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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 keep in 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 average number of any animal or plant depends only indirectly on the number of its eggs or seeds.
In looking at Nature, it is most necessary to keep the foregoing considerations always in mind—never to forget that every single organic being may be said to be striving to the utmost to increase in numbers; that each lives by a struggle at some period of its life; that heavy destruction inevitably falls either on the young or old, during each generation or at recurrent intervals. Lighten any check, mitigate the destruction ever so little, and the number of the species will almost instantaneously increase to any amount.
Nature of the Chccks to Increase.
The causes which check the natural tendency of cash species to increase are most obscure. Look at the most vigorous species; by as much as it swarms in numbers, by so much will it tend to increase still further. We know not exactly what the checks are even in a single instance. Nor will this surprise any one who reflects how ignorant we are on this head, even in regard to mankind, although so incomparably better known than any other animal. This subject of the checks to increase has been ably treated by several authors, and I hope in a future work to discuss it at considerable length, more especially in regard to the feral animals of South America. Here I will make only a few remarks, just to recall to the reader's mind some of the chief points. Eggs or very young animals seem generally to suffer most, but this is not invariably the case. With plants there is a vast destruction of seeds, but, from some observations which I have made it appears that the seedlings suffer most from germinating in ground already thickly stocked with other plants. Seedlings, also, are destroyed in vast numbers by various enemies; for instance, on a piece of ground three feet long and two wide, dug and cleared, and where there could be no choking from other plants, I marked all the seedlings of our native weeds as they came up, and out of 357 no less than 295 were destroyed, chiefly by slugs and insects. If turf which has long been mown, and the case would be the same with turf closely browsed by quadrupeds, be let to grow, the more vigorous plants gradually kill the less vigorous, though fully grown plants; thus out of twenty species growing on a little plot of mown turf (three feet by four) nine species perished, from the other species being allowed to grow up freely.
The amount of food for each species of course gives the extreme limit to which each can increase; but very frequently it is not the obtaining food, but the serving as prey to other animals, which determines the average numbers of a species. Thus, there seems to be little doubt that the stock of partridges, grouse, and hares on any large estate depends chiefly on the destruction of vermin. If not one head of game were shot during the next twenty years in England, and, at the same time, if no vermin were destroyed, there would, in all probability, be less game than at present, although hundreds of thousands of game animals are now annually shot. On the other hand, in some cases, as with the elephant, none are destroyed by beasts of prey; for even the tiger in India most rarely dares to attack a young elephant protected by its dam.
Climate plays an important part in determining the average numbers of a species, and periodical seasons of extreme cold or drought seem to be the most effective of all checks. I estimated (chiefly from the greatly reduced numbers of nests in the spring) that the winter of 1854-5 destroyed four-fifths of the birds in my own grounds; and this is a tremendous destruction, when we remember that ten per cent, is an extraordinarily severe mortality from epidemics with man. The action of climate seems at first sight to be quite independent of the struggle for existence; but in so far as climate chiefly acts in reducing food, it bring on the most severe struggle between the individuals, whether of the same or of distinct species, which