranged in a long series, from those which simply twine round a support, to those which I have called leaf-climbers, and to those provided with tendrils. In these two latter classes the stems have generally, but not always, lost the power of twining, though they retain the power of revolving, which the tendrils likewise possess. The gradations from leaf-climbers to tendril-bearers are wonderfully close, and certain plants may be indifferently placed in either class. But in ascending the series from simple twiners to leaf-climbers, an important quality is added, namely sensitiveness to a touch, by which means the foot-stalks of the leaves or flowers, or these modified and converted into tendrils, are excited to bend round and clasp the touching object. He who will read my memoir on these plants will, I think, admit that all the many gradations in function and structure between simple twiners and tendril-bearers are in each case beneficial in a high degree to the species. For instance, it is clearly a great advantage to a twining plant to become a leaf-climber; and it is probable that every twiner which possessed leaves with long foot-stalks would have been developed into a leaf-climber if the footstalks had possessed in any slight degree the requisite sensitiveness to a touch.
As twining is the simplest means of ascending a support, and forms the basis of our series, it may naturally be asked how did plants acquire this power in an incipient degree, afterwards to be improved and increased through natural selection. The power of twining depends, firstly, on the stems whilst young being extremely flexible (but this is a character common to many plants which are not climbers); and, secondly, on their continually bending to all points of the compass, one after the other in succession, in the same order. By this movement the stems are inclined to all sides, and are made to move round and round. As soon as the lower part of a stem strikes against any object and is stopped, the upper part still goes on bending and revolving, and thus necessarily twines round and up the support. The revolving movement ceases after the early growth of each shoot. As in many widely separated families of plants, single species and single genera possess the power of revolving, and have thus become twiners, they must have independently acquired it, and cannot have inherited it from a common progenitor. Hence I was led to predict that some slight tendency to a movement of this kind would be found to be far from uncommon with plants which did not climb; and that this had afforded the basis for natural selection to work on and improve. When I made this prediction, I knew of only one imperfect case, namely, of the young flower-peduncles of a Maurandia which revolved slightly and irregularly, like the stems of twining plants, but without making any use of this habit. Soon afterwards Fritz Müller discovered that the young stems of an Alisima and of a Linum, - plants which do not climb and are widely separated in the natural system, - revolved plainly, though irregularly; and he states that he has reason to suspect that this occurs with some other plants. These slight movements appear to be of no service to the plants in question; anyhow, they are not of the least use in the way of climbing, which is the point that concerns us. Nevertheless we can see that if the stems of these plants had been flexible, and if under the conditions to which they are exposed it had profited them to ascend to a height, then the habit of slightly and irregularly revolving might have been increased and utilised through natural selection, until they had become converted into well-developed twining species.
With respect to the sensitiveness of the footstalks of the leaves and flowers, and of tendrils, nearly the same remarks are applicable as in the case of the revolving movements of twining plants. As a vast number of species, belonging to widely distinct groups, are endowed with this kind of sensitiveness, it ought to be found in a nascent condition in many plants which have not become climbers. This is the case: I observed that the young flower-peduncles of the above Maurandia curved themselves a little toward the side which was touched. Morren found in several species of Oxalis that the leaves and their foot-stalks moved, especially after exposure to a hot sun, when they were gently and repeatedly touched, or when the plant was shaken. I repeated these observations on some other species of Oxalis with the same result; in some of them the movement was distinct, but was best seen in the young leaves; in others it was extremely slight. It is a more important fact that according to the high authority of Hofmeister, the young shoots and leaves of all plants move after being shaken; and with climbing plants it is, as we know, only during the early stages of growth that the foot-stalks and tendrils are sensitive.
It is scarcely possible that the above slight movements, due to a touch or shake, in the young and growing organs of plants, can be of any functional importance to them. But plants possess, in obedience to various stimuli, powers of movement, which are of manifest importance to them; for instance, towards and more rarely from the light, - in opposition to, and more rarely in the direction of, the attraction of gravity. When the nerves and muscles of an animal are excited by galvanism or by the absorption of strychnine, the consequent movements may be called an incidental result, for the nerves and muscles have not been rendered specially sensitive to these stimuli. So with plants it appears that, from having the power of movement in obedience to certain stimuli, they are excited in an incidental manner by a touch, or by being shaken. Hence there is no great difficulty in admitting that in the case of leaf-climbers and tendril-bearers, it is this tendency which has been taken advantage of and increased through natural selection. It is, however, probable, from reasons which I have assigned in my memoir, that this will have occurred only with plants which had already acquired the power of revolving, and had thus become twiners.
I have already endeavoured to explain how plants became twiners, namely, by the increase of a tendency to slight and irregular revolving movements, which were at first of no use to them; this movement, as well as that due to a touch or shake, being the incidental result of the power of moving, gained for other and beneficial purposes. Whether, during the gradual development of climbing plants, natural selection has been aided by the inherited effects of use, I will not pretend to decide; but we know that certain periodical movements, for instance the so-called sleep of plants, are governed by habit.
I have now considered enough, perhaps more than enough, of the cases, selected with care by a skilful naturalist, to prove that natural selection is incompetent to account for the incipient stages of useful structures; and I have shown, as I hope, that there is no great difficulty on this head. A good opportunity has thus been afforded for enlarging a little on gradations of structure, often associated with changed functions, - an important subject which was not treated at sufficient length in the former editions of this work. I will now briefly recapitulate the foregoing cases.
With the giraffe, the continued preservation of the individuals of some extinct high-reaching ruminant, which had the longest necks, legs, etc., and could browse a little above the average height, and the continued destruction of those which could not browse so high, would have sufficed for the production of this remarkable quadruped; but the prolonged use of all the parts together with inheritance will have aided in an important manner in their co-ordination. With the many insects which imitate various objects, there is no improbability in the belief that an accidental resemblance to some common object was in each case the foundation for the work of natural selection, since perfected through the occasional preservation of slight variations which ma de the resemblance at all closer; and this will have been carried on as long as the insect continued to vary, and as long as a more and more perfect resemblance led to its escape from sharp-sighted enemies. In certain species of whales there is a tendency to the formation of irregular little points of horn on the palate; and it seems to be quite within the scope of natural selection to preserve all favourable variations, until the points were converted first into lamellated knobs or teeth, like those on the beak of a goose, - then into short lamellæ, like those of the domestic ducks, - and then into lamellæ, as perfect as those of the shoveller-duck, - and finally into the gigantic plates of baleen, as in the mouth of the Greenland whale. In the family of the ducks, the lamellæ are first used as teeth, then partly as teeth, and partly as a sifting apparatus, and at last almost exclusively for this latter purpose.
With such structures as the above lamellæ of horn or whalebone, habit or use can have done little or nothing, as far as we can judge, towards their development. On the other hand, the transportal of the lower eye of a flat-fish to the upper side of the head, and the formation of a prehensile tail, may be attributed almost wholly to continued use, together with inheritance. With respect to the mammæ of the higher animals, the most probable conjecture is that primordially the cutaneous glands over the whole surface of a marsupial sack secreted a nutritious fluid; and that these glands were improved in function through natural selection, and concentrated into a confined area, in which case they would have formed a mamma. There is no more difficulty in understanding how the branched spines of some ancient echinoderm, which served as a defence, became developed through natural selection into tridactyle pedicellariæ, than in understanding the development of the pincers of crustaceans, through slight, serviceable modifications in the ultimate and penultimate segments of a limb, which was at first used solely for locomotion. In the avicularia and vibracula of the Polyzoa we have organs widely different in appearance developed from the same source; and with the vibracula we can understand how the successive gradations might have been of service. With the pollinia of orchids, the threads which originally served to tie together the pollen-grains, can be traced cohering into caudicles; and the steps can likewise be followed by which viscid matter, such as that secreted by the stigmas of ordinary flowers, and still subserving nearly but not quite the same purpose, became attached to the free ends of the caudicles; - all these gradations being of modest benefit to the plants in question. With respect to climbing plants, I need not repeat what has been so lately said.
It has often been asked, if natural selection be so potent, why has not this or that structure been gained by certain species, to which it would apparently have been advantageous? But it is unreasonable to expect a precise answer to such questions, considering our ignorance of the past history of each species, and of the conditions which at the present day determine its numbers and range. In most cases only general reasons, but in some few cases special reasons, can be assigned. Thus to adapt a species to new habits of life, many co-ordinated modifications are almost indispensable, and it may often have happened that the requisite parts did not vary in the right manner or to the right degree. Many species must have been prevented from increasing in numbers through destructive agencies, which stood in no relation to certain structures, which we imagine would have been gained through natural selection from appearing to us advantageous to the species. In this case, as the struggle for life did not depend on such structures, they could not have been acquired through natural selection. In many cases complex and long-enduring conditions, often of a peculiar nature, are necessary for the development of a structure; and the requisite conditions may seldom have concurred. The belief that any given structure, which we think, often erroneously, would have been beneficial to a species, would have been gained under all circumstances through natural selection, is opposed to what we can understand of its manner of action. Mr. Mivart does not deny that natural selection has effected something; but he considers it as demonstrably insufficient to account for the phenomena which I explain by its agency. His chief arguments have now been considered, and the others will hereafter be considered. They seem to me to partake little of the character of demonstration, and to have little weight in comparison with those in favour of the power of natural selection, aided by the other agencies often specified. I am bound to add, that some of the facts and arguments here used by me, have been advanced for the same purpose in an able article lately published in the Medico-Chirurgical Review.
At the present day almost all naturalists admit evolution under some form. Mr. Mivart believes that species change through an internal force or tendency, about which it is not pretended that anything is known. That species have a capacity for change will be admitted by all evolutionists; but there is no need, as it seems to me, to invoke any internal force beyond the tendency to ordinary variability, which through the aid of selection by man has given rise to many well-adapted domestic races, and which through the aid of natural selection would equally well give rise by graduated steps to natural races or species. The final result will generally have been, as already explained, an advance, but in some few cases a retrogression, in organisation.
Mr. Mivart is further inclined to believe, and some naturalists agree with him, that new species manifest themselves with suddenness and by modifications appearing at once. For instance, he supposes that the differences between the extinct three-toed Hipparion and the horse arose suddenly. He thinks it difficult to believe that the wing of a bird was developed in any other way than by a comparatively sudden modification of a marked and important kind; and apparently he would extend the same view to the wings of bats and pterodactyles. This conclusion, which implies great breaks or discontinuity in the series, appears to me improbable in the highest degree.
Every one who believes in slow and gradual evolution, will of course admit that specific changes may have been as abrupt and as great as any single variation which we meet with under nature, or even under domestication. But as species are more variable when domesticated or cultivated than under their natural conditions, it is not probable that such great and abrupt variations have often occurred under nature, as are known occasionally to arise under domestication. Of these latter variations several may be attributed to reversion; and the characters which thus reappear were, it is probable, in many cases at first gained in a gradual manner. A still greater number must be called monstrosities, such as six-fingered men, porcupine men, Ancon sheep, Niata cattle, etc.; and as they are widely different in character from natural species, they throw very little light on our subject. Excluding such cases of abrupt variations, the few which remain would at best constitute, if found in a state of nature, doubtful species, closely related to their parental types.
My reasons for doubting whether natural species have changed as abruptly as have occasionally domestic races, and for entirely disbelieving that they have changed in the wonderful manner indicated by Mr. Mivart, are as follows. According to our experience, abrupt and strongly marked variations occur in our domesticated productions, singly and at rather long intervals of time. If such occurred under nature, they would be liable, as formerly explained, to be lost by accidental causes of destruction and by subsequent inter-crossing; and so it is known to be under domestication, unless abrupt variations of this kind are specially preserved and separated by the care of man. Hence in order that a new species should suddenly appear in the manner supposed by Mr. Mivart, it is almost necessary to believe, in opposition to all analogy, that several wonderfully changed individuals appeared simultaneously within the same district. This difficulty, as in the case of unconscious selection by man, is avoided on the theory of gradual evolution, through the preservation of a large number of individuals, which varied more or less in any favourable direction, and of the destruction of a large number which varied in an opposite manner.
That many species have been evolved in an extremely gradual manner, there can hardly be a doubt. The species and even the genera of many large natural families are so closely allied together, that it is difficult to distinguish not a few of them. On every continent in proceeding from north to south, from lowland to upland, etc., we meet with a host of closely related or representative species; as we likewise do on certain distinct continents, which we have reason to believe were formerly connected. But in making these and the following remarks, I am compelled to allude to subjects hereafter to be discussed. Look at the many outlying islands round a continent, and see how many of their inhabitants can be raised only to the rank of doubtful species. So it is if we look to past times, and compare the species which have just passed away with those still living within the same areas; or if we compare the fossil species embedded in the sub-stages of the same geological formation. It is indeed manifest that multitudes of species are related in the closest manner to other species that still exist, or have lately existed; and it will hardly be maintained that such species have been developed in an abrupt or sudden manner. Nor should it be forgotten, when we look to the special parts of allied species, instead of to distinct species, that numerous and wonderfully fine gradations can be traced, connecting together widely different structures.
Many large groups of facts are intelligible only on the principle that species have been evolved by very small steps: for instance, the fact that the species included in the larger genera are more closely related to each other, and present a greater number of varieties than do the species in the smaller genera. The former are also grouped in little clusters, like varieties round species, and they present other analogies with varieties, as was shown in our second chapter. On this same principle we can understand how it is that specific characters are more variable than generic characters; and how the parts which are developed in an extraordinary degree or manner are more variable than other parts of the same species. Many analogous facts, all pointing in the same direction, could be added.
Although very many species have almost certainly been produced by steps not greater than those separating fine varieties; yet it may be maintained that some have been developed in a different and abrupt manner. Such an admission, however, ought not to be made without strong evidence being assigned. The vague and in some respects false analogies, as they have been shown to be by Mr. Chauncey Wright, which have been advanced in favour of this view, such as the sudden crystallisation of inorganic substances, or the falling of a facetted spheroid from one facet to another, hardly deserve consideration. One class of facts, however, namely, the sudden appearance of new and distinct forms of life in our geological formations, supports at first sight the belief in abrupt development. But the value of this evidence depends entirely on the perfection of the geological record, in relation to periods remote in the history of the world. If the record is as fragmentary as many geologists strenuously assert, there is nothing strange in new forms appearing as if suddenly developed.
Unless we admit transformations as prodigious as those advocated by Mr. Mivart, such as the sudden development of the wings of birds or bats, or the sudden conversion of a Hipparion into a horse, hardly any light is thrown by the belief in abrupt modifications on the deficiency of connecting links in our geological formations. But against the belief in such abrupt changes, embryology enters a strong protest. It is notorious that the wings of birds and bats, and the legs of horses or other quadrupeds, are undistinguishable at an early embryonic period, and that they become differentiated by insensibly fine steps. Embryological resemblances of all kinds can be accounted for, as we shall hereafter see, by the progenitors of our existing species having varied after early youth, and having transmitted their newly acquired characters to their offspring, at a corresponding age. The embryo is thus left almost unaffected, and serves as a record of the past condition of the species. Hence it is that existing species during the early stages of their development so often resemble ancient and extinct forms belonging to the same class. On this view of the meaning of embryological resemblances, and indeed on any view, it is incredible that an animal should have undergone such momentous and abrupt transformations, as those above indicated; and yet should not bear even a trace in its embryonic condition of any sudden modification; every detail in its structure being developed by insensibly fine steps.
He who believes that some ancient form was transformed suddenly through an internal force or tendency into, for instance, one furnished with wings, will be almost compelled to assume, in opposition to all analogy, that many individuals varied simultaneously. It cannot be denied that such abrupt and great changes of structure are widely different from those which most species apparently have undergone. He will further be compelled to believe that many structures beautifully adapted to all the other parts of the same creature and to the surrounding conditions, have been suddenly produced; and of such complex and wonderful co-adaptations, he will not be able to assign a shadow of an explanation. He will be forced to admit that these great and sudden transformations have left no trace of their action on the embryo. To admit all this is, as it seems to me, to enter into the realms of miracle, and to leave those of Science.
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