The Origin of Species by Means of Natural Selection

Naturalists, as we have seen, try to arrange the species, genera and families in each class, on what is called the Natural System. But what is meant by this system? Some authors look at it merely as a scheme for arranging together those living objects which are most alike, and for separating those which are most unlike; or as an artificial method of enunciating, as briefly as possible, general propositions – that is, by one sentence to give the characters common, for instance, to all mammals, by another those common to all carnivora, by another those common to the dog-genus, and then, by adding a single sentence, a full description is given of each kind of dog. The ingenuity and utility of this system are indisputable. But many naturalists think that something more is meant by the Natural System; they believe that it reveals the plan of the Creator; but unless it be specified whether order in time or space, or both, or what else is meant by the plan of the Creator, it seems to me that nothing is thus added to our knowledge. Expressions such as that famous one by Linnaeus, which we often meet with in a more or less concealed form, namely, that the characters do not make the genus, but that the genus gives the characters, seem to imply that some deeper bond is included in our classifications than mere resemblance. I believe that this is the case, and that community of descent – the one known cause of close similarity in organic beings – is the bond, which, though observed by various degrees of modification, is partially revealed to us by our classifications.

Let us now consider the rules followed in classification, and the difficulties which are encountered on the view that classification either gives some unknown plan of creation, or is simply a scheme for enunciating general propositions and of placing together the forms most like each other. It might have been thought (and was in ancient times thought) that those parts of the structure which determined the habits of life, and the general place of each being in the economy of nature, would be of very high importance in classification. Nothing can be more false. No one regards the external similarity of a mouse to a shrew, of a dugong to a whale, of a whale to a fish, as of any importance. These resemblances, though so intimately connected with the whole life of the being, are ranked as merely "adaptive or analogical characters;" but to the consideration of these resemblances we shall recur. It may even be given as a general rule, that the less any part of the organisation is concerned with special habits, the more important it becomes for classification. As an instance: Owen, in speaking of the dugong, says, "The generative organs, being those which are most remotely related to the habits and food of an animal, I have always regarded as affording very clear indications of its true affinities. We are least likely in the modifications of these organs to mistake a merely adaptive for an essential character." With plants how remarkable it is that the organs of vegetation, on which their nutrition and life depend, are of little signification; whereas the organs of reproduction, with their product the seed and embryo, are of paramount importance! So again, in formerly discussing certain morphological characters which are not functionally important, we have seen that they are often of the highest service in classification. This depends on their constancy throughout many allied groups; and their constancy chiefly depends on any slight deviations not having been preserved and accumulated by natural selection, which acts only on serviceable characters.

That the mere physiological importance of an organ does not determine its classificatory value, is almost proved by the fact, that in allied groups, in which the same organ, as we have every reason to suppose, has nearly the same physiological value, its classificatory value is widely different. No naturalist can have worked at any group without being struck with this fact; and it has been fully acknowledged in the writings of almost every author. It will suffice to quote the highest authority, Robert Brown, who, in speaking of certain organs in the Proteaceae, says their generic importance, "like that of all their parts, not only in this, but, as I apprehend in every natural family, is very unequal, and in some cases seems to be entirely lost." Again, in another work he says, the genera of the Connaraceae "differ in having one or more ovaria, in the existence or absence of albumen, in the imbricate or valvular aestivation. Any one of these characters singly is frequently of more than generic importance, though here even, when all taken together, they appear insufficient to separate Cnestis from Connarus." To give an example among insects: in one great division of the Hymenoptera, the antennae, as Westwood has remarked, are most constant in structure; in another division they differ much, and the differences are of quite subordinate value in classification; yet no one will say that the antennae in these two divisions of the same order are of unequal physiological importance. Any number of instances could be given of the varying importance for classification of the same important organ within the same group of beings.

Again, no one will say that rudimentary or atrophied organs are of high physiological or vital importance; yet, undoubtedly, organs in this condition are often of much value in classification. No one will dispute that the rudimentary teeth in the upper jaws of young ruminants, and certain rudimentary bones of the leg, are highly serviceable in exhibiting the close affinity between Ruminants and Pachyderms. Robert Brown has strongly insisted on the fact that the position of the rudimentary florets is of the highest importance in the classification of the Grasses.

Numerous instances could be given of characters derived from parts which must be considered of very trifling physiological importance, but which are universally admitted as highly serviceable in the definition of whole groups. For instance, whether or not there is an open passage from the nostrils to the mouth, the only character, according to Owen, which absolutely distinguishes fishes and reptiles – the inflection of the angle of the lower jaw in Marsupials – the manner in which the wings of insects are folded – mere colour in certain Algae – mere pubescence on parts of the flower in grasses – the nature of the dermal covering, as hair or feathers, in the Vertebrata. If the Ornithorhynchus had been covered with feathers instead of hair, this external and trifling character would have been considered by naturalists as an important aid in determining the degree of affinity of this strange creature to birds.

The importance, for classification, of trifling characters, mainly depends on their being correlated with many other characters of more or less importance. The value indeed of an aggregate of characters is very evident in natural history. Hence, as has often been remarked, a species may depart from its allies in several characters, both of high physiological importance, and of almost universal prevalence, and yet leave us in no doubt where it should be ranked. Hence, also, it has been found that a classification founded on any single character, however important that may be, has always failed; for no part of the organisation is invariably constant. The importance of an aggregate of characters, even when none are important, alone explains the aphorism enunciated by Linnaeus, namely, that the characters do not give the genus, but the genus gives the character; for this seems founded on the appreciation of many trifling points of resemblance, too slight to be defined. Certain plants, belonging to the Malpighiaceae, bear perfect and degraded flowers; in the latter, as A. de Jussieu has remarked, "The greater number of the characters proper to the species, to the genus, to the family, to the class, disappear, and thus laugh at our classification." When Aspicarpa produced in France, during several years, only these degraded flowers, departing so wonderfully in a number of the most important points of structure from the proper type of the order, yet M. Richard sagaciously saw, as Jussieu observes, that this genus should still be retained among the Malpighiaceae. This case well illustrates the spirit of our classifications.

Practically, when naturalists are at work, they do not trouble themselves about the physiological value of the characters which they use in defining a group or in allocating any particular species. If they find a character nearly uniform, and common to a great number of forms, and not common to others, they use it as one of high value; if common to some lesser number, they use it as of subordinate value. This principle has been broadly confessed by some naturalists to be the true one; and by none more clearly than by that excellent botanist, Aug. St. Hilaire. If several trifling characters are always found in combination, though no apparent bond of connexion can be discovered between them, especial value is set on them. As in most groups of animals, important organs, such as those for propelling the blood, or for aerating it, or those for propagating the race, are found nearly uniform, they are considered as highly serviceable in classification; but in some groups all these, the most important vital organs, are found to offer characters of quite subordinate value. Thus, as Fritz Muller has lately remarked, in the same group of crustaceans, Cypridina is furnished with a heart, while in two closely allied genera, namely Cypris and Cytherea, there is no such organ; one species of Cypridina has well-developed branchiae, while another species is destitute of them.

We can see why characters derived from the embryo should be of equal importance with those derived from the adult, for a natural classification of course includes all ages. But it is by no means obvious, on the ordinary view, why the structure of the embryo should be more important for this purpose than that of the adult, which alone plays its full part in the economy of nature. Yet it has been strongly urged by those great naturalists, Milne Edwards and Agassiz, that embryological characters are the most important of all; and this doctrine has very generally been admitted as true. Nevertheless, their importance has sometimes been exaggerated, owing to the adaptive characters of larvae not having been excluded; in order to show this, Fritz Muller arranged, by the aid of such characters alone, the great class of crustaceans, and the arrangement did not prove a natural one. But there can be no doubt that embryonic, excluding larval characters, are of the highest value for classification, not only with animals but with plants. Thus the main divisions of flowering plants are founded on differences in the embryo – on the number and position of the cotyledons, and on the mode of development of the plumule and radicle. We shall immediately see why these characters possess so high a value in classification, namely, from the natural system being genealogical in its arrangement.

Our classifications are often plainly influenced by chains of affinities. Nothing can be easier than to define a number of characters common to all birds; but with crustaceans, any such definition has hitherto been found impossible. There are crustaceans at the opposite ends of the series, which have hardly a character in common; yet the species at both ends, from being plainly allied to others, and these to others, and so onwards, can be recognised as unequivocally belonging to this, and to no other class of the Articulata.

Geographical distribution has often been used, though perhaps not quite logically, in classification, more especially in very large groups of closely allied forms. Temminck insists on the utility or even necessity of this practice in certain groups of birds; and it has been followed by several entomologists and botanists.

Finally, with respect to the comparative value of the various groups of species, such as orders, suborders, families, subfamilies, and genera, they seem to be, at least at present, almost arbitrary. Several of the best botanists, such as Mr. Bentham and others, have strongly insisted on their arbitrary value. Instances could be given among plants and insects, of a group first ranked by practised naturalists as only a genus, and then raised to the rank of a subfamily or family; and this has been done, not because further research has detected important structural differences, at first overlooked, but because numerous allied species, with slightly different grades of difference, have been subsequently discovered.

All the foregoing rules and aids and difficulties in classification may be explained, if I do not greatly deceive myself, on the view that the natural system is founded on descent with modification – that the characters which naturalists consider as showing true affinity between any two or more species, are those which have been inherited from a common parent, all true classification being genealogical – that community of descent is the hidden bond which naturalists have been unconsciously seeking, and not some unknown plan of creation, or the enunciation of general propositions, and the mere putting together and separating objects more or less alike.

But I must explain my meaning more fully. I believe that the ARRANGEMENT of the groups within each class, in due subordination and relation to each other, must be strictly genealogical in order to be natural; but that the AMOUNT of difference in the several branches or groups, though allied in the same degree in blood to their common progenitor, may differ greatly, being due to the different degrees of modification which they have undergone; and this is expressed by the forms being ranked under different genera, families, sections or orders. The reader will best understand what is meant, if he will take the trouble to refer to the diagram in the fourth chapter. We will suppose the letters A to L to represent allied genera existing during the Silurian epoch, and descended from some still earlier form. In three of these genera (A, F, and I) a species has transmitted modified descendants to the present day, represented by the fifteen genera (a14 to z14) on the uppermost horizontal line. Now, all these modified descendants from a single species are related in blood or descent in the same degree. They may metaphorically be called cousins to the same millionth degree, yet they differ widely and in different degrees from each other. The forms descended from A, now broken up into two or three families, constitute a distinct order from those descended from I, also broken up into two families. Nor can the existing species descended from A be ranked in the same genus with the parent A, or those from I with parent I. But the existing genus F14 may be supposed to have been but slightly modified, and it will then rank with the parent genus F; just as some few still living organisms belong to Silurian genera. So that the comparative value of the differences between these organic beings, which are all related to each other in the same degree in blood, has come to be widely different. Nevertheless, their genealogical ARRANGEMENT remains strictly true, not only at the present time, but at each successive period of descent. All the modified descendants from A will have inherited something in common from their common parent, as will all the descendants from I; so will it be with each subordinate branch of descendants at each successive stage. If, however, we suppose any descendant of A or of I to have become so much modified as to have lost all traces of its parentage in this case, its place in the natural system will be lost, as seems to have occurred with some few existing organisms. All the descendants of the genus F, along its whole line of descent, are supposed to have been but little modified, and they form a single genus. But this genus, though much isolated, will still occupy its proper intermediate position. The representation of the groups as here given in the diagram on a flat surface, is much too simple. The branches ought to have diverged in all directions. If the names of the groups had been simply written down in a linear series the representation would have been still less natural; and it is notoriously not possible to represent in a series, on a flat surface, the affinities which we discover in nature among the beings of the same group. Thus, the natural system is genealogical in its arrangement, like a pedigree. But the amount of modification which the different groups have undergone has to be expressed by ranking them under different so-called genera, subfamilies, families, sections, orders, and classes.

It may be worth while to illustrate this view of classification, by taking the case of languages. If we possessed a perfect pedigree of mankind, a genealogical arrangement of the races of man would afford the best classification of the various languages now spoken throughout the world; and if all extinct languages, and all intermediate and slowly changing dialects, were to be included, such an arrangement would be the only possible one. Yet it might be that some ancient languages had altered very little and had given rise to few new languages, whilst others had altered much owing to the spreading, isolation and state of civilisation of the several co-descended races, and had thus given rise to many new dialects and languages. The various degrees of difference between the languages of the same stock would have to be expressed by groups subordinate to groups; but the proper or even the only possible arrangement would still be genealogical; and this would be strictly natural, as it would connect together all languages, extinct and recent, by the closest affinities, and would give the filiation and origin of each tongue.

In confirmation of this view, let us glance at the classification of varieties, which are known or believed to be descended from a single species. These are grouped under the species, with the subvarieties under the varieties; and in some cases, as with the domestic pigeon, with several other grades of difference. Nearly the same rules are followed as in classifying species. Authors have insisted on the necessity of arranging varieties on a natural instead of an artificial system; we are cautioned, for instance, not to class two varieties of the pine-apple together, merely because their fruit, though the most important part, happens to be nearly identical; no one puts the Swedish and common turnip together, though the esculent and thickened stems are so similar. Whatever part is found to be most constant, is used in classing varieties: thus the great agriculturist Marshall says the horns are very useful for this purpose with cattle, because they are less variable than the shape or colour of the body, etc.; whereas with sheep the horns are much less serviceable, because less constant. In classing varieties, I apprehend that if we had a real pedigree, a genealogical classification would be universally preferred; and it has been attempted in some cases. For we might feel sure, whether there had been more or less modification, that the principle of inheritance would keep the forms together which were allied in the greatest number of points. In tumbler pigeons, though some of the subvarieties differ in the important character of the length of the beak, yet all are kept together from having the common habit of tumbling; but the short-faced breed has nearly or quite lost this habit; nevertheless, without any thought on the subject, these tumblers are kept in the same group, because allied in blood and alike in some other respects.

With species in a state of nature, every naturalist has in fact brought descent into his classification; for he includes in his lowest grade, that of species, the two sexes; and how enormously these sometimes differ in the most important characters is known to every naturalist: scarcely a single fact can be predicated in common of the adult males and hermaphrodites of certain cirripedes, and yet no one dreams of separating them. As soon as the three Orchidean forms, Monachanthus, Myanthus, and Catasetum, which had previously been ranked as three distinct genera, were known to be sometimes produced on the same plant, they were immediately considered as varieties; and now I have been able to show that they are the male, female, and hermaphrodite forms of the same species. The naturalist includes as one species the various larval stages of the same individual, however much they may differ from each other and from the adult; as well as the so-called alternate generations of Steenstrup, which can only in a technical sense be considered as the same individual. He includes monsters and varieties, not from their partial resemblance to the parent-form, but because they are descended from it.

As descent has universally been used in classing together the individuals of the same species, though the males and females and larvae are sometimes extremely different; and as it has been used in classing varieties which have undergone a certain, and sometimes a considerable amount of modification, may not this same element of descent have been unconsciously used in grouping species under genera, and genera under higher groups, all under the so-called natural system? I believe it has been unconsciously used; and thus only can I understand the several rules and guides which have been followed by our best systematists. As we have no written pedigrees, we are forced to trace community of descent by resemblances of any kind. Therefore, we choose those characters which are the least likely to have been modified, in relation to the conditions of life to which each species has been recently exposed. Rudimentary structures on this view are as good as, or even sometimes better than other parts of the organisation. We care not how trifling a character may be – let it be the mere inflection of the angle of the jaw, the manner in which an insect's wing is folded, whether the skin be covered by hair or feathers – if it prevail throughout many and different species, especially those having very different habits of life, it assumes high value; for we can account for its presence in so many forms with such different habits, only by inheritance from a common parent. We may err in this respect in regard to single points of structure, but when several characters, let them be ever so trifling, concur throughout a large group of beings having different habits, we may feel almost sure, on the theory of descent, that these characters have been inherited from a common ancestor; and we know that such aggregated characters have especial value in classification.

We can understand why a species or a group of species may depart from its allies, in several of its most important characteristics, and yet be safely classed with them. This may be safely done, and is often done, as long as a sufficient number of characters, let them be ever so unimportant, betrays the hidden bond of community of descent. Let two forms have not a single character in common, yet, if these extreme forms are connected together by a chain of intermediate groups, we may at once infer their community of descent, and we put them all into the same class. As we find organs of high physiological importance – those which serve to preserve life under the most diverse conditions of existence – are generally the most constant, we attach especial value to them; but if these same organs, in another group or section of a group, are found to differ much, we at once value them less in our classification. We shall presently see why embryological characters are of such high classificatory importance. Geographical distribution may sometimes be brought usefully into play in classing large genera, because all the species of the same genus, inhabiting any distinct and isolated region, are in all probability descended from the same parents.

ANALOGICAL RESEMBLANCES.

We can understand, on the above views, the very important distinction between real affinities and analogical or adaptive resemblances. Lamarck first called attention to this subject, and he has been ably followed by Macleay and others. The resemblance in the shape of the body and in the fin-like anterior limbs between dugongs and whales, and between these two orders of mammals and fishes, are analogical. So is the resemblance between a mouse and a shrew-mouse (Sorex), which belong to different orders; and the still closer resemblance, insisted on by Mr. Mivart, between the mouse and a small marsupial animal (Antechinus) of Australia. These latter resemblances may be accounted for, as it seems to me, by adaptation for similarly active movements through thickets and herbage, together with concealment from enemies.

Among insects there are innumerable instances; thus Linnaeus, misled by external appearances, actually classed an homopterous insect as a moth. We see something of the same kind even with our domestic varieties, as in the strikingly similar shape of the body in the improved breeds of the Chinese and common pig, which are descended from distinct species; and in the similarly thickened stems of the common and specifically distinct Swedish turnip. The resemblance between the greyhound and race-horse is hardly more fanciful than the analogies which have been drawn by some authors between widely different animals.

On the view of characters being of real importance for classification, only in so far as they reveal descent, we can clearly understand why analogical or adaptive characters, although of the utmost importance to the welfare of the being, are almost valueless to the systematist. For animals, belonging to two most distinct lines of descent, may have become adapted to similar conditions, and thus have assumed a close external resemblance; but such resemblances will not reveal – will rather tend to conceal their blood-relationship. We can thus also understand the apparent paradox, that the very same characters are analogical when one group is compared with another, but give true affinities when the members of the same group are compared together: thus the shape of the body and fin-like limbs are only analogical when whales are compared with fishes, being adaptations in both classes for swimming through the water; but between the the several members of the whale family, the shape of the body and the fin-like limbs offer characters exhibiting true affinity; for as these parts are so nearly similar throughout the whole family, we cannot doubt that they have been inherited from a common ancestor. So it is with fishes.

Numerous cases could be given of striking resemblances in quite distinct beings between single parts or organs, which have been adapted for the same functions. A good instance is afforded by the close resemblance of the jaws of the dog and Tasmanian wolf or Thylacinus – animals which are widely sundered in the natural system. But this resemblance is confined to general appearance, as in the prominence of the canines, and in the cutting shape of the molar teeth. For the teeth really differ much: thus the dog has on each side of the upper jaw four pre-molars and only two molars; while the Thylacinus has three pre-molars and four molars. The molars also differ much in the two animals in relative size and structure. The adult dentition is preceded by a widely different milk dentition. Any one may, of course, deny that the teeth in either case have been adapted for tearing flesh, through the natural selection of successive variations; but if this be admitted in the one case, it is unintelligible to me that it should be denied in the other. I am glad to find that so high an authority as Professor Flower has come to this same conclusion.

The extraordinary cases given in a former chapter, of widely different fishes possessing electric organs – of widely different insects possessing luminous organs – and of orchids and asclepiads having pollen-masses with viscid discs, come under this same head of analogical resemblances. But these cases are so wonderful that they were introduced as difficulties or objections to our theory. In all such cases some fundamental difference in the growth or development of the parts, and generally in their matured structure, can be detected. The end gained is the same, but the means, though appearing superficially to be the same, are essentially different. The principle formerly alluded to under the term of ANALOGICAL VARIATION has probably in these cases often come into play; that is, the members of the same class, although only distantly allied, have inherited so much in common in their constitution, that they are apt to vary under similar exciting causes in a similar manner; and this would obviously aid in the acquirement through natural selection of parts or organs, strikingly like each other, independently of their direct inheritance from a common progenitor.

As species belonging to distinct classes have often been adapted by successive slight modifications to live under nearly similar circumstances – to inhabit, for instance, the three elements of land, air and water – we can perhaps understand how it is that a numerical parallelism has sometimes been observed between the subgroups of distinct classes. A naturalist, struck with a parallelism of this nature, by arbitrarily raising or sinking the value of the groups in several classes (and all our experience shows that their valuation is as yet arbitrary), could easily extend the parallelism over a wide range; and thus the septenary, quinary, quaternary and ternary classifications have probably arisen.

There is another and curious class of cases in which close external resemblance does not depend on adaptation to similar habits of life, but has been gained for the sake of protection. I allude to the wonderful manner in which certain butterflies imitate, as first described by Mr. Bates, other and quite distinct species. This excellent observer has shown that in some districts of South America, where, for instance, an Ithomia abounds in gaudy swarms, another butterfly, namely, a Leptalis, is often found mingled in the same flock; and the latter so closely resembles the Ithomia in every shade and stripe of colour, and even in the shape of its wings, that Mr. Bates, with his eyes sharpened by collecting during eleven years, was, though always on his guard, continually deceived. When the mockers and the mocked are caught and compared, they are found to be very different in essential structure, and to belong not only to distinct genera, but often to distinct families. Had this mimicry occurred in only one or two instances, it might have been passed over as a strange coincidence. But, if we proceed from a district where one Leptalis imitates an Ithomia, another mocking and mocked species, belonging to the same two genera, equally close in their resemblance, may be found. Altogether no less than ten genera are enumerated, which include species that imitate other butterflies. The mockers and mocked always inhabit the same region; we never find an imitator living remote from the form which it imitates. The mockers are almost invariably rare insects; the mocked in almost every case abounds in swarms. In the same district in which a species of Leptalis closely imitates an Ithomia, there are sometimes other Lepidoptera mimicking the same Ithomia: so that in the same place, species of three genera of butterflies and even a moth are found all closely resembling a butterfly belonging to a fourth genus. It deserves especial notice that many of the mimicking forms of the Leptalis, as well as of the mimicked forms, can be shown by a graduated series to be merely varieties of the same species; while others are undoubtedly distinct species. But why, it may be asked, are certain forms treated as the mimicked and others as the mimickers? Mr. Bates satisfactorily answers this question by showing that the form which is imitated keeps the usual dress of the group to which it belongs, while the counterfeiters have changed their dress and do not resemble their nearest allies.

We are next led to enquire what reason can be assigned for certain butterflies and moths so often assuming the dress of another and quite distinct form; why, to the perplexity of naturalists, has nature condescended to the tricks of the stage? Mr. Bates has, no doubt, hit on the true explanation. The mocked forms, which always abound in numbers, must habitually escape destruction to a large extent, otherwise they could not exist in such swarms; and a large amount of evidence has now been collected, showing that they are distasteful to birds and other insect-devouring animals. The mocking forms, on the other hand, that inhabit the same district, are comparatively rare, and belong to rare groups; hence, they must suffer habitually from some danger, for otherwise, from the number of eggs laid by all butterflies, they would in three or four generations swarm over the whole country. Now if a member of one of these persecuted and rare groups were to assume a dress so like that of a well-protected species that it continually deceived the practised eyes of an entomologist, it would often deceive predaceous birds and insects, and thus often escape destruction. Mr. Bates may almost be said to have actually witnessed the process by which the mimickers have come so closely to resemble the mimicked; for he found that some of the forms of Leptalis which mimic so many other butterflies, varied in an extreme degree. In one district several varieties occurred, and of these one alone resembled, to a certain extent, the common Ithomia of the same district. In another district there were two or three varieties, one of which was much commoner than the others, and this closely mocked another form of Ithomia. From facts of this nature, Mr. Bates concludes that the Leptalis first varies; and when a variety happens to resemble in some degree any common butterfly inhabiting the same district, this variety, from its resemblance to a flourishing and little persecuted kind, has a better chance of escaping destruction from predaceous birds and insects, and is consequently oftener preserved; "the less perfect degrees of resemblance being generation after generation eliminated, and only the others left to propagate their kind." So that here we have an excellent illustration of natural selection.

Messrs. Wallace and Trimen have likewise described several equally striking cases of imitation in the Lepidoptera of the Malay Archipelago and Africa, and with some other insects. Mr. Wallace has also detected one such case with birds, but we have none with the larger quadrupeds. The much greater frequency of imitation with insects than with other animals, is probably the consequence of their small size; insects cannot defend themselves, excepting indeed the kinds furnished with a sting, and I have never heard of an instance of such kinds mocking other insects, though they are mocked; insects cannot easily escape by flight from the larger animals which prey on them; therefore, speaking metaphorically, they are reduced, like most weak creatures, to trickery and dissimulation.

It should be observed that the process of imitation probably never commenced between forms widely dissimilar in colour. But, starting with species already somewhat like each other, the closest resemblance, if beneficial, could readily be gained by the above means, and if the imitated form was subsequently and gradually modified through any agency, the imitating form would be led along the same track, and thus be altered to almost any extent, so that it might ultimately assume an appearance or colouring wholly unlike that of the other members of the family to which it belonged. There is, however, some difficulty on this head, for it is necessary to suppose in some cases that ancient members belonging to several distinct groups, before they had diverged to their present extent, accidentally resembled a member of another and protected group in a sufficient degree to afford some slight protection, this having given the basis for the subsequent acquisition of the most perfect resemblance.

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.