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amount of difference considered necessary to give to any two forms the rank of species cannot be defined. In genera having more than the average number of species in any country, the species of these genera have more than the average number of varieties. In large genera the species are apt to be closely, but unequally, allied together, forming little clusters round other species. Species very closely allied to other species apparently have restricted ranges. In all these respects the species of large genera present a strong analogy with varieties. And we can clearly understand these analogies, if species once existed as varieties, and thus originated: whereas, these analogies are utterly inexplicable if species are independent creations.
We have, also, seen that it is the most flourishing or dominant species of the larger genera within each class which on an average yield the greatest number of varieties; and varieties, as we shall hereafter see, tend to become converted into new and distinct species. Thus the larger genera tend to become larger ; and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. But by steps hereafter to be explained, the larger genera also tend to break up into smaller genera. And thus, the forms of life throughout the universe become divided into groups subordinate to groups.
Its bearing on natural selection—The term used in a wide sense—Geometrical ratio of increase–Rapid increase of naturalised animals and plants— Nature of the checks to increase—Competition universal–Effects of climate—Protection from the number of individuals—Complex relations of all animals and plants throughout nature—Struggle for life most severe between individuals and varieties of the same species: often severe between species of the same genus—The relation of organism to organism the most important of all relations.
BEFoRE entering on the subject of this chapter, I must make a few preliminary remarks, to show how the struggle for existence bears on Natural Selection. It has been seen in the last chapter that amongst organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties
be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life, and of one organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water: in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world. Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species 7 How do those groups of species, which constitute what are called distinct genera, and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow from the struggle for life. Owing to this struggle, variations, however slight and from whatever cause proceeding, if they be in any degree profitable to the individuals of a species, in their infinitely complex relations to other organic beings and to their physical conditions of life, will tend to the preservation of such individuals, and will generally be inherited by the offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term Natural Selection, in order to mark its relation to man's power of selection. But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man's feeble efforts, as the works of Nature are to those of Art. We will now discuss in a little more detail the struggle for existence. In my future work this subject will be treated, as it well deserves, at greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no
one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult—at least I have found it so—than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladmess, we often see superabundance of food; we do not see or we forget, that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that, though food may be now superabundant, it is not so at all seasons of each recurring year.
The Term, Struggle for Existence, used in a large sense.
I should premise that I use this term in a large and metaphorical sense including dependence of one being on another, and including (which is more important) not only the life of the individual, but success in leaving progeny. Two canine animals, in a time of dearth, may be truly said to struggle with each other which shall get food and live. But a plant on the edge of a desert is said to struggle for life against the drought, though more properly it should be said to be dependent on the moisture. A plant which annually produces a thousand seeds, of which only one of an average comes to maturity, may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. The mistletoe is dependent on the apple and a few other trees, but can only in a far-fetched sense be said to struggle with these trees, for, if too many of these parasites grow on the same tree, it languishes and dies. But several seedling mistletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the mistletoe is disseminated by birds, its existence depends on them ; and it may metaphorically be said to struggle with other fruit-bearing plants, in tempting the birds to devour and thus disseminate its seeds. In these several senses, which pass into each other, I use for convenience’ sake the general term of Struggle for Existence.
Geometrical Ratio of Increase.
A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season or occasional year, otherwise, on the principle of geometrical increase, its numbers would quickly become so inordinately great that no country could support the product. Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence, either one individual with another of the same species, or with the individuals of distinct species, or with the physical conditions of life. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms; for in this case there can be no artificial increase of food, and no prudential restraint from marriage. Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them. There is no exception to the rule that every organic being naturally increases at so high a rate, that, if not destroyed, the earth would soon be covered by the progeny of a single pair. Even slow-breeding man has doubled in twenty-five years, and at this rate, in less than a thousand years, there would literally not be standing-room for his progeny. Linnaeus has calculated that if an annual plant produced only two seeds—and there is no plant so unproductive as this—and their seedlings next year produced two, and so on, then in twenty years there would be a million plants. The elephant is reckoned the slowest breeder of all known animals, and I have taken some pains to estimate its probable minimum rate of natural increase; it will be safest to assume that it begins breeding when thirty years old, and goes on breeding till ninety years old, bringing forth six young in the interval, and surviving till one hundred years old; if this be so, after a period of from 740 to 750 years there would be nearly nineteen million elephants alive, descended from the first pair. But we have better evidence on this subject than mere theoretical calculations, namely, the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature, when circumstances have been favourable to them during two or three following seasons. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world; if the statements of the rate of increase of slow-breeding cattle and horses in South America, and latterly in Australia, had not been well authenticated, they would have been incredible. So it is with plants; cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. Several of the plants, such as the cardoon and a tall thistle, which are now the commonest over the wide plains of La Plata, clothing square leagues of surface almost to the exclusion of every other plant, have been introduced from Europe; and there are plants which now range in India, as I hear from Dr. Falconer, from Cape Comorin to the Himalaya, which have been imported from America since its discovery. In such cases, and endless others could be given, no one supposes, that the fertility of the animals or plants has been suddenly and temporarily increased in any sensible degree. The obvious explanation is that the conditions of life have been highly favourable, and that there has consequently been less destruction of the old and young, and that nearly all the young have been enabled to breed. Their geometrical ratio of increase, the result of which never fails to be surprising, simply explains their extraordinarily rapid increase and wide diffusion in their new homes. In a state of nature almost every full-grown plant annually produces seed, and amongst animals there are very few which do not annually pair. Hence we may confidently assert, that all plants and animals are tending to increase at a geometrical ratio, that all would rapidly stock every station in which they could anyhow exist,-and that this geometrical tendency to increase must be checked by destruction at some period of life. Our familiarity with the larger domestic animals tends, I think, to mislead us: we see no great destruction falling on them, but we do not keepin mind that thousands are annually slaughtered for food, and that in a state of nature an equal number would have somehow to be disposed of. The only difference between organisms which annually produce eggs or seeds by the thousand, and those which produce extremely few, is, that the slow-breeders would require a few more years to people, under favourable conditions, a whole district, let it be ever so large. The condor lays a couple of eggs and the ostrich a score, and yet in the same country the condor may be the more numerous of the two; the Fulmar petrel lays but one egg, yet it is believed to be the most numerous bird in the world. One fly deposits hundreds of eggs, and another, like the hippobosca, a single one; but this difference does not determine how many individuals of the two species can be supported in a district. A large number of eggs is of some importance to those species which depend on a fluctuating amount of food, for it allows them rapidly to increase in number. But the real importance of a large number of eggs or seeds is to make up for much destruction at some period of life; and this period in the great majority of cases is an early one. If an animal can in any way protect its own eggs or young, a small number may be produced, and yet the average stock be fully kept up ; but if many eggs or young are destroyed, many must be produced, or the species will become extinct. It would suffice to keep up the full number of a tree, which lived on an average for a thousand years, if a single seed were produced once in a thousand years, supposing that this seed were never destroyed, and could be ensured to germinate in a fitting place. So that, in all cases, the