The Origin of Species by Means of Natural Selection

It may be worth notice that when Wagner's remarkable discovery was first announced, I was asked how was it possible to account for the larvae of this fly having acquired the power of a sexual reproduction. As long as the case remained unique no answer could be given. But already Grimm has shown that another fly, a Chironomus, reproduces itself in nearly the same manner, and he believes that this occurs frequently in the order. It is the pupa, and not the larva, of the Chironomus which has this power; and Grimm further shows that this case, to a certain extent, "unites that of the Cecidomyia with the parthenogenesis of the Coccidae;" the term parthenogenesis implying that the mature females of the Coccidae are capable of producing fertile eggs without the concourse of the male. Certain animals belonging to several classes are now known to have the power of ordinary reproduction at an unusually early age; and we have only to accelerate parthenogenetic reproduction by gradual steps to an earlier and earlier age – Chironomus showing us an almost exactly intermediate stage, viz., that of the pupa – and we can perhaps account for the marvellous case of the Cecidomyia.

It has already been stated that various parts in the same individual, which are exactly alike during an early embryonic period, become widely different and serve for widely different purposes in the adult state. So again it has been shown that generally the embryos of the most distinct species belonging to the same class are closely similar, but become, when fully developed, widely dissimilar. A better proof of this latter fact cannot be given than the statement by Von Baer that "the embryos of mammalia, of birds, lizards and snakes, probably also of chelonia, are in the earliest states exceedingly like one another, both as a whole and in the mode of development of their parts; so much so, in fact, that we can often distinguish the embryos only by their size. In my possession are two little embryos in spirit, whose names I have omitted to attach, and at present I am quite unable to say to what class they belong. They may be lizards or small birds, or very young mammalia, so complete is the similarity in the mode of formation of the head and trunk in these animals. The extremities, however, are still absent in these embryos. But even if they had existed in the earliest stage of their development we should learn nothing, for the feet of lizards and mammals, the wings and feet of birds, no less than the hands and feet of man, all arise from the same fundamental form." The larvae of most crustaceans, at corresponding stages of development, closely resemble each other, however different the adults may become; and so it is with very many other animals. A trace of the law of embryonic resemblance occasionally lasts till a rather late age: thus birds of the same genus, and of allied genera, often resemble each other in their immature plumage; as we see in the spotted feathers in the young of the thrush group. In the cat tribe, most of the species when adult are striped or spotted in lines; and stripes or spots can be plainly distinguished in the whelp of the lion and the puma. We occasionally, though rarely, see something of the same kind in plants; thus the first leaves of the ulex or furze, and the first leaves of the phyllodineous acacias, are pinnate or divided like the ordinary leaves of the leguminosae.

The points of structure, in which the embryos of widely different animals within the same class resemble each other, often have no direct relation to their conditions of existence. We cannot, for instance, suppose that in the embryos of the vertebrata the peculiar loop-like courses of the arteries near the branchial slits are related to similar conditions – in the young mammal which is nourished in the womb of its mother, in the egg of the bird which is hatched in a nest, and in the spawn of a frog under water. We have no more reason to believe in such a relation than we have to believe that the similar bones in the hand of a man, wing of a bat, and fin of a porpoise, are related to similar conditions of life. No one supposes that the stripes on the whelp of a lion, or the spots on the young blackbird, are of any use to these animals.

The case, however, is different when an animal, during any part of its embryonic career, is active, and has to provide for itself. The period of activity may come on earlier or later in life; but whenever it comes on, the adaptation of the larva to its conditions of life is just as perfect and as beautiful as in the adult animal. In how important a manner this has acted, has recently been well shown by Sir J. Lubbock in his remarks on the close similarity of the larvae of some insects belonging to very different orders, and on the dissimilarity of the larvae of other insects within the same order, according to their habits of life. Owing to such adaptations the similarity of the larvae of allied animals is sometimes greatly obscured; especially when there is a division of labour during the different stages of development, as when the same larva has during one stage to search for food, and during another stage has to search for a place of attachment. Cases can even be given of the larvae of allied species, or groups of species, differing more from each other than do the adults. In most cases, however, the larvae, though active, still obey, more or less closely, the law of common embryonic resemblance. Cirripedes afford a good instance of this: even the illustrious Cuvier did not perceive that a barnacle was a crustacean: but a glance at the larva shows this in an unmistakable manner. So again the two main divisions of cirripedes, the pedunculated and sessile, though differing widely in external appearance, have larvae in all their stages barely distinguishable.

The embryo in the course of development generally rises in organisation. I use this expression, though I am aware that it is hardly possible to define clearly what is meant by organisation being higher or lower. But no one probably will dispute that the butterfly is higher than the caterpillar. In some cases, however, the mature animal must be considered as lower in the scale than the larva, as with certain parasitic crustaceans. To refer once again to cirripedes: the larvae in the first stage have three pairs of locomotive organs, a simple single eye, and a probosciformed mouth, with which they feed largely, for they increase much in size. In the second stage, answering to the chrysalis stage of butterflies, they have six pairs of beautifully constructed natatory legs, a pair of magnificent compound eyes, and extremely complex antennae; but they have a closed and imperfect mouth, and cannot feed: their function at this stage is, to search out by their well-developed organs of sense, and to reach by their active powers of swimming, a proper place on which to become attached and to undergo their final metamorphosis. When this is completed they are fixed for life: their legs are now converted into prehensile organs; they again obtain a well-constructed mouth; but they have no antennae, and their two eyes are now reconverted into a minute, single, simple eye-spot. In this last and complete state, cirripedes may be considered as either more highly or more lowly organised than they were in the larval condition. But in some genera the larvae become developed into hermaphrodites having the ordinary structure, or into what I have called complemental males; and in the latter the development has assuredly been retrograde; for the male is a mere sack, which lives for a short time and is destitute of mouth, stomach, and every other organ of importance, excepting those for reproduction.

We are so much accustomed to see a difference in structure between the embryo and the adult, that we are tempted to look at this difference as in some necessary manner contingent on growth. But there is no reason why, for instance, the wing of a bat, or the fin of a porpoise, should not have been sketched out with all their parts in proper proportion, as soon as any part became visible. In some whole groups of animals and in certain members of other groups this is the case, and the embryo does not at any period differ widely from the adult: thus Owen has remarked in regard to cuttle-fish, "there is no metamorphosis; the cephalopodic character is manifested long before the parts of the embryo are completed." Land-shells and fresh-water crustaceans are born having their proper forms, while the marine members of the same two great classes pass through considerable and often great changes during their development. Spiders, again, barely undergo any metamorphosis. The larvae of most insects pass through a worm-like stage, whether they are active and adapted to diversified habits, or are inactive from being placed in the midst of proper nutriment, or from being fed by their parents; but in some few cases, as in that of Aphis, if we look to the admirable drawings of the development of this insect, by Professor Huxley, we see hardly any trace of the vermiform stage.

Sometimes it is only the earlier developmental stages which fail. Thus, Fritz Muller has made the remarkable discovery that certain shrimp-like crustaceans (allied to Penoeus) first appear under the simple nauplius-form, and after passing through two or more zoea-stages, and then through the mysis-stage, finally acquire their mature structure: now in the whole great malacostracan order, to which these crustaceans belong, no other member is as yet known to be first developed under the nauplius-form, though many appear as zoeas; nevertheless Muller assigns reasons for his belief, that if there had been no suppression of development, all these crustaceans would have appeared as nauplii.

How, then, can we explain these several facts in embryology – namely, the very general, though not universal, difference in structure between the embryo and the adult; the various parts in the same individual embryo, which ultimately become very unlike, and serve for diverse purposes, being at an early period of growth alike; the common, but not invariable, resemblance between the embryos or larvae of the most distinct species in the same class; the embryo often retaining, while within the egg or womb, structures which are of no service to it, either at that or at a later period of life; on the other hand, larvae which have to provide for their own wants, being perfectly adapted to the surrounding conditions; and lastly, the fact of certain larvae standing higher in the scale of organisation than the mature animal into which they are developed? I believe that all these facts can be explained as follows.

It is commonly assumed, perhaps from monstrosities affecting the embryo at a very early period, that slight variations or individual differences necessarily appear at an equally early period. We have little evidence on this head, but what we have certainly points the other way; for it is notorious that breeders of cattle, horses and various fancy animals, cannot positively tell, until some time after birth, what will be the merits and demerits of their young animals. We see this plainly in our own children; we cannot tell whether a child will be tall or short, or what its precise features will be. The question is not, at what period of life any variation may have been caused, but at what period the effects are displayed. The cause may have acted, and I believe often has acted, on one or both parents before the act of generation. It deserves notice that it is of no importance to a very young animal, as long as it is nourished and protected by its parent, whether most of its characters are acquired a little earlier or later in life. It would not signify, for instance, to a bird which obtained its food by having a much-curved beak whether or not while young it possessed a beak of this shape, as long as it was fed by its parents.

I have stated in the first chapter, that at whatever age any variation first appears in the parent, it tends to reappear at a corresponding age in the offspring. Certain variations can only appear at corresponding ages; for instance, peculiarities in the caterpillar, cocoon, or imago states of the silk-moth; or, again, in the full-grown horns of cattle. But variations which, for all that we can see might have appeared either earlier or later in life, likewise tend to reappear at a corresponding age in the offspring and parent. I am far from meaning that this is invariably the case, and I could give several exceptional cases of variations (taking the word in the largest sense) which have supervened at an earlier age in the child than in the parent.

These two principles, namely, that slight variations generally appear at a not very early period of life, and are inherited at a corresponding not early period, explain, as I believe, all the above specified leading facts in embryology. But first let us look to a few analogous cases in our domestic varieties. Some authors who have written on Dogs maintain that the greyhound and bull-dog, though so different, are really closely allied varieties, descended from the same wild stock, hence I was curious to see how far their puppies differed from each other. I was told by breeders that they differed just as much as their parents, and this, judging by the eye, seemed almost to be the case; but on actually measuring the old dogs and their six-days-old puppies, I found that the puppies had not acquired nearly their full amount of proportional difference. So, again, I was told that the foals of cart and race-horses – breeds which have been almost wholly formed by selection under domestication – differed as much as the full-grown animals; but having had careful measurements made of the dams and of three-days-old colts of race and heavy cart-horses, I find that this is by no means the case.

As we have conclusive evidence that the breeds of the Pigeon are descended from a single wild species, I compared the young pigeons within twelve hours after being hatched. I carefully measured the proportions (but will not here give the details) of the beak, width of mouth, length of nostril and of eyelid, size of feet and length of leg, in the wild parent species, in pouters, fantails, runts, barbs, dragons, carriers, and tumblers. Now, some of these birds, when mature, differ in so extraordinary a manner in the length and form of beak, and in other characters, that they would certainly have been ranked as distinct genera if found in a state of nature. But when the nestling birds of these several breeds were placed in a row, though most of them could just be distinguished, the proportional differences in the above specified points were incomparably less than in the full-grown birds. Some characteristic points of difference – for instance, that of the width of mouth – could hardly be detected in the young. But there was one remarkable exception to this rule, for the young of the short-faced tumbler differed from the young of the wild rock-pigeon, and of the other breeds, in almost exactly the same proportions as in the adult stage.

These facts are explained by the above two principles. Fanciers select their dogs, horses, pigeons, etc., for breeding, when nearly grown up. They are indifferent whether the desired qualities are acquired earlier or later in life, if the full-grown animal possesses them. And the cases just given, more especially that of the pigeons, show that the characteristic differences which have been accumulated by man's selection, and which give value to his breeds, do not generally appear at a very early period of life, and are inherited at a corresponding not early period. But the case of the short-faced tumbler, which when twelve hours old possessed its proper characters, proves that this is not the universal rule; for here the characteristic differences must either have appeared at an earlier period than usual, or, if not so, the differences must have been inherited, not at a corresponding, but at an earlier age.

Now, let us apply these two principles to species in a state of nature. Let us take a group of birds, descended from some ancient form and modified through natural selection for different habits. Then, from the many slight successive variations having supervened in the several species at a not early age, and having been inherited at a corresponding age, the young will have been but little modified, and they will still resemble each other much more closely than do the adults, just as we have seen with the breeds of the pigeon. We may extend this view to widely distinct structures and to whole classes. The fore-limbs, for instance, which once served as legs to a remote progenitor, may have become, through a long course of modification, adapted in one descendant to act as hands, in another as paddles, in another as wings; but on the above two principles the fore-limbs will not have been much modified in the embryos of these several forms; although in each form the fore-limb will differ greatly in the adult state. Whatever influence long continued use or disuse may have had in modifying the limbs or other parts of any species, this will chiefly or solely have affected it when nearly mature, when it was compelled to use its full powers to gain its own living; and the effects thus produced will have been transmitted to the offspring at a corresponding nearly mature age. Thus the young will not be modified, or will be modified only in a slight degree, through the effects of the increased use or disuse of parts.

With some animals the successive variations may have supervened at a very early period of life, or the steps may have been inherited at an earlier age than that at which they first occurred. In either of these cases the young or embryo will closely resemble the mature parent-form, as we have seen with the short-faced tumbler. And this is the rule of development in certain whole groups, or in certain sub-groups alone, as with cuttle-fish, land-shells, fresh-water crustaceans, spiders, and some members of the great class of insects. With respect to the final cause of the young in such groups not passing through any metamorphosis, we can see that this would follow from the following contingencies: namely, from the young having to provide at a very early age for their own wants, and from their following the same habits of life with their parents; for in this case it would be indispensable for their existence that they should be modified in the same manner as their parents. Again, with respect to the singular fact that many terrestrial and fresh-water animals do not undergo any metamorphosis, while marine members of the same groups pass through various transformations, Fritz Muller has suggested that the process of slowly modifying and adapting an animal to live on the land or in fresh water, instead of in the sea, would be greatly simplified by its not passing through any larval stage; for it is not probable that places well adapted for both the larval and mature stages, under such new and greatly changed habits of life, would commonly be found unoccupied or ill-occupied by other organisms. In this case the gradual acquirement at an earlier and earlier age of the adult structure would be favoured by natural selection; and all traces of former metamorphoses would finally be lost.

If, on the other hand, it profited the young of an animal to follow habits of life slightly different from those of the parent-form, and consequently to be constructed on a slightly different plan, or if it profited a larva already different from its parent to change still further, then, on the principle of inheritance at corresponding ages, the young or the larvae might be rendered by natural selection more and more different from their parents to any conceivable extent. Differences in the larva might, also, become correlated with successive stages of its development; so that the larva, in the first stage, might come to differ greatly from the larva in the second stage, as is the case with many animals. The adult might also become fitted for sites or habits, in which organs of locomotion or of the senses, etc., would be useless; and in this case the metamorphosis would be retrograde.

From the remarks just made we can see how by changes of structure in the young, in conformity with changed habits of life, together with inheritance at corresponding ages, animals might come to pass through stages of development, perfectly distinct from the primordial condition of their adult progenitors. Most of our best authorities are now convinced that the various larval and pupal stages of insects have thus been acquired through adaptation, and not through inheritance from some ancient form. The curious case of Sitaris – a beetle which passes through certain unusual stages of development – will illustrate how this might occur. The first larval form is described by M. Fabre, as an active, minute insect, furnished with six legs, two long antennae, and four eyes. These larvae are hatched in the nests of bees; and when the male bees emerge from their burrows, in the spring, which they do before the females, the larvae spring on them, and afterwards crawl on to the females while paired with the males. As soon as the female bee deposits her eggs on the surface of the honey stored in the cells, the larvae of the Sitaris leap on the eggs and devour them. Afterwards they undergo a complete change; their eyes disappear; their legs and antennae become rudimentary, and they feed on honey; so that they now more closely resemble the ordinary larvae of insects; ultimately they undergo a further transformation, and finally emerge as the perfect beetle. Now, if an insect, undergoing transformations like those of the Sitaris, were to become the progenitor of a whole new class of insects, the course of development of the new class would be widely different from that of our existing insects; and the first larval stage certainly would not represent the former condition of any adult and ancient form.

On the other hand it is highly probable that with many animals the embryonic or larval stages show us, more or less completely, the condition of the progenitor of the whole group in its adult state. In the great class of the Crustacea, forms wonderfully distinct from each other, namely, suctorial parasites, cirripedes, entomostraca, and even the malacostraca, appear at first as larvae under the nauplius-form; and as these larvae live and feed in the open sea, and are not adapted for any peculiar habits of life, and from other reasons assigned by Fritz Muller, it is probable that at some very remote period an independent adult animal, resembling the Nauplius, existed, and subsequently produced, along several divergent lines of descent, the above-named great Crustacean groups. So again, it is probable, from what we know of the embryos of mammals, birds, fishes and reptiles, that these animals are the modified descendants of some ancient progenitor, which was furnished in its adult state with branchiae, a swim-bladder, four fin-like limbs, and a long tail, all fitted for an aquatic life.

As all the organic beings, extinct and recent, which have ever lived, can be arranged within a few great classes; and as all within each class have, according to our theory, been connected together by fine gradations, the best, and, if our collections were nearly perfect, the only possible arrangement, would be genealogical; descent being the hidden bond of connexion which naturalists have been seeking under the term of the Natural System. On this view we can understand how it is that, in the eyes of most naturalists, the structure of the embryo is even more important for classification than that of the adult. In two or more groups of animals, however much they may differ from each other in structure and habits in their adult condition, if they pass through closely similar embryonic stages, we may feel assured that they are all descended from one parent-form, and are therefore closely related. Thus, community in embryonic structure reveals community of descent; but dissimilarity in embryonic development does not prove discommunity of descent, for in one of two groups the developmental stages may have been suppressed, or may have been so greatly modified through adaptation to new habits of life as to be no longer recognisable. Even in groups, in which the adults have been modified to an extreme degree, community of origin is often revealed by the structure of the larvae; we have seen, for instance, that cirripedes, though externally so like shell-fish, are at once known by their larvae to belong to the great class of crustaceans. As the embryo often shows us more or less plainly the structure of the less modified and ancient progenitor of the group, we can see why ancient and extinct forms so often resemble in their adult state the embryos of existing species of the same class. Agassiz believes this to be a universal law of nature; and we may hope hereafter to see the law proved true. It can, however, be proved true only in those cases in which the ancient state of the progenitor of the group has not been wholly obliterated, either by successive variations having supervened at a very early period of growth, or by such variations having been inherited at an earlier age than that at which they first appeared. It should also be borne in mind, that the law may be true, but yet, owing to the geological record not extending far enough back in time, may remain for a long period, or for ever, incapable of demonstration. The law will not strictly hold good in those cases in which an ancient form became adapted in its larval state to some special line of life, and transmitted the same larval state to a whole group of descendants; for such larval state will not resemble any still more ancient form in its adult state.

Thus, as it seems to me, the leading facts in embryology, which are second to none in importance, are explained on the principle of variations in the many descendants from some one ancient progenitor, having appeared at a not very early period of life, and having been inherited at a corresponding period. Embryology rises greatly in interest, when we look at the embryo as a picture, more or less obscured, of the progenitor, either in its adult or larval state, of all the members of the same great class.

RUDIMENTARY, ATROPHIED, AND ABORTED ORGANS.

Organs or parts in this strange condition, bearing the plain stamp of inutility, are extremely common, or even general, throughout nature. It would be impossible to name one of the higher animals in which some part or other is not in a rudimentary condition. In the mammalia, for instance, the males possess rudimentary mammae; in snakes one lobe of the lungs is rudimentary; in birds the "bastard-wing" may safely be considered as a rudimentary digit, and in some species the whole wing is so far rudimentary that it cannot be used for flight. What can be more curious than the presence of teeth in foetal whales, which when grown up have not a tooth in their heads; or the teeth, which never cut through the gums, in the upper jaws of unborn calves?

Rudimentary organs plainly declare their origin and meaning in various ways. There are beetles belonging to closely allied species, or even to the same identical species, which have either full-sized and perfect wings, or mere rudiments of membrane, which not rarely lie under wing-covers firmly soldered together; and in these cases it is impossible to doubt, that the rudiments represent wings. Rudimentary organs sometimes retain their potentiality: this occasionally occurs with the mammae of male mammals, which have been known to become well developed and to secrete milk. So again in the udders of the genus Bos, there are normally four developed and two rudimentary teats; but the latter in our domestic cows sometimes become well developed and yield milk. In regard to plants, the petals are sometimes rudimentary, and sometimes well developed in the individuals of the same species. In certain plants having separated sexes Kolreuter found that by crossing a species, in which the male flowers included a rudiment of a pistil, with an hermaphrodite species, having of course a well-developed pistil, the rudiment in the hybrid offspring was much increased in size; and this clearly shows that the rudimentary and perfect pistils are essentially alike in nature. An animal may possess various parts in a perfect state, and yet they may in one sense be rudimentary, for they are useless: thus the tadpole of the common salamander or water-newt, as Mr. G.H. Lewes remarks, "has gills, and passes its existence in the water; but the Salamandra atra, which lives high up among the mountains, brings forth its young full-formed. This animal never lives in the water. Yet if we open a gravid female, we find tadpoles inside her with exquisitely feathered gills; and when placed in water they swim about like the tadpoles of the water-newt. Obviously this aquatic organisation has no reference to the future life of the animal, nor has it any adaptation to its embryonic condition; it has solely reference to ancestral adaptations, it repeats a phase in the development of its progenitors."

An organ, serving for two purposes, may become rudimentary or utterly aborted for one, even the more important purpose, and remain perfectly efficient for the other. Thus, in plants, the office of the pistil is to allow the pollen-tubes to reach the ovules within the ovarium. The pistil consists of a stigma supported on the style; but in some Compositae, the male florets, which of course cannot be fecundated, have a rudimentary pistil, for it is not crowned with a stigma; but the style remains well developed and is clothed in the usual manner with hairs, which serve to brush the pollen out of the surrounding and conjoined anthers. Again, an organ may become rudimentary for its proper purpose, and be used for a distinct one: in certain fishes the swim-bladder seems to be rudimentary for its proper function of giving buoyancy, but has become converted into a nascent breathing organ or lung. Many similar instances could be given.

Useful organs, however little they may be developed, unless we have reason to suppose that they were formerly more highly developed, ought not to be considered as rudimentary. They may be in a nascent condition, and in progress towards further development. Rudimentary organs, on the other hand, are either quite useless, such as teeth which never cut through the gums, or almost useless, such as the wings of an ostrich, which serve merely as sails. As organs in this condition would formerly, when still less developed, have been of even less use than at present, they cannot formerly have been produced through variation and natural selection, which acts solely by the preservation of useful modifications. They have been partially retained by the power of inheritance, and relate to a former state of things. It is, however, often difficult to distinguish between rudimentary and nascent organs; for we can judge only by analogy whether a part is capable of further development, in which case alone it deserves to be called nascent. Organs in this condition will always be somewhat rare; for beings thus provided will commonly have been supplanted by their successors with the same organ in a more perfect state, and consequently will have become long ago extinct. The wing of the penguin is of high service, acting as a fin; it may, therefore, represent the nascent state of the wing: not that I believe this to be the case; it is more probably a reduced organ, modified for a new function: the wing of the Apteryx, on the other hand, is quite useless, and is truly rudimentary. Owen considers the simple filamentary limbs of the Lepidosiren as the "beginnings of organs which attain full functional development in higher vertebrates;" but, according to the view lately advocated by Dr. Gunther, they are probably remnants, consisting of the persistent axis of a fin, with the lateral rays or branches aborted. The mammary glands of the Ornithorhynchus may be considered, in comparison with the udders of a cow, as in a nascent condition. The ovigerous frena of certain cirripedes, which have ceased to give attachment to the ova and are feebly developed, are nascent branchiae.

Rudimentary organs in the individuals of the same species are very liable to vary in the degree of their development and in other respects. In closely allied species, also, the extent to which the same organ has been reduced occasionally differs much. This latter fact is well exemplified in the state of the wings of female moths belonging to the same family. Rudimentary organs may be utterly aborted; and this implies, that in certain animals or plants, parts are entirely absent which analogy would lead us to expect to find in them, and which are occasionally found in monstrous individuals. Thus in most of the Scrophulariaceae the fifth stamen is utterly aborted; yet we may conclude that a fifth stamen once existed, for a rudiment of it is found in many species of the family, and this rudiment occasionally becomes perfectly developed, as may sometimes be seen in the common snap-dragon. In tracing the homologies of any part in different members of the same class, nothing is more common, or, in order fully to understand the relations of the parts, more useful than the discovery of rudiments. This is well shown in the drawings given by Owen of the leg bones of the horse, ox, and rhinoceros.

It is an important fact that rudimentary organs, such as teeth in the upper jaws of whales and ruminants, can often be detected in the embryo, but afterwards wholly disappear. It is also, I believe, a universal rule, that a rudimentary part is of greater size in the embryo relatively to the adjoining parts, than in the adult; so that the organ at this early age is less rudimentary, or even cannot be said to be in any degree rudimentary. Hence rudimentary organs in the adult are often said to have retained their embryonic condition.

I have now given the leading facts with respect to rudimentary organs. In reflecting on them, every one must be struck with astonishment; for the same reasoning power which tells us that most parts and organs are exquisitely adapted for certain purposes, tells us with equal plainness that these rudimentary or atrophied organs are imperfect and useless. In works on natural history, rudimentary organs are generally said to have been created "for the sake of symmetry," or in order "to complete the scheme of nature." But this is not an explanation, merely a restatement of the fact. Nor is it consistent with itself: thus the boa-constrictor has rudiments of hind limbs and of a pelvis, and if it be said that these bones have been retained "to complete the scheme of nature," why, as Professor Weismann asks, have they not been retained by other snakes, which do not possess even a vestige of these same bones? What would be thought of an astronomer who maintained that the satellites revolve in elliptic courses round their planets "for the sake of symmetry," because the planets thus revolve round the sun? An eminent physiologist accounts for the presence of rudimentary organs, by supposing that they serve to excrete matter in excess, or matter injurious to the system; but can we suppose that the minute papilla, which often represents the pistil in male flowers, and which is formed of mere cellular tissue, can thus act? Can we suppose that rudimentary teeth, which are subsequently absorbed, are beneficial to the rapidly growing embryonic calf by removing matter so precious as phosphate of lime? When a man's fingers have been amputated, imperfect nails have been known to appear on the stumps, and I could as soon believe that these vestiges of nails are developed in order to excrete horny matter, as that the rudimentary nails on the fin of the manatee have been developed for this same purpose.

On the view of descent with modification, the origin of rudimentary organs is comparatively simple; and we can understand to a large extent the laws governing their imperfect development. We have plenty of cases of rudimentary organs in our domestic productions, as the stump of a tail in tailless breeds, the vestige of an ear in earless breeds of sheep – the reappearance of minute dangling horns in hornless breeds of cattle, more especially, according to Youatt, in young animals – and the state of the whole flower in the cauliflower. We often see rudiments of various parts in monsters; but I doubt whether any of these cases throw light on the origin of rudimentary organs in a state of nature, further than by showing that rudiments can be produced; for the balance of evidence clearly indicates that species under nature do not undergo great and abrupt changes. But we learn from the study of our domestic productions that the disuse of parts leads to their reduced size; and that the result is inherited.

It appears probable that disuse has been the main agent in rendering organs rudimentary. It would at first lead by slow steps to the more and more complete reduction of a part, until at last it became rudimentary – as in the case of the eyes of animals inhabiting dark caverns, and of the wings of birds inhabiting oceanic islands, which have seldom been forced by beasts of prey to take flight, and have ultimately lost the power of flying. Again, an organ, useful under certain conditions, might become injurious under others, as with the wings of beetles living on small and exposed islands; and in this case natural selection will have aided in reducing the organ, until it was rendered harmless and rudimentary.

Any change in structure and function, which can be effected by small stages, is within the power of natural selection; so that an organ rendered, through changed habits of life, useless or injurious for one purpose, might be modified and used for another purpose. An organ might, also, be retained for one alone of its former functions. Organs, originally formed by the aid of natural selection, when rendered useless may well be variable, for their variations can no longer be checked by natural selection. All this agrees well with what we see under nature. Moreover, at whatever period of life either disuse or selection reduces an organ, and this will generally be when the being has come to maturity and to exert its full powers of action, the principle of inheritance at corresponding ages will tend to reproduce the organ in its reduced state at the same mature age, but will seldom affect it in the embryo. Thus we can understand the greater size of rudimentary organs in the embryo relatively to the adjoining parts, and their lesser relative size in the adult. If, for instance, the digit of an adult animal was used less and less during many generations, owing to some change of habits, or if an organ or gland was less and less functionally exercised, we may infer that it would become reduced in size in the adult descendants of this animal, but would retain nearly its original standard of development in the embryo.

There remains, however, this difficulty. After an organ has ceased being used, and has become in consequence much reduced, how can it be still further reduced in size until the merest vestige is left; and how can it be finally quite obliterated? It is scarcely possible that disuse can go on producing any further effect after the organ has once been rendered functionless. Some additional explanation is here requisite which I cannot give. If, for instance, it could be proved that every part of the organisation tends to vary in a greater degree towards diminution than toward augmentation of size, then we should be able to understand how an organ which has become useless would be rendered, independently of the effects of disuse, rudimentary and would at last be wholly suppressed; for the variations towards diminished size would no longer be checked by natural selection. The principle of the economy of growth, explained in a former chapter, by which the materials forming any part, if not useful to the possessor, are saved as far as is possible, will perhaps come into play in rendering a useless part rudimentary. But this principle will almost necessarily be confined to the earlier stages of the process of reduction; for we cannot suppose that a minute papilla, for instance, representing in a male flower the pistil of the female flower, and formed merely of cellular tissue, could be further reduced or absorbed for the sake of economising nutriment.

Finally, as rudimentary organs, by whatever steps they may have been degraded into their present useless condition, are the record of a former state of things, and have been retained solely through the power of inheritance – we can understand, on the genealogical view of classification, how it is that systematists, in placing organisms in their proper places in the natural system, have often found rudimentary parts as useful as, or even sometimes more useful than, parts of high physiological importance. Rudimentary organs may be compared with the letters in a word, still retained in the spelling, but become useless in the pronunciation, but which serve as a clue for its derivation. On the view of descent with modification, we may conclude that the existence of organs in a rudimentary, imperfect, and useless condition, or quite aborted, far from presenting a strange difficulty, as they assuredly do on the old doctrine of creation, might even have been anticipated in accordance with the views here explained.

SUMMARY.

In this chapter I have attempted to show that the arrangement of all organic beings throughout all time in groups under groups – that the nature of the relationships by which all living and extinct organisms are united by complex, radiating, and circuitous lines of affinities into a few grand classes – the rules followed and the difficulties encountered by naturalists in their classifications – the value set upon characters, if constant and prevalent, whether of high or of the most trifling importance, or, as with rudimentary organs of no importance – the wide opposition in value between analogical or adaptive characters, and characters of true affinity; and other such rules – all naturally follow if we admit the common parentage of allied forms, together with their modification through variation and natural selection, with the contingencies of extinction and divergence of character. In considering this view of classification, it should be borne in mind that the element of descent has been universally used in ranking together the sexes, ages, dimorphic forms, and acknowledged varieties of the same species, however much they may differ from each other in structure. If we extend the use of this element of descent – the one certainly known cause of similarity in organic beings – we shall understand what is meant by the Natural System: it is genealogical in its attempted arrangement, with the grades of acquired difference marked by the terms, varieties, species, genera, families, orders, and classes.

On this same view of descent with modification, most of the great facts in Morphology become intelligible – whether we look to the same pattern displayed by the different species of the same class in their homologous organs, to whatever purpose applied, or to the serial and lateral homologies in each individual animal and plant.

On the principle of successive slight variations, not necessarily or generally supervening at a very early period of life, and being inherited at a corresponding period, we can understand the leading facts in embryology; namely, the close resemblance in the individual embryo of the parts which are homologous, and which when matured become widely different in structure and function; and the resemblance of the homologous parts or organs in allied though distinct species, though fitted in the adult state for habits as different as is possible. Larvae are active embryos, which have become specially modified in a greater or less degree in relation to their habits of life, with their modifications inherited at a corresponding early age. On these same principles, and bearing in mind that when organs are reduced in size, either from disuse or through natural selection, it will generally be at that period of life when the being has to provide for its own wants, and bearing in mind how strong is the force of inheritance – the occurrence of rudimentary organs might even have been anticipated. The importance of embryological characters and of rudimentary organs in classification is intelligible, on the view that a natural arrangement must be genealogical.

Finally, the several classes of facts which have been considered in this chapter, seem to me to proclaim so plainly, that the innumerable species, genera and families, with which this world is peopled, are all descended, each within its own class or group, from common parents, and have all been modified in the course of descent, that I should without hesitation adopt this view, even if it were unsupported by other facts or arguments.

CHAPTER XV. RECAPITULATION AND CONCLUSION

Recapitulation of the objections to the theory of Natural Selection – Recapitulation of the general and special circumstances in its favour – Causes of the general belief in the immutability of species – How far the theory of Natural Selection may be extended – Effects of its adoption on the study of Natural History – Concluding remarks.

As this whole volume is one long argument, it may be convenient to the reader to have the leading facts and inferences briefly recapitulated.

That many and serious objections may be advanced against the theory of descent with modification through variation and natural selection, I do not deny. I have endeavoured to give to them their full force. Nothing at first can appear more difficult to believe than that the more complex organs and instincts have been perfected, not by means superior to, though analogous with, human reason, but by the accumulation of innumerable slight variations, each good for the individual possessor. Nevertheless, this difficulty, though appearing to our imagination insuperably great, cannot be considered real if we admit the following propositions, namely, that all parts of the organisation and instincts offer, at least individual differences – that there is a struggle for existence leading to the preservation of profitable deviations of structure or instinct – and, lastly, that gradations in the state of perfection of each organ may have existed, each good of its kind. The truth of these propositions cannot, I think, be disputed.

It is, no doubt, extremely difficult even to conjecture by what gradations many structures have been perfected, more especially among broken and failing groups of organic beings, which have suffered much extinction; but we see so many strange gradations in nature, that we ought to be extremely cautious in saying that any organ or instinct, or any whole structure, could not have arrived at its present state by many graduated steps. There are, it must be admitted, cases of special difficulty opposed to the theory of natural selection; and one of the most curious of these is the existence in the same community of two or three defined castes of workers or sterile female ants; but I have attempted to show how these difficulties can be mastered.

With respect to the almost universal sterility of species when first crossed, which forms so remarkable a contrast with the almost universal fertility of varieties when crossed, I must refer the reader to the recapitulation of the facts given at the end of the ninth chapter, which seem to me conclusively to show that this sterility is no more a special endowment than is the incapacity of two distinct kinds of trees to be grafted together; but that it is incidental on differences confined to the reproductive systems of the intercrossed species. We see the truth of this conclusion in the vast difference in the results of crossing the same two species reciprocally – that is, when one species is first used as the father and then as the mother. Analogy from the consideration of dimorphic and trimorphic plants clearly leads to the same conclusion, for when the forms are illegitimately united, they yield few or no seed, and their offspring are more or less sterile; and these forms belong to the same undoubted species, and differ from each other in no respect except in their reproductive organs and functions.

Although the fertility of varieties when intercrossed, and of their mongrel offspring, has been asserted by so many authors to be universal, this cannot be considered as quite correct after the facts given on the high authority of Gartner and Kolreuter. Most of the varieties which have been experimented on have been produced under domestication; and as domestication (I do not mean mere confinement) almost certainly tends to eliminate that sterility which, judging from analogy, would have affected the parent-species if intercrossed, we ought not to expect that domestication would likewise induce sterility in their modified descendants when crossed. This elimination of sterility apparently follows from the same cause which allows our domestic animals to breed freely under diversified circumstances; and this again apparently follows from their having been gradually accustomed to frequent changes in their conditions of life.

A double and parallel series of facts seems to throw much light on the sterility of species, when first crossed, and of their hybrid offspring. On the one side, there is good reason to believe that slight changes in the conditions of life give vigour and fertility to all organic beings. We know also that a cross between the distinct individuals of the same variety, and between distinct varieties, increases the number of their offspring, and certainly gives to them increased size and vigour. This is chiefly owing to the forms which are crossed having been exposed to somewhat different conditions of life; for I have ascertained by a labourious series of experiments that if all the individuals of the same variety be subjected during several generations to the same conditions, the good derived from crossing is often much diminished or wholly disappears. This is one side of the case. On the other side, we know that species which have long been exposed to nearly uniform conditions, when they are subjected under confinement to new and greatly changed conditions, either perish, or if they survive, are rendered sterile, though retaining perfect health. This does not occur, or only in a very slight degree, with our domesticated productions, which have long been exposed to fluctuating conditions. Hence when we find that hybrids produced by a cross between two distinct species are few in number, owing to their perishing soon after conception or at a very early age, or if surviving that they are rendered more or less sterile, it seems highly probable that this result is due to their having been in fact subjected to a great change in their conditions of life, from being compounded of two distinct organisations. He who will explain in a definite manner why, for instance, an elephant or a fox will not breed under confinement in its native country, whilst the domestic pig or dog will breed freely under the most diversified conditions, will at the same time be able to give a definite answer to the question why two distinct species, when crossed, as well as their hybrid offspring, are generally rendered more or less sterile, while two domesticated varieties when crossed and their mongrel offspring are perfectly fertile.

Turning to geographical distribution, the difficulties encountered on the theory of descent with modification are serious enough. All the individuals of the same species, and all the species of the same genus, or even higher group, are descended from common parents; and therefore, in however distant and isolated parts of the world they may now be found, they must in the course of successive generations have travelled from some one point to all the others. We are often wholly unable even to conjecture how this could have been effected. Yet, as we have reason to believe that some species have retained the same specific form for very long periods of time, immensely long as measured by years, too much stress ought not to be laid on the occasional wide diffusion of the same species; for during very long periods there will always have been a good chance for wide migration by many means. A broken or interrupted range may often be accounted for by the extinction of the species in the intermediate regions. It cannot be denied that we are as yet very ignorant as to the full extent of the various climatical and geographical changes which have affected the earth during modern periods; and such changes will often have facilitated migration. As an example, I have attempted to show how potent has been the influence of the Glacial period on the distribution of the same and of allied species throughout the world. We are as yet profoundly ignorant of the many occasional means of transport. With respect to distinct species of the same genus, inhabiting distant and isolated regions, as the process of modification has necessarily been slow, all the means of migration will have been possible during a very long period; and consequently the difficulty of the wide diffusion of the species of the same genus is in some degree lessened.

As according to the theory of natural selection an interminable number of intermediate forms must have existed, linking together all the species in each group by gradations as fine as our existing varieties, it may be asked, Why do we not see these linking forms all around us? Why are not all organic beings blended together in an inextricable chaos? With respect to existing forms, we should remember that we have no right to expect (excepting in rare cases) to discover DIRECTLY connecting links between them, but only between each and some extinct and supplanted form. Even on a wide area, which has during a long period remained continuous, and of which the climatic and other conditions of life change insensibly in proceeding from a district occupied by one species into another district occupied by a closely allied species, we have no just right to expect often to find intermediate varieties in the intermediate zones. For we have reason to believe that only a few species of a genus ever undergo change; the other species becoming utterly extinct and leaving no modified progeny. Of the species which do change, only a few within the same country change at the same time; and all modifications are slowly effected. I have also shown that the intermediate varieties which probably at first existed in the intermediate zones, would be liable to be supplanted by the allied forms on either hand; for the latter, from existing in greater numbers, would generally be modified and improved at a quicker rate than the intermediate varieties, which existed in lesser numbers; so that the intermediate varieties would, in the long run, be supplanted and exterminated.

On this doctrine of the extermination of an infinitude of connecting links, between the living and extinct inhabitants of the world, and at each successive period between the extinct and still older species, why is not every geological formation charged with such links? Why does not every collection of fossil remains afford plain evidence of the gradation and mutation of the forms of life? Although geological research has undoubtedly revealed the former existence of many links, bringing numerous forms of life much closer together, it does not yield the infinitely many fine gradations between past and present species required on the theory, and this is the most obvious of the many objections which may be urged against it. Why, again, do whole groups of allied species appear, though this appearance is often false, to have come in suddenly on the successive geological stages? Although we now know that organic beings appeared on this globe, at a period incalculably remote, long before the lowest bed of the Cambrian system was deposited, why do we not find beneath this system great piles of strata stored with the remains of the progenitors of the Cambrian fossils? For on the theory, such strata must somewhere have been deposited at these ancient and utterly unknown epochs of the world's history.

I can answer these questions and objections only on the supposition that the geological record is far more imperfect than most geologists believe. The number of specimens in all our museums is absolutely as nothing compared with the countless generations of countless species which have certainly existed. The parent form of any two or more species would not be in all its characters directly intermediate between its modified offspring, any more than the rock-pigeon is directly intermediate in crop and tail between its descendants, the pouter and fantail pigeons. We should not be able to recognise a species as the parent of another and modified species, if we were to examine the two ever so closely, unless we possessed most of the intermediate links; and owing to the imperfection of the geological record, we have no just right to expect to find so many links. If two or three, or even more linking forms were discovered, they would simply be ranked by many naturalists as so many new species, more especially if found in different geological substages, let their differences be ever so slight. Numerous existing doubtful forms could be named which are probably varieties; but who will pretend that in future ages so many fossil links will be discovered, that naturalists will be able to decide whether or not these doubtful forms ought to be called varieties? Only a small portion of the world has been geologically explored. Only organic beings of certain classes can be preserved in a fossil condition, at least in any great number. Many species when once formed never undergo any further change but become extinct without leaving modified descendants; and the periods during which species have undergone modification, though long as measured by years, have probably been short in comparison with the periods during which they retained the same form. It is the dominant and widely ranging species which vary most frequently and vary most, and varieties are often at first local – both causes rendering the discovery of intermediate links in any one formation less likely. Local varieties will not spread into other and distant regions until they are considerably modified and improved; and when they have spread, and are discovered in a geological formation, they appear as if suddenly created there, and will be simply classed as new species. Most formations have been intermittent in their accumulation; and their duration has probably been shorter than the average duration of specific forms. Successive formations are in most cases separated from each other by blank intervals of time of great length, for fossiliferous formations thick enough to resist future degradation can, as a general rule, be accumulated only where much sediment is deposited on the subsiding bed of the sea. During the alternate periods of elevation and of stationary level the record will generally be blank. During these latter periods there will probably be more variability in the forms of life; during periods of subsidence, more extinction.