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Origin of Species by Means of Natural Selection; or, the Preservation of...

ON THE NATURE OF THE AFFINITIES CONNECTING ORGANIC BEINGS.

As the modified descendants of dominant species, belonging to the larger genera, tend to inherit the advantages which made the groups to which they belong large and their parents dominant, they are almost sure to spread widely, and to seize on more and more places in the economy of nature. The larger and more dominant groups within each class thus tend to go on increasing in size, and they consequently supplant many smaller and feebler groups. Thus, we can account for the fact that all organisms, recent and extinct, are included under a few great orders and under still fewer classes. As showing how few the higher groups are in number, and how widely they are spread throughout the world, the fact is striking that the discovery of Australia has not added an insect belonging to a new class, and that in the vegetable kingdom, as I learn from Dr. Hooker, it has added only two or three families of small size.

In the chapter on geological succession I attempted to show, on the principle of each group having generally diverged much in character during the long-continued process of modification, how it is that the more ancient forms of life often present characters in some degree intermediate between existing groups. As some few of the old and intermediate forms having transmitted to the present day descendants but little modified, these constitute our so-called osculant or aberrant groups. The more aberrant any form is, the greater must be the number of connecting forms which have been exterminated and utterly lost. And we have evidence of aberrant groups having suffered severely from extinction, for they are almost always represented by extremely few species; and such species as do occur are generally very distinct from each other, which again implies extinction. The genera Ornithorhynchus and Lepidosiren, for example, would not have been less aberrant had each been represented by a dozen species, instead of as at present by a single one, or by two or three. We can, I think, account for this fact only by looking at aberrant groups as forms which have been conquered by more successful competitors, with a few members still preserved under unusually favourable conditions.

Mr. Waterhouse has remarked that when a member belonging to one group of animals exhibits an affinity to a quite distinct group, this affinity in most cases is general and not special: thus, according to Mr. Waterhouse, of all Rodents, the bizcacha is most nearly related to Marsupials; but in the points in which it approaches this order, its relations are general, that is, not to any one Marsupial species more than to another. As these points of affinity are believed to be real and not merely adaptive, they must be due in accordance with our view to inheritance from a common progenitor. Therefore, we must suppose either that all Rodents, including the bizcacha, branched off from some ancient Marsupial, which will naturally have been more or less intermediate in character with respect to all existing Marsupials; or that both Rodents and Marsupials branched off from a common progenitor, and that both groups have since undergone much modification in divergent directions. On either view we must suppose that the bizcacha has retained, by inheritance, more of the character of its ancient progenitor than have other Rodents; and therefore it will not be specially related to any one existing Marsupial, but indirectly to all or nearly all Marsupials, from having partially retained the character of their common progenitor, or of some early member of the group. On the other hand, of all Marsupials, as Mr. Waterhouse has remarked, the Phascolomys resembles most nearly, not any one species, but the general order of Rodents. In this case, however, it may be strongly suspected that the resemblance is only analogical, owing to the Phascolomys having become adapted to habits like those of a Rodent. The elder De Candolle has made nearly similar observations on the general nature of the affinities of distinct families of plants.

On the principle of the multiplication and gradual divergence in character of the species descended from a common progenitor, together with their retention by inheritance of some characters in common, we can understand the excessively complex and radiating affinities by which all the members of the same family or higher group are connected together. For the common progenitor of a whole family, now broken up by extinction into distinct groups and subgroups, will have transmitted some of its characters, modified in various ways and degrees, to all the species; and they will consequently be related to each other by circuitous lines of affinity of various lengths (as may be seen in the diagram so often referred to), mounting up through many predecessors. As it is difficult to show the blood-relationship between the numerous kindred of any ancient and noble family, even by the aid of a genealogical tree, and almost impossible to do so without this aid, we can understand the extraordinary difficulty which naturalists have experienced in describing, without the aid of a diagram, the various affinities which they perceive between the many living and extinct members of the same great natural class.

Extinction, as we have seen in the fourth chapter, has played an important part in defining and widening the intervals between the several groups in each class. We may thus account for the distinctness of whole classes from each other—for instance, of birds from all other vertebrate animals—by the belief that many ancient forms of life have been utterly lost, through which the early progenitors of birds were formerly connected with the early progenitors of the other and at that time less differentiated vertebrate classes. There has been much less extinction of the forms of life which once connected fishes with Batrachians. There has been still less within some whole classes, for instance the Crustacea, for here the most wonderfully diverse forms are still linked together by a long and only partially broken chain of affinities. Extinction has only defined the groups: it has by no means made them; for if every form which has ever lived on this earth were suddenly to reappear, though it would be quite impossible to give definitions by which each group could be distinguished, still a natural classification, or at least a natural arrangement, would be possible. We shall see this by turning to the diagram: the letters, A to L, may represent eleven Silurian genera, some of which have produced large groups of modified descendants, with every link in each branch and sub-branch still alive; and the links not greater than those between existing varieties. In this case it would be quite impossible to give definitions by which the several members of the several groups could be distinguished from their more immediate parents and descendants. Yet the arrangement in the diagram would still hold good and would be natural; for, on the principle of inheritance, all the forms descended, for instance from A, would have something in common. In a tree we can distinguish this or that branch, though at the actual fork the two unite and blend together. We could not, as I have said, define the several groups; but we could pick out types, or forms, representing most of the characters of each group, whether large or small, and thus give a general idea of the value of the differences between them. This is what we should be driven to, if we were ever to succeed in collecting all the forms in any one class which have lived throughout all time and space. Assuredly we shall never succeed in making so perfect a collection: nevertheless, in certain classes, we are tending toward this end; and Milne Edwards has lately insisted, in an able paper, on the high importance of looking to types, whether or not we can separate and define the groups to which such types belong.

Finally, we have seen that natural selection, which follows from the struggle for existence, and which almost inevitably leads to extinction and divergence of character in the descendants from any one parent-species, explains that great and universal feature in the affinities of all organic beings, namely, their subordination in group under group. We use the element of descent in classing the individuals of both sexes and of all ages under one species, although they may have but few characters in common; we use descent in classing acknowledged varieties, however different they may be from their parents; and I believe that this element of descent is the hidden bond of connexion which naturalists have sought under the term of the Natural System. On this idea of the natural system being, in so far as it has been perfected, genealogical in its arrangement, with the grades of difference expressed by the terms genera, families, orders, etc., we can understand the rules which we are compelled to follow in our classification. We can understand why we value certain resemblances far more than others; why we use rudimentary and useless organs, or others of trifling physiological importance; why, in finding the relations between one group and another, we summarily reject analogical or adaptive characters, and yet use these same characters within the limits of the same group. We can clearly see how it is that all living and extinct forms can be grouped together within a few great classes; and how the several members of each class are connected together by the most complex and radiating lines of affinities. We shall never, probably, disentangle the inextricable web of the affinities between the members of any one class; but when we have a distinct object in view, and do not look to some unknown plan of creation, we may hope to make sure but slow progress.

Professor Haeckel in his "Generelle Morphologie" and in another works, has recently brought his great knowledge and abilities to bear on what he calls phylogeny, or the lines of descent of all organic beings. In drawing up the several series he trusts chiefly to embryological characters, but receives aid from homologous and rudimentary organs, as well as from the successive periods at which the various forms of life are believed to have first appeared in our geological formations. He has thus boldly made a great beginning, and shows us how classification will in the future be treated.

MORPHOLOGY.

We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation. This resemblance is often expressed by the term "unity of type;" or by saying that the several parts and organs in the different species of the class are homologous. The whole subject is included under the general term of Morphology. This is one of the most interesting departments of natural history, and may almost be said to be its very soul. What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include similar bones, in the same relative positions? How curious it is, to give a subordinate though striking instance, that the hind feet of the kangaroo, which are so well fitted for bounding over the open plains—those of the climbing, leaf-eating koala, equally well fitted for grasping the branches of trees—those of the ground-dwelling, insect or root-eating, bandicoots—and those of some other Australian marsupials—should all be constructed on the same extraordinary type, namely with the bones of the second and third digits extremely slender and enveloped within the same skin, so that they appear like a single toe furnished with two claws. Notwithstanding this similarity of pattern, it is obvious that the hind feet of these several animals are used for as widely different purposes as it is possible to conceive. The case is rendered all the more striking by the American opossums, which follow nearly the same habits of life as some of their Australian relatives, having feet constructed on the ordinary plan. Professor Flower, from whom these statements are taken, remarks in conclusion: "We may call this conformity to type, without getting much nearer to an explanation of the phenomenon;" and he then adds "but is it not powerfully suggestive of true relationship, of inheritance from a common ancestor?"

Geoffroy St. Hilaire has strongly insisted on the high importance of relative position or connexion in homologous parts; they may differ to almost any extent in form and size, and yet remain connected together in the same invariable order. We never find, for instance, the bones of the arm and forearm, or of the thigh and leg, transposed. Hence the same names can be given to the homologous bones in widely different animals. We see the same great law in the construction of the mouths of insects: what can be more different than the immensely long spiral proboscis of a sphinx-moth, the curious folded one of a bee or bug, and the great jaws of a beetle? Yet all these organs, serving for such widely different purposes, are formed by infinitely numerous modifications of an upper lip, mandibles, and two pairs of maxillae. The same law governs the construction of the mouths and limbs of crustaceans. So it is with the flowers of plants.

Nothing can be more hopeless than to attempt to explain this similarity of pattern in members of the same class, by utility or by the doctrine of final causes. The hopelessness of the attempt has been expressly admitted by Owen in his most interesting work on the "Nature of Limbs." On the ordinary view of the independent creation of each being, we can only say that so it is; that it has pleased the Creator to construct all the animals and plants in each great class on a uniform plan; but this is not a scientific explanation.

The explanation is to a large extent simple, on the theory of the selection of successive slight modifications, each being profitable in some way to the modified form, but often affecting by correlation other parts of the organisation. In changes of this nature, there will be little or no tendency to alter the original pattern, or to transpose the parts. The bones of a limb might be shortened and flattened to any extent, becoming at the same time enveloped in thick membrane, so as to serve as a fin; or a webbed hand might have all its bones, or certain bones, lengthened to any extent, with the membrane connecting them increased, so as to serve as a wing; yet all these modifications would not tend to alter the framework of the bones or the relative connexion of the parts. If we suppose that an early progenitor—the archetype, as it may be called—of all mammals, birds and reptiles, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain signification of the homologous construction of the limbs throughout the class. So with the mouths of insects, we have only to suppose that their common progenitor had an upper lip, mandibles, and two pairs of maxillae, these parts being perhaps very simple in form; and then natural selection will account for the infinite diversity in structure and function of the mouths of insects. Nevertheless, it is conceivable that the general pattern of an organ might become so much obscured as to be finally lost, by the reduction and ultimately by the complete abortion of certain parts, by the fusion of other parts, and by the doubling or multiplication of others, variations which we know to be within the limits of possibility. In the paddles of the gigantic extinct sea-lizards, and in the mouths of certain suctorial crustaceans, the general pattern seems thus to have become partially obscured.

There is another and equally curious branch of our subject; namely, serial homologies, or the comparison of the different parts or organs in the same individual, and not of the same parts or organs in different members of the same class. Most physiologists believe that the bones of the skull are homologous—that is, correspond in number and in relative connexion—with the elemental parts of a certain number of vertebrae. The anterior and posterior limbs in all the higher vertebrate classes are plainly homologous. So it is with the wonderfully complex jaws and legs of crustaceans. It is familiar to almost every one, that in a flower the relative position of the sepals, petals, stamens, and pistils, as well as their intimate structure, are intelligible on the view that they consist of metamorphosed leaves, arranged in a spire. In monstrous plants, we often get direct evidence of the possibility of one organ being transformed into another; and we can actually see, during the early or embryonic stages of development in flowers, as well as in crustaceans and many other animals, that organs, which when mature become extremely different are at first exactly alike.

How inexplicable are the cases of serial homologies on the ordinary view of creation! Why should the brain be enclosed in a box composed of such numerous and such extraordinarily shaped pieces of bone apparently representing vertebrae? As Owen has remarked, the benefit derived from the yielding of the separate pieces in the act of parturition by mammals, will by no means explain the same construction in the skulls of birds and reptiles. Why should similar bones have been created to form the wing and the leg of a bat, used as they are for such totally different purposes, namely flying and walking? Why should one crustacean, which has an extremely complex mouth formed of many parts, consequently always have fewer legs; or conversely, those with many legs have simpler mouths? Why should the sepals, petals, stamens, and pistils, in each flower, though fitted for such distinct purposes, be all constructed on the same pattern?

On the theory of natural selection, we can, to a certain extent, answer these questions. We need not here consider how the bodies of some animals first became divided into a series of segments, or how they became divided into right and left sides, with corresponding organs, for such questions are almost beyond investigation. It is, however, probable that some serial structures are the result of cells multiplying by division, entailing the multiplication of the parts developed from such cells. It must suffice for our purpose to bear in mind that an indefinite repetition of the same part or organ is the common characteristic, as Owen has remarked, of all low or little specialised forms; therefore the unknown progenitor of the Vertebrata probably possessed many vertebrae; the unknown progenitor of the Articulata, many segments; and the unknown progenitor of flowering plants, many leaves arranged in one or more spires. We have also formerly seen that parts many times repeated are eminently liable to vary, not only in number, but in form. Consequently such parts, being already present in considerable numbers, and being highly variable, would naturally afford the materials for adaptation to the most different purposes; yet they would generally retain, through the force of inheritance, plain traces of their original or fundamental resemblance. They would retain this resemblance all the more, as the variations, which afforded the basis for their subsequent modification through natural selection, would tend from the first to be similar; the parts being at an early stage of growth alike, and being subjected to nearly the same conditions. Such parts, whether more or less modified, unless their common origin became wholly obscured, would be serially homologous.

In the great class of molluscs, though the parts in distinct species can be shown to be homologous, only a few serial homologies; such as the valves of Chitons, can be indicated; that is, we are seldom enabled to say that one part is homologous with another part in the same individual. And we can understand this fact; for in molluscs, even in the lowest members of the class, we do not find nearly so much indefinite repetition of any one part as we find in the other great classes of the animal and vegetable kingdoms.

But morphology is a much more complex subject than it at first appears, as has lately been well shown in a remarkable paper by Mr. E. Ray Lankester, who has drawn an important distinction between certain classes of cases which have all been equally ranked by naturalists as homologous. He proposes to call the structures which resemble each other in distinct animals, owing to their descent from a common progenitor with subsequent modification, "homogenous"; and the resemblances which cannot thus be accounted for, he proposes to call "homoplastic". For instance, he believes that the hearts of birds and mammals are as a whole homogenous—that is, have been derived from a common progenitor; but that the four cavities of the heart in the two classes are homoplastic—that is, have been independently developed. Mr. Lankester also adduces the close resemblance of the parts on the right and left sides of the body, and in the successive segments of the same individual animal; and here we have parts commonly called homologous which bear no relation to the descent of distinct species from a common progenitor. Homoplastic structures are the same with those which I have classed, though in a very imperfect manner, as analogous modifications or resemblances. Their formation may be attributed in part to distinct organisms, or to distinct parts of the same organism, having varied in an analogous manner; and in part to similar modifications, having been preserved for the same general purpose or function, of which many instances have been given.

Naturalists frequently speak of the skull as formed of metamorphosed vertebrae; the jaws of crabs as metamorphosed legs; the stamens and pistils in flowers as metamorphosed leaves; but it would in most cases be more correct, as Professor Huxley has remarked, to speak of both skull and vertebrae, jaws and legs, etc., as having been metamorphosed, not one from the other, as they now exist, but from some common and simpler element. Most naturalists, however, use such language only in a metaphorical sense: they are far from meaning that during a long course of descent, primordial organs of any kind—vertebrae in the one case and legs in the other—have actually been converted into skulls or jaws. Yet so strong is the appearance of this having occurred that naturalists can hardly avoid employing language having this plain signification. According to the views here maintained, such language may be used literally; and the wonderful fact of the jaws, for instance, of a crab retaining numerous characters, which they probably would have retained through inheritance, if they had really been metamorphosed from true though extremely simple legs, is in part explained.

DEVELOPMENT AND EMBRYOLOGY.

This is one of the most important subjects in the whole round of natural history. The metamorphoses of insects, with which every one is familiar, are generally effected abruptly by a few stages; but the transformations are in reality numerous and gradual, though concealed. A certain ephemerous insect (Chloeon) during its development, moults, as shown by Sir J. Lubbock, above twenty times, and each time undergoes a certain amount of change; and in this case we see the act of metamorphosis performed in a primary and gradual manner. Many insects, and especially certain crustaceans, show us what wonderful changes of structure can be effected during development. Such changes, however, reach their acme in the so-called alternate generations of some of the lower animals. It is, for instance, an astonishing fact that a delicate branching coralline, studded with polypi, and attached to a submarine rock, should produce, first by budding and then by transverse division, a host of huge floating jelly-fishes; and that these should produce eggs, from which are hatched swimming animalcules, which attach themselves to rocks and become developed into branching corallines; and so on in an endless cycle. The belief in the essential identity of the process of alternate generation and of ordinary metamorphosis has been greatly strengthened by Wagner's discovery of the larva or maggot of a fly, namely the Cecidomyia, producing asexually other larvae, and these others, which finally are developed into mature males and females, propagating their kind in the ordinary manner by eggs.

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.


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