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there would be no struggle for existence, and, consequently, no natural selection, until many forms had been produced: variations in a single species inhabiting an isolated station might be beneficial, and thus the whole mass of individuals might be modified, or two distinct forms might arise. But, as I remarked towards the close of the Introduction, no one ought to feel surprise at much remaining as yet unexplained on the origin of species, if we make due allowance for our profound ignorance on the mutual relations of the inhabitants of the world at the present time, and still more so during past ages.
Convergence of Character.
Mr. H. C. Watson thinks that I have overrated the importance of divergence of character (in which, however, he apparently believes), and that convergence, as it may be called, has likewise played a part. If two species, belonging to two distinct though allied genera, had both produced a large number of new and divergent forms, it is conceivable that these might approach each other so closely that they would have all to be classed under the same genus; and thus the descendants of two distinct genera would converge into one. But it would in most cases be extremely rash to attribute to convergence a close and general similarity of structure in the modified descendants of widely distinct forms. The shape of a crystal is determined solely by the molecular forces, and it is not surprising that dissimilar substances should sometimes assume the same form; but with organic beings we should bearin mind that the form of each depends on an infinitude of complex relations, namely on the variations which have arisen, these being due to causes far too intricate to be followed out, on the nature of the variations which have been preserved or selected, and this depends on the surrounding physical conditions, and in a still higher degree on the surrounding organisms with which each being has come into competition,-and lastly, on inheritance (in itself a fluctuating element) from innumerable progenitors, all of which have had their forms determined through equally complex relations. It is incredible that the descendants of two organisms, which had originally differed in a marked manner, should ever afterwards converge so closely as to lead to a near approach to identity throughout their whole organisation. If this had occurred, we should meet with the same form, independently of genetic connection, recurring in widely separated geological formations; and the balance of evidence is opposed to any such an admission.
Mr. Watson has also objected that the continued action of natural selection, together with divergence of character, would tend to make an indefinite number of specific forms. As far as mere inorganic conditions are concerned, it seems probable that a sufficient number of species would soon become adapted to all considerable diversities of heat, moisture, &c.; but I fully admit that the mutual relations of organic beings are more important; and as the number of species in any country goes on increasing, the organic conditions of life must become more and more complex. Consequently there seems at first sight no limit to the amount of profitable diversification of structure, and therefore no limit to the number of species which might be produced. We do not know that even the most prolific area is fully stocked with specific forms: at the Cape of Good Hope and in Australia, which support such an astonishing number of species, many European plants have become naturalised. But geology shows us, that from an early part of the tertiary period the number of species of shells, and that from the middle part of this same period the number of mammals, has not greatly or at all increased. What then checks an indefinite increase in the number of species? The amount of life (I do not mean the number of specific forms) supported on an area must have a limit, depending so largely as it does on physical conditions; therefore, if an area be inhabited by very many species, each or nearly each species will be represented by few individuals; and such species will be liable to extermination from accidental fluctuations in the nature of the seasons or in the number of their enemies. The process of extermination in such cases would be rapid, whereas the production of new species must always be slow. Imagine the extreme case of as many species as individuals in England, and the first severe winter or very dry summer would exterminate thousands on thousands of species. Rare species, and each species will become rare if the number of species in any country becomes indefinitely increased, will, on the principle often explained, present within a given period few favourable variations; consequently, the process of giving birth to new specific forms would thus be retarded. When any species becomes very rare, close interbreeding will help to exterminate it; authors have thought that this comes into play in accounting for the deterioration of the Aurochs in Lithuania, of Red Deer in Scotland, and of Bears in Norway, &c. Lastly, and this I am inclined to think is the most important element, a dominant species, which has already beaten many competitors in its own home, will tend to spread and supplant many others. Alph. de Candolle has shown that those species which spread widely, tend generally to spread very widely; consequently, they will tend to supplant and exterminate several species in several areas, and thus check the inordinate increase of specific forms throughout the world. Dr. Hooker has recently shown that in the S.E. corner of Australia, where, apparently, there are many invaders from different quarters of the globe, the endemic Australian species have been greatly reduced in number. How much weight to attribute to these several considerations I will not pretend to say; but conjointly they must limit in each country the tendency to an indefinite augmentation of specific forms.
Summary of Chapter.
If under changing conditions of life organic beings present individual differences in almost every part of their structure, and this cannot be disputed; if there be, owing to their geometrical rate of increase, a severe struggle for life at some age, season, or year, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of life, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, it would be a most extraordinary fact if no variations had ever occurred useful to each being's own welfare, in the same manner as so many variations have occurred useful to man. But if variations useful to any organic being ever do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance, these will tend to produce offspring similarly characterised. This principle of preservation, or the survival of the fittest, I have called Natural Selection. It leads to the improvement of each creature in relation to its organic and inorganic conditions of life; and consequently, in most cases, to what must be regarded as an advance in organisation. Nevertheless, low and simple forms will long endure if well fitted for their simple conditions of life.
Natural selection, on the principle of qualities being inherited at corresponding ages, can modify the egg, seed, or young, as easily as the adult. Amongst many animals, sexual selection will have given its aid to ordinary selection, by assuring to the most vigorous and best adapted males the greatest number of offspring. Sexual selection will also give characters useful to the males alone, in their struggles or rivalry with other males; and these characters will be transmitted to one sex or to both sexes, according to the form of inheritance which prevails.
Whether natural selection has really thus acted in adapting the various forms of life to their several conditions and stations, must be judged by the general tenor and balance of evidence given in the following chapters. But we have already seen how it entails extinction; and how largely extinction has acted in the world's history, geology plainly declares. Natural selection, also, leads to divergence of character; for the more organic beings diverge in structure, habits, and constitution, by so much the more can a large number be supported on the area, of which we see proof by looking to the inhabitants of any small spot, and to the productions naturalised in foreign lands. Therefore, during the modification of the descendants of any one species, and during the incessant struggle of all species to increase in numbers, the more diversified the descendants become, the better will be their chance of success in the battle for life. Thus the small differences distinguishing varieties of the same species, steadily tend to increase, till they equal the greater differences between species of the same genus, or even of distinct genera. We have seen that it is the common, the widely-diffused and widely-ranging species, belonging to the larger genera within each class, which vary most ; and these tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. Natural selection, as has just been remarked, leads to divergence of character and to much extinction of the less improved and intermediate forms of life. On these principles, the nature of the affinities, and the generally well-defined distinctions between the innumerable organic beings in each class throughout the world, may be explained. It is a truly wonderful fact—the wonder of which we are apt to overlook from familiarity—that all animals and all plants throughout all time and space should be related to each other in groups, subordinate to groups, in the manner which we everywhere behold—namely, varieties of the same species most closely related, species of the same genus less closely and unequally related, forming sections and sub-genera, species of distinct genera much less closely related, and genera related in different degrees, forming sub-families, families, orders, sub-classes and classes. The several subordinate groups in any class cannot be ranked in a single file, but seem clustered round points, and these round other points, and so on in almost endless cycles. If species had been independently created, no explanation would have been possible of this kind of classification; but it is explained through inheritance and the complex action of natural selection, entailing extinction and divergence of character, as we have seen illustrated in the diagram. The affinities of all the beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth. The green and budding twigs may represent existing species; and those produced during former years may represent the long succession of extinct species. At each period of growth all the growing twigs have tried to branch out on all sides, and to overtop and kill the surrounding twigs and branches, in the same manner as species and groups of species have at all times overmastered other species in the great battle for life. The H
limbs divided into great branches, and these into lesser and lesser branches, were themselves once, when the tree was young, budding twigs; and this connection of the former and present buds by ramifying branches may well represent the classification of all extinct and living species in groups subordinate to groups. Of the many twigs which flourished when the tree was a mere bush, only two or three, now grown into great branches, yet survive and bear the other branches; so with the species which lived during long-past geological periods, very few have left living and modified descendants. From the first growth of the tree, many a limb and branch has decayed and dropped off; and these fallen branches of various sizes may represent those whole orders, families, and genera which have now no living representatives, and which are known to us only in a fossil state. As we here and there see a thin straggling branch springing from a fork low down in a tree, and which by some chance has been favoured and is still alive on its summit, so we occasionally see an animal like the Ornithorhynchus or Lepidosiren, which in some small degree connects by its affinities two large branches of life, and which has apparently been saved from fatal competition by having inhabited a protected station. As buds give rise by growth to fresh buds, and these, if vigorous, branch out and overtop on all sides many a feebler branch, so by generation I believe it has been with the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever-branching and beautiful ramifications.
Effects of changed conditions—Use and disuse, combined with natural selection; organs of flight and of vision—Acclimatisation—Correlated variation— Compensation and economy of growth—False correlations—Multiple, rudimentary, and lowly organised structures variable—Parts do in an unusual manner are highly variable: specific characters more variable than generic : secondary sexual characters variable—Species of the same enus vary in an analogous manner—Reversions to long-lost characters— so
I HAVE hitherto sometimes spoken as if the variations—so common and multiform with organic beings under domestication, and in a lesser degree with those under nature—were due to chance. This, of course, is a wholly incorrect expression, but it serves to acknow