Bears on natural selection—The term used in a wide sense— Geometrical powers 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 distinct organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and missletoe; 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? 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 inevitably from the struggle for life. Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its 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 of Natural Selection, in order to mark its relation to man's power of selection. 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 shall be treated, as it well deserves, at much 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, I am convinced that 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 gladness, 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.
I should premise that I use the term Struggle for Existence 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 on an average only one 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 missletoe 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 will languish and die. But several seedling missletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the missletoe is disseminated by birds, its existence depends on birds; and it may metaphorically be said to struggle with other fruit-bearing plants, in order to tempt birds to devour and thus disseminate its seeds rather than those of other plants. In these several senses, which pass into each other, I use for convenience sake the general term of struggle for existence.
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 a few 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 to be 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 under the mark to assume that it breeds when thirty years old, and goes on breeding till ninety years old, bringing forth three pair of young in this interval; if this be so, at the end of the fifth century there would be alive fifteen million elephants, 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 quite 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 now most numerous over the wide plains of La Plata, clothing square leagues of surface almost to the exclusion of all other plants, 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 instances could be given, no one supposes that the fertility of these animals or plants has been suddenly and temporarily increased in any sensible degree. The obvious explanation is that the conditions of life have been very 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. In such cases the geometrical ratio of increase, the result of which never fails to be surprising, simply explains the extraordinarily rapid increase and wide diffusion of naturalised productions in their new homes.
In a state of nature almost every plant 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 most rapidly stock every station in which they could any how exist, and that the 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, and we forget 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 rapidly 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 around us 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. The face of Nature may be compared to a yielding surface, with ten thousand sharp wedges packed close together and driven inwards by incessant blows, sometimes one wedge being struck, and then another with greater force.
What checks the natural tendency of each species to increase in number is most obscure. Look at the most vigorous species; by as much as it swarms in numbers, by so much will its tendency to increase be still further increased. We know not exactly what the checks are in even one single instance. Nor will this surprise any one who reflects how ignorant we are on this head, even in regard to mankind, so incomparably better known than any other animal. This subject has been ably treated by several authors, and I shall, in my future work, discuss some of the checks 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, I believe that it is the seedlings which 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 the 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 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 killed. On the other hand, in some cases, as with the elephant and rhinoceros, none are destroyed by beasts of prey: 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, I believe to be the most effective of all checks. I estimated that the winter of 1854-55 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 brings on the most severe struggle between the individuals, whether of the same or of distinct species, which subsist on the same kind of food. Even when climate, for instance extreme cold, acts directly, it will be the least vigorous, or those which have got least food through the advancing winter, which will suffer most. When we travel from south to north, or from a damp region to a dry, we invariably see some species gradually getting rarer and rarer, and finally disappearing; and the change of climate being conspicuous, we are tempted to attribute the whole effect to its direct action. But this is a very false view: we forget that each species, even where it most abounds, is constantly suffering enormous destruction at some period of its life, from enemies or from competitors for the same place and food; and if these enemies or competitors be in the least degree favoured by any slight change of climate, they will increase in numbers, and, as each area is already fully stocked with inhabitants, the other species will decrease. When we travel southward and see a species decreasing in numbers, we may feel sure that the cause lies quite as much in other species being favoured, as in this one being hurt. So it is when we travel northward, but in a somewhat lesser degree, for the number of species of all kinds, and therefore of competitors, decreases northwards; hence in going northward, or in ascending a mountain, we far oftener meet with stunted forms, due to the directly injurious action of climate, than we do in proceeding southwards or in descending a mountain. When we reach the Arctic regions, or snow-capped summits, or absolute deserts, the struggle for life is almost exclusively with the elements.
That climate acts in main part indirectly by favouring other species, we may clearly see in the prodigious number of plants in our gardens which can perfectly well endure our climate, but which never become naturalised, for they cannot compete with our native plants, nor resist destruction by our native animals.
When a species, owing to highly favourable circumstances, increases inordinately in numbers in a small tract, epidemics—at least, this seems generally to occur with our game animals—often ensue: and here we have a limiting check independent of the struggle for life. But even some of these so-called epidemics appear to be due to parasitic worms, which have from some cause, possibly in part through facility of diffusion amongst the crowded animals, been disproportionably favoured: and here comes in a sort of struggle between the parasite and its prey.
On the other hand, in many cases, a large stock of individuals of the same species, relatively to the numbers of its enemies, is absolutely necessary for its preservation. Thus we can easily raise plenty of corn and rape-seed, etc., in our fields, because the seeds are in great excess compared with the number of birds which feed on them; nor can the birds, though having a superabundance of food at this one season, increase in number proportionally to the supply of seed, as their numbers are checked during winter: but any one who has tried, knows how troublesome it is to get seed from a few wheat or other such plants in a garden; I have in this case lost every single seed. This view of the necessity of a large stock of the same species for its preservation, explains, I believe, some singular facts in nature, such as that of very rare plants being sometimes extremely abundant in the few spots where they do occur; and that of some social plants being social, that is, abounding in individuals, even on the extreme confines of their range. For in such cases, we may believe, that a plant could exist only where the conditions of its life were so favourable that many could exist together, and thus save each other from utter destruction. I should add that the good effects of frequent intercrossing, and the ill effects of close interbreeding, probably come into play in some of these cases; but on this intricate subject I will not here enlarge.
Many cases are on record showing how complex and unexpected are the checks and relations between organic beings, which have to struggle together in the same country. I will give only a single instance, which, though a simple one, has interested me. In Staffordshire, on the estate of a relation where I had ample means of investigation, there was a large and extremely barren heath, which had never been touched by the hand of man; but several hundred acres of exactly the same nature had been enclosed twenty-five years previously and planted with Scotch fir. The change in the native vegetation of the planted part of the heath was most remarkable, more than is generally seen in passing from one quite different soil to another: not only the proportional numbers of the heath-plants were wholly changed, but twelve species of plants (not counting grasses and carices) flourished in the plantations, which could not be found on the heath. The effect on the insects must have been still greater, for six insectivorous birds were very common in the plantations, which were not to be seen on the heath; and the heath was frequented by two or three distinct insectivorous birds. Here we see how potent has been the effect of the introduction of a single tree, nothing whatever else having been done, with the exception that the land had been enclosed, so that cattle could not enter. But how important an element enclosure is, I plainly saw near Farnham, in Surrey. Here there are extensive heaths, with a few clumps of old Scotch firs on the distant hill-tops: within the last ten years large spaces have been enclosed, and self-sown firs are now springing up in multitudes, so close together that all cannot live. When I ascertained that these young trees had not been sown or planted, I was so much surprised at their numbers that I went to several points of view, whence I could examine hundreds of acres of the unenclosed heath, and literally I could not see a single Scotch fir, except the old planted clumps. But on looking closely between the stems of the heath, I found a multitude of seedlings and little trees, which had been perpetually browsed down by the cattle. In one square yard, at a point some hundred yards distant from one of the old clumps, I counted thirty-two little trees; and one of them, judging from the rings of growth, had during twenty-six years tried to raise its head above the stems of the heath, and had failed. No wonder that, as soon as the land was enclosed, it became thickly clothed with vigorously growing young firs. Yet the heath was so extremely barren and so extensive that no one would ever have imagined that cattle would have so closely and effectually searched it for food.
Here we see that cattle absolutely determine the existence of the Scotch fir; but in several parts of the world insects determine the existence of cattle. Perhaps Paraguay offers the most curious instance of this; for here neither cattle nor horses nor dogs have ever run wild, though they swarm southward and northward in a feral state; and Azara and Rengger have shown that this is caused by the greater number in Paraguay of a certain fly, which lays its eggs in the navels of these animals when first born. The increase of these flies, numerous as they are, must be habitually checked by some means, probably by birds. Hence, if certain insectivorous birds (whose numbers are probably regulated by hawks or beasts of prey) were to increase in Paraguay, the flies would decrease—then cattle and horses would become feral, and this would certainly greatly alter (as indeed I have observed in parts of South America) the vegetation: this again would largely affect the insects; and this, as we just have seen in Staffordshire, the insectivorous birds, and so onwards in ever-increasing circles of complexity. We began this series by insectivorous birds, and we have ended with them. Not that in nature the relations can ever be as simple as this. Battle within battle must ever be recurring with varying success; and yet in the long-run the forces are so nicely balanced, that the face of nature remains uniform for long periods of time, though assuredly the merest trifle would often give the victory to one organic being over another. Nevertheless so profound is our ignorance, and so high our presumption, that we marvel when we hear of the extinction of an organic being; and as we do not see the cause, we invoke cataclysms to desolate the world, or invent laws on the duration of the forms of life!
I am tempted to give one more instance showing how plants and animals, most remote in the scale of nature, are bound together by a web of complex relations. I shall hereafter have occasion to show that the exotic Lobelia fulgens, in this part of England, is never visited by insects, and consequently, from its peculiar structure, never can set a seed. Many of our orchidaceous plants absolutely require the visits of moths to remove their pollen-masses and thus to fertilise them. I have, also, reason to believe that humble-bees are indispensable to the fertilisation of the heartsease (Viola tricolor), for other bees do not visit this flower. From experiments which I have tried, I have found that the visits of bees, if not indispensable, are at least highly beneficial to the fertilisation of our clovers; but humble-bees alone visit the common red clover (Trifolium pratense), as other bees cannot reach the nectar. Hence I have very little doubt, that if the whole genus of humble-bees became extinct or very rare in England, the heartsease and red clover would become very rare, or wholly disappear. The number of humble-bees in any district depends in a great degree on the number of field-mice, which destroy their combs and nests; and Mr. H. Newman, who has long attended to the habits of humble-bees, believes that “more than two-thirds of them are thus destroyed all over England.” Now the number of mice is largely dependent, as every one knows, on the number of cats; and Mr. Newman says, “Near villages and small towns I have found the nests of humble-bees more numerous than elsewhere, which I attribute to the number of cats that destroy the mice.” Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine, through the intervention first of mice and then of bees, the frequency of certain flowers in that district!
In the case of every species, many different checks, acting at different periods of life, and during different seasons or years, probably come into play; some one check or some few being generally the most potent, but all concurring in determining the average number or even the existence of the species. In some cases it can be shown that widely-different checks act on the same species in different districts. When we look at the plants and bushes clothing an entangled bank, we are tempted to attribute their proportional numbers and kinds to what we call chance. But how false a view is this! Every one has heard that when an American forest is cut down, a very different vegetation springs up; but it has been observed that the trees now growing on the ancient Indian mounds, in the Southern United States, display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. What a struggle between the several kinds of trees must here have gone on during long centuries, each annually scattering its seeds by the thousand; what war between insect and insect—between insects, snails, and other animals with birds and beasts of prey—all striving to increase, and all feeding on each other or on the trees or their seeds and seedlings, or on the other plants which first clothed the ground and thus checked the growth of the trees! Throw up a handful of feathers, and all must fall to the ground according to definite laws; but how simple is this problem compared to the action and reaction of the innumerable plants and animals which have determined, in the course of centuries, the proportional numbers and kinds of trees now growing on the old Indian ruins!
The dependency of one organic being on another, as of a parasite on its prey, lies generally between beings remote in the scale of nature. This is often the case with those which may strictly be said to struggle with each other for existence, as in the case of locusts and grass-feeding quadrupeds. But the struggle almost invariably will be most severe between the individuals of the same species, for they frequent the same districts, require the same food, and are exposed to the same dangers. In the case of varieties of the same species, the struggle will generally be almost equally severe, and we sometimes see the contest soon decided: for instance, if several varieties of wheat be sown together, and the mixed seed be resown, some of the varieties which best suit the soil or climate, or are naturally the most fertile, will beat the others and so yield more seed, and will consequently in a few years quite supplant the other varieties. To keep up a mixed stock of even such extremely close varieties as the variously coloured sweet-peas, they must be each year harvested separately, and the seed then mixed in due proportion, otherwise the weaker kinds will steadily decrease in numbers and disappear. So again with the varieties of sheep: it has been asserted that certain mountain-varieties will starve out other mountain-varieties, so that they cannot be kept together. The same result has followed from keeping together different varieties of the medicinal leech. It may even be doubted whether the varieties of any one of our domestic plants or animals have so exactly the same strength, habits, and constitution, that the original proportions of a mixed stock could be kept up for half a dozen generations, if they were allowed to struggle together, like beings in a state of nature, and if the seed or young were not annually sorted.
As species of the same genus have usually, though by no means invariably, some similarity in habits and constitution, and always in structure, the struggle will generally be more severe between species of the same genus, when they come into competition with each other, than between species of distinct genera. We see this in the recent extension over parts of the United States of one species of swallow having caused the decrease of another species. The recent increase of the missel-thrush in parts of Scotland has caused the decrease of the song-thrush. How frequently we hear of one species of rat taking the place of another species under the most different climates! In Russia the small Asiatic cockroach has everywhere driven before it its great congener. One species of charlock will supplant another, and so in other cases. We can dimly see why the competition should be most severe between allied forms, which fill nearly the same place in the economy of nature; but probably in no one case could we precisely say why one species has been victorious over another in the great battle of life.
A corollary of the highest importance may be deduced from the foregoing remarks, namely, that the structure of every organic being is related, in the most essential yet often hidden manner, to that of all other organic beings, with which it comes into competition for food or residence, or from which it has to escape, or on which it preys. This is obvious in the structure of the teeth and talons of the tiger; and in that of the legs and claws of the parasite which clings to the hair on the tiger's body. But in the beautifully plumed seed of the dandelion, and in the flattened and fringed legs of the water-beetle, the relation seems at first confined to the elements of air and water. Yet the advantage of plumed seeds no doubt stands in the closest relation to the land being already thickly clothed by other plants; so that the seeds may be widely distributed and fall on unoccupied ground. In the water-beetle, the structure of its legs, so well adapted for diving, allows it to compete with other aquatic insects, to hunt for its own prey, and to escape serving as prey to other animals.
The store of nutriment laid up within the seeds of many plants seems at first sight to have no sort of relation to other plants. But from the strong growth of young plants produced from such seeds (as peas and beans), when sown in the midst of long grass, I suspect that the chief use of the nutriment in the seed is to favour the growth of the young seedling, whilst struggling with other plants growing vigorously all around.
Look at a plant in the midst of its range, why does it not double or quadruple its numbers? We know that it can perfectly well withstand a little more heat or cold, dampness or dryness, for elsewhere it ranges into slightly hotter or colder, damper or drier districts. In this case we can clearly see that if we wished in imagination to give the plant the power of increasing in number, we should have to give it some advantage over its competitors, or over the animals which preyed on it. On the confines of its geographical range, a change of constitution with respect to climate would clearly be an advantage to our plant; but we have reason to believe that only a few plants or animals range so far, that they are destroyed by the rigour of the climate alone. Not until we reach the extreme confines of life, in the arctic regions or on the borders of an utter desert, will competition cease. The land may be extremely cold or dry, yet there will be competition between some few species, or between the individuals of the same species, for the warmest or dampest spots.
Hence, also, we can see that when a plant or animal is placed in a new country amongst new competitors, though the climate may be exactly the same as in its former home, yet the conditions of its life will generally be changed in an essential manner. If we wished to increase its average numbers in its new home, we should have to modify it in a different way to what we should have done in its native country; for we should have to give it some advantage over a different set of competitors or enemies.
It is good thus to try in our imagination to give any form some advantage over another. Probably in no single instance should we know what to do, so as to succeed. It will convince us of our ignorance on the mutual relations of all organic beings; a conviction as necessary, as it seems to be difficult to acquire. All that we can do, is to keep steadily in mind that each organic being is striving to increase at a geometrical ratio; that each at some period of its life, during some season of the year, during each generation or at intervals, has to struggle for life, and to suffer great destruction. When we reflect on this struggle, we may console ourselves with the full belief, that the war of nature is not incessant, that no fear is felt, that death is generally prompt, and that the vigorous, the healthy, and the happy survive and multiply.
對(duì)自然選擇的影響——該術(shù)語的廣義——幾何級(jí)數(shù)的增加——?dú)w化動(dòng)植物的迅速增加——抑制增加的性質(zhì)——競爭的普遍性——?dú)夂虻挠绊憽獋€(gè)體數(shù)目的保護(hù)——全體動(dòng)植物在自然界的復(fù)雜關(guān)系——同種的個(gè)體和變種間生存斗爭最劇烈;同屬的物種間也往往劇烈——生物與生物的關(guān)系是一切關(guān)系中最重要的
進(jìn)入本章的主題之前,必須先說幾句開場(chǎng)白,表明生存斗爭對(duì)于“自然選擇”的影響。前一章已經(jīng)談到,自然狀況下的生物是有個(gè)體變異的;這一點(diǎn)從未聽說有爭論。把一群存疑類型叫作物種、亞種或變種,對(duì)于我們無關(guān)緊要。例如,只要承認(rèn)有顯著變種存在,把英國植物中二三百個(gè)存疑類型列入哪一級(jí)都沒有關(guān)系。但是,僅僅有個(gè)體變異和少數(shù)顯著變種的存在,雖然為本書打基礎(chǔ)是必要的,但很少能夠幫我們理解物種在自然狀況下是怎樣發(fā)生的。體制的這一部分對(duì)于另一部分及其對(duì)于生活條件的一切巧妙適應(yīng),生物之間的一切巧妙適應(yīng),是怎樣完善的呢?在啄木鳥和槲寄生身上,我們明顯看到了這種美妙的相互適應(yīng);在依附獸毛、羽毛之上的最下等寄生物上,在潛水甲蟲的構(gòu)造上,在微風(fēng)中飄蕩的冠毛種子上,也差不多同樣明顯;簡而言之,無論何地和生物界的每一部分,都能看到美妙的適應(yīng)。
不妨再問一下,變種即我所謂的初始物種,最終怎樣變成貨真價(jià)實(shí)的物種了呢?在大多數(shù)情形下,物種間的差異,顯然遠(yuǎn)遠(yuǎn)超過了同一物種的變種間的差異。那些組成所謂屬的種群間的差異比同屬物種間的差異為大,這些種群是怎樣發(fā)生的呢?所有這些結(jié)果都不可避免地是從生存斗爭中得來的,下一章將充分論述。由于生存斗爭,變異無論多么輕微,無論由于什么原因發(fā)生,只要任何物種的個(gè)體在與其他生物、與自然界的無限復(fù)雜關(guān)系中多少從中得益,就會(huì)傾向于保存該個(gè)體,并且一般會(huì)讓后代繼承下來。后代也因此有了較好的生存機(jī)會(huì),因?yàn)槿魏挝锓N間歇性產(chǎn)生的許多個(gè)體,只有少數(shù)能夠生存。我把保存每一個(gè)有用的微小變異的這一原則稱為“自然選擇”,表明它和人工選擇力的關(guān)系。我們已經(jīng)看到,人類利用選擇,確能產(chǎn)生偉大的結(jié)果,并且通過累積自然之手所給予的微小而有用的變異,能使生物適合于自己的用途。但是我們以后將看到,自然選擇是一種不斷隨時(shí)激活的力量,它無比地優(yōu)越于微弱的人力。天工無限優(yōu)于人工。
現(xiàn)在就生存斗爭稍加詳論。我以后的著作還要大事討論這個(gè)問題,完全值得討論。老德康多爾和賴爾已經(jīng)富于哲理性地闡明了,一切生物都暴露在劇烈的競爭之中。關(guān)于植物,曼徹斯特區(qū)監(jiān)督牧師赫伯特(W. Herbert)以無人能及的氣魄和才華進(jìn)行了討論,顯然來源于淵博的園藝學(xué)知識(shí)。口頭上承認(rèn)生存斗爭的普遍性,是再容易不過的事情,但至少我認(rèn)為,對(duì)這一結(jié)論要念念不忘卻難上加難。然而,我認(rèn)為,除非在思想上徹底體會(huì)這一點(diǎn),否則我們對(duì)于包含著分布、稀少、繁盛、滅絕以及變異等各種事實(shí)的整個(gè)自然系統(tǒng),就是認(rèn)識(shí)模糊或完全誤解。我們看見自然界的外貌喜氣洋洋,我們常常看見食物過剩,卻看不見或者忘卻安閑地在周圍唱歌的鳥,多數(shù)是以昆蟲或種子為生的,因而經(jīng)常性地在毀滅生命。我們會(huì)忘記這些鳴禽,它們的蛋或幼鳥,會(huì)被猛禽猛獸所大批毀滅。我們并非總是記得,食物雖然現(xiàn)在是過剩的,但并不見得每年的所有季節(jié)都是這樣。
應(yīng)當(dāng)先設(shè)定,術(shù)語生存斗爭采取廣義的比喻義,包含生物的相互依存關(guān)系,更重要的,不僅僅是個(gè)體保命且成功留下后代。兩只狗類動(dòng)物在饑餓的時(shí)候,為了生存爭食,可以說實(shí)實(shí)在在在互相搏斗。但是,生長在沙漠邊緣的植物,可以說是在抗旱求生存,但適當(dāng)?shù)貞?yīng)該說,它是依存于潮氣。一株植物,每年結(jié)一千粒種子,但平均只有一粒能成熟結(jié)籽,可以確切地說,它是在和覆被地面的同類和異類植物做斗爭。槲寄生依存于蘋果樹和少數(shù)其他樹木,但只能牽強(qiáng)附會(huì)地說它在和這些樹木做斗爭,一株樹上這種寄生物過多會(huì)枯死。但是如果幾株槲寄生苗密集地寄生在同一枝條上,就可以實(shí)實(shí)在在地說是在互相斗爭。槲寄生是由鳥類散布的,所以生存便取決于鳥類;可以比喻地說,為了引誘鳥類來吞吃果實(shí)從而散布種子,就是在和其他結(jié)籽植物做斗爭了。在這幾種彼此交叉的意義中,我出于方便,采用了生存斗爭這一通用術(shù)語。
所有生物都有高速增殖的傾向,生存斗爭不可避免。各種生物在其自然壽命中都會(huì)產(chǎn)生若干卵或種子,在生命的某一時(shí)期,某一季節(jié),或者某一年,必定要遭到毀滅,否則按照幾何級(jí)數(shù)增加的原則,數(shù)量就會(huì)很快多得泛濫,沒有地方能夠容納。因此,由于產(chǎn)生的個(gè)體比能生存的多,無論如何一定會(huì)發(fā)生生存斗爭,或者同種個(gè)體之間,或者同異種的個(gè)體斗爭,或者同外界的生活條件斗爭。這是馬爾薩斯學(xué)說成倍地應(yīng)用于整個(gè)的動(dòng)植物界;在這種情形下,既不能人為地增加食物,也不能謹(jǐn)慎地限制交配。雖然某些物種現(xiàn)在可以或快或慢地增加數(shù)目,但是所有的物種并不能這樣,因?yàn)槭澜缛菁{不下它們。
毫無例外,各種生物都自然地高速增殖,如果不加以毀滅,一對(duì)生物的后代很快就會(huì)充滿這個(gè)地球。即使生殖緩慢的人類,也在二十五年間增加了一倍,照此速率類推,幾千年以后,后代就沒有立足之地了。林奈(Linnaeus)計(jì)算過,如果一株一年生的植物只結(jié)兩粒種子(生殖力這樣低的植物是沒有的),幼株翌年也只結(jié)兩粒種子,這樣下去,二十年后就會(huì)有一百萬株了。大象在所有已知的動(dòng)物中可謂是生殖最慢的,我曾費(fèi)力去計(jì)算它在自然增殖方面最小的可能速率;可以保守地假定,它在三十歲開始生育,直到九十歲,在這期間共生三對(duì)小象;如果這樣,五百年以后就會(huì)有一千五百萬只象生存,都是第一對(duì)的后裔。
但是,這個(gè)問題除了理論計(jì)算外,還有更好的證據(jù);大量記載事例表明,自然狀況下的各種動(dòng)物如遇環(huán)境連續(xù)兩三季都適宜的話,便會(huì)神速增殖。還有更觸目驚心的證據(jù),來自世界若干地方已返歸野生狀態(tài)的許多種類的家畜:生育慢的馬和牛在南美洲以及近年來在澳洲的增殖率記錄,若非確有實(shí)據(jù),實(shí)難以置信。植物也是這樣,以外地移入的植物為例,不到十年時(shí)間,就布滿了全島。現(xiàn)在阿根廷拉普拉塔(La Plata)廣大平原上最普通的若干種植物,原來是歐洲引進(jìn)的,可以密布數(shù)里格(1里格=3平方英里)的地面上,幾乎排除了一切他種植物。還有,我聽福爾克納(Falconer)博士說,在美洲發(fā)現(xiàn)后從那里移入到印度的一些植物,已從科摩林角(Cape Comorin)分布到喜馬拉雅了。這些例子真是不勝枚舉。在這些個(gè)案中,沒有人假定這些動(dòng)植物的能育性突如其來暫時(shí)地明顯增加了。解釋不言而喻,生活條件是十分適宜的,結(jié)果,老幼動(dòng)植物的毀滅減少了,幾乎所有新生者都能生育。結(jié)果按幾何級(jí)數(shù)增殖,令人瞠目結(jié)舌,這干脆地說明了歸化動(dòng)植物在新家為什么會(huì)神速增殖和廣泛散布。
自然狀況下,幾乎每一植株都產(chǎn)生種子,而動(dòng)物很少不是每年交配的。因此我們可以斷定,一切動(dòng)植物都有幾何級(jí)數(shù)增殖的傾向,凡是能生存下去的地方,每一處都要迅速滿員,而幾何級(jí)數(shù)增加的傾向必須在生命某一時(shí)期加以毀滅抑制。我想,對(duì)大家畜熟門熟路,會(huì)把我們引入歧途,對(duì)大量毀滅視而不見,也就忘記了每年有成千上萬家畜遭屠宰食用,而且在自然狀況下好歹也得有相等的數(shù)目消滅掉。
生物有每年生產(chǎn)成千上萬枚卵或種子的,也有只生產(chǎn)極少數(shù)卵或種子的,兩者僅有的差別是,生殖慢的生物,在適宜的條件下需要稍稍長一些年限去布滿整個(gè)地區(qū),哪怕地方很大很大。神鷹(condor)產(chǎn)兩三枚卵,鴕鳥(ostrich)產(chǎn)二三十個(gè)卵,然而在同一地區(qū),神鷹可能為數(shù)更多;管鼻鹱(Fulmar petrel)只產(chǎn)一枚卵,但公認(rèn)是世界上最多的鳥。一種蠅產(chǎn)卵成百上千,另一種蠅,如虱蠅(hippobosca)只產(chǎn)一枚卵;但生卵數(shù)量多少,并不能決定兩個(gè)物種在一個(gè)地區(qū)內(nèi)可以養(yǎng)活多少個(gè)體。所依賴的食物大起大落的物種,多產(chǎn)卵是較重要的,因?yàn)榭梢匝杆僭鲋?。但是大量產(chǎn)卵或種子的真正重要性,卻在于補(bǔ)償生命某一階段的大量毀滅;大多數(shù)情況下這個(gè)階段就是初始期。如果一頭動(dòng)物好歹能保護(hù)住卵或幼仔,少量生產(chǎn)仍然能充分保持平均數(shù)量;如果卵或幼仔遭到大量毀滅,就必須大量生產(chǎn),否則物種就要滅絕,假如有一種樹平均能活一千年,哪怕千年產(chǎn)一粒種,假定種子不毀,又能保證在適宜的地方萌發(fā),這就足以保持這種樹的數(shù)目了。所以在所有情況下,任何動(dòng)植物,平均數(shù)目只間接地取決于卵或種子的數(shù)目。
觀察大自然的時(shí)候,千萬記住上述論點(diǎn),千萬不要忘記周圍每一個(gè)生物可以說都在竭力增殖,每一種生物在生命的某一時(shí)期要靠斗爭而生活;千萬不要忘記在每一世代或者間隔幾代,大毀滅不可避免地要降臨幼者或老者。只要少許減輕抑制作用,只要緩和毀滅,物種的數(shù)量幾乎立刻就會(huì)大事增加。大自然的面孔可以比作高產(chǎn)的表面,密密麻麻打入了萬千尖利的楔子,不停地?fù)舸蛳騼?nèi)插,有時(shí)候擊打一根楔子,然后加大力氣擊打另一根。
各個(gè)物種有增殖的自然傾向,其抑制因素極其含糊。看一看最生機(jī)勃勃的物種,其數(shù)量越是密密匝匝,進(jìn)一步增殖的傾向也越強(qiáng)。抑制增殖的因素究竟是什么,我們連一個(gè)事例也弄不明白。這也不足為怪,只要想一想,我們?cè)谶@方面是何等無知,哪怕對(duì)于遠(yuǎn)比任何其他動(dòng)物更了解的人類也是如此。這一主題已有若干作者高論過了,我期望將來在自己的著作里詳論抑制增殖的因素,特別是對(duì)于南美洲的野生動(dòng)物。這里我只稍微談一談,讓讀者注意幾個(gè)要點(diǎn)。卵或幼小動(dòng)物一般看起來受害最多,但不能一概而論。植物的種子被毀的極多,但從我所做的某些觀察得知,在已布滿他種植物的土地上發(fā)芽時(shí),幼苗受害最多。幼苗還會(huì)被各種敵害大量毀滅。例如,有一塊三英尺長二英尺寬的土地,翻耕除草后,不會(huì)再受其他植物的抑制,土著雜草出秧時(shí),我在所有幼苗上做了記號(hào),357株中,不下295株毀滅了,主要是蛞蝓、昆蟲吃掉了。在長期修剪的草皮,四腳獸細(xì)嚼慢咽過的草皮也一樣,如果讓草任意生長,強(qiáng)壯的植物會(huì)逐漸滅掉不強(qiáng)的,哪怕后者已經(jīng)長大。例如在一小塊草皮(三英尺乘四英尺)上生長著二十個(gè)物種,其中九個(gè)物種由于其他物種的自由生長而死亡了。
每個(gè)物種所能吃到的食物數(shù)量,當(dāng)然為各物種的增殖劃了極限;但決定一個(gè)物種的平均數(shù)量,往往不在于獲得食物,而在于他種動(dòng)物的捕食。例如,似乎很少有人懷疑,任何大莊園的鷓鴣、松雞、野兔的數(shù)量主要決定于有害獸的消滅。如果今后二十年中英格蘭不射殺一個(gè)獵物,同時(shí)也不消滅有害獸,那么獵物很有可能比現(xiàn)在還要來得少,雖然現(xiàn)在每年要射殺百十萬只。相反,在某些情形下,例如象和河馬,是不會(huì)被食肉獸捕殺的;在印度甚至老虎也極少敢于攻擊母象保護(hù)下的小象。
決定物種的平均數(shù)量,氣候至關(guān)重要,我認(rèn)為極端寒冷或干旱季節(jié)的不時(shí)出現(xiàn),是最有效的抑制因素。我估算過,1854—1855年冬季,我的居住地消滅的鳥類達(dá)五分之四;這真是重大的毀滅,我們知道,如果人類因傳染病而死去百分之十,便是慘重的死亡率了。氣候的作用乍看似乎同生存斗爭無關(guān),而由于氣候的主要作用在于減少食物,便引發(fā)了同種、異種的個(gè)體間最激烈的斗爭,因?yàn)樗鼈兛客瑯邮澄锷?。哪怕是氣候,例如?yán)寒直接發(fā)生作用時(shí),受害最大的還是最不健壯的個(gè)體,或者入冬后獲得食物最少的個(gè)體。我們從南往北走,或從濕潤地區(qū)到干燥地區(qū),必定會(huì)看出某些物種漸次稀少,最后絕跡。氣候變化顯而易見,我們不免把這整個(gè)的效果歸因于它的直接作用。但這種見解大錯(cuò)特錯(cuò)了,我們忘記了,各個(gè)物種即使在其最繁盛的地方,也經(jīng)常在生命的某一時(shí)期由于敵害或同一地方同一食物的競爭者而大量毀滅。只要?dú)夂蛴休p微變化而稍有利于這些敵害或競爭者,它們便會(huì)增殖;由于各個(gè)地區(qū)都已布滿了生物,其他物種便要減少。我們向南走,如果看見某一物種數(shù)量越來越少,就可以斷定,其原因可以是別的物種受了益,也可以是這個(gè)物種受了損。向北走的情形也是這樣,不過程度稍輕,因?yàn)楦黝惖奈锓N數(shù)量向北去都在減少,所以競爭者也減少了;因此向北走或登山時(shí),往往就比向南走或下山時(shí)見到的植物矮小,這是由于氣候的直接有害作用所致。我們到達(dá)北極區(qū)、積雪的山頂、純粹的沙漠時(shí),生物幾乎單單是同自然環(huán)境進(jìn)行生存斗爭了。
花園里巨大數(shù)量的植物完全能夠忍受我們的氣候,但是永遠(yuǎn)不能歸化,因?yàn)闊o法和土著植物進(jìn)行斗爭,也不能抵抗土著動(dòng)物的侵害。顯而易見,氣候主要是間接起作用,有利于其他物種。
如果一個(gè)物種由于高度適宜的環(huán)境條件在一個(gè)小地域內(nèi)過分增殖了,常常會(huì)引起傳染病的發(fā)生,至少我們的獵物一般是如此。這里的限制性抑制因素同生存斗爭不相干。但是,甚至有些所謂傳染病似乎是由寄生蟲所致,由于某原因,部分地可能是由于動(dòng)物擁擠易于傳播,寄生蟲不對(duì)稱地受益,這里就發(fā)生了某種寄生物和寄主間的斗爭。
另一方面,在許多情形下,面對(duì)敵害,同種個(gè)體絕對(duì)需要大數(shù)量才能保存。例如,我們能輕易地在田間種植大量的五谷和油菜籽等等,因?yàn)榉N子和以此為食的鳥類數(shù)量相比,大為過剩,鳥在這一季里雖然食物異常豐富,卻不能按照種子供給的比例增殖,其數(shù)量在冬季受抑制。人們一試便知,要想從花園里的少量小麥這類植物獲得種子是多么麻煩;我就曾顆粒無收。同種的大群個(gè)體對(duì)于自身保存是必要的,這一觀點(diǎn),我相信可以解釋自然界某些奇特的事實(shí),例如極稀少的植物有時(shí)會(huì)在所生存的少數(shù)地方長得極其繁盛;某些叢生性植物,甚至在分布范圍的邊緣還能叢生,這就是說,個(gè)體是繁盛的。在這種情形下,可以相信,只有在許多個(gè)體能夠共存的有利生活條件下,一種植物才能生存,這樣才能抱團(tuán)互助,免于全部覆滅。我還要補(bǔ)充一句,頻繁雜交的優(yōu)良效果,近親交配的不良效果,也許在這些個(gè)案中起了作用;不過這一問題太復(fù)雜,這里不預(yù)備詳述。
記載下來的很多個(gè)案表明,在同一地方勢(shì)必進(jìn)行斗爭的生物之間的抑制因素和相互關(guān)系,是何等的復(fù)雜和出人意料。我只準(zhǔn)備舉一個(gè)例子,雖然簡單,但我感興趣。我親戚在斯塔福德郡(Staffordshire)有一莊園,我在那里可以進(jìn)行大量的調(diào)查。那里有一大片極度荒蕪的荒地,從來沒有耕種過;但有數(shù)百英畝性質(zhì)完全一致的土地,曾在二十五年前圈了起來,種上了歐洲赤松(Scotch fir)。荒地種植部分的土著植物群落發(fā)生了極顯著的變化,遠(yuǎn)非兩片不同的土壤上可以見到的一般變化程度可比:不但荒地植物的比例數(shù)完全改變了,且有十二個(gè)植物種(不算禾本草類及苔草類)在種植園內(nèi)繁生,而它們根本不見于荒地。對(duì)于昆蟲的影響想必更大些,有六種不見于荒地的食蟲鳥,在種植園內(nèi)很普遍;而經(jīng)常光顧荒地的卻是兩三種食蟲鳥。這里我們看到,只是引進(jìn)一種樹便會(huì)發(fā)生多么大的影響,當(dāng)時(shí)除了把土地圈起來防止牛踏進(jìn)去之外,什么也沒有做。但是,圈地這種要素的重要性,我曾在薩里郡(Surrey)的費(fèi)勒姆(Farnham)鄰近地方清楚地看到了。那里有廣袤的荒地,遠(yuǎn)處小山頂上生長著幾片老齡歐洲赤松。最近十年內(nèi),大塊地方已圈地了,于是自然播種的赤松樹層出不窮,密密麻麻擠著,無法全部存活。當(dāng)我確定這些幼樹并非人工播種或移植,對(duì)于它們的數(shù)量之多大感驚異,于是去了數(shù)處觀測(cè)點(diǎn),觀察了未圈地的數(shù)百英畝荒地,除了舊時(shí)種植的幾叢外,簡直看不到一株歐洲赤松。但在荒地灌木的莖干之間細(xì)察時(shí),我發(fā)現(xiàn)了許多幼苗和小樹不時(shí)被牛吃掉了尖頭。離一片老樹百把碼地方,一平方碼的地上,共計(jì)有三十二株小樹;其中一株,有二十六圈年輪,看來多年來曾試圖把樹頂伸出荒地灌木的樹干之上,但沒有成功。難怪一經(jīng)圈地,便有生氣勃勃的幼齡松樹密布在土地上面了??墒沁@片荒地曾經(jīng)極端荒蕪而且遼闊,沒有人會(huì)想象到牛竟能這樣細(xì)密地來覓食,而且頗有斬獲。
由此可見,牛絕對(duì)決定著歐洲赤松的生存;但在若干地區(qū),昆蟲決定著牛的生存。大概巴拉圭在這方面有最奇異的事例;那里從來沒有牛馬或狗變成野生,但南來北往都有這些動(dòng)物在野生狀態(tài)下成群行動(dòng);亞莎拉(Azara)和倫格(Rengger)闡明,這是由于巴拉圭的某種蠅過多所致,這種蠅就在初生幼畜的臍中產(chǎn)卵。此蠅雖多,但其增殖想必常遇到某種抑制,大概是鳥類吧。因此,如果巴拉圭某種食蟲鳥(其數(shù)量大概受老鷹或猛獸調(diào)節(jié))增多了,蠅就要減少——于是牛馬便可能成為野生的了,而這一定會(huì)使植物群落大為改變(我確在南美洲一些地方看到過這種現(xiàn)象);同時(shí)這又會(huì)大大地影響昆蟲;從而又會(huì)影響食蟲鳥,恰如我們?cè)谒顾5驴に?,如此循環(huán)往復(fù),復(fù)雜關(guān)系不斷擴(kuò)大。這個(gè)系列從食蟲鳥始,又以食蟲鳥終。倒不是自然界里的各種關(guān)系都可以這樣簡單。戰(zhàn)斗之中套著戰(zhàn)斗,必定反復(fù)發(fā)生,成敗無常;盡管區(qū)區(qū)瑣事往往能使一種生物戰(zhàn)勝另一種生物,然而從長遠(yuǎn)看,各種勢(shì)力是微妙平衡的,自然界可以長期保持劃一的面貌。然而我們是多么無知,又是多么自說自話,一聽到一種生物的滅絕就大驚小怪;又不知道其原因,就提出毀滅世界的災(zāi)變說,或者創(chuàng)造出一些法則來規(guī)定生物類型的壽命!
我想再舉一個(gè)事例,說明自然界等級(jí)中相距甚遠(yuǎn)的動(dòng)植物如何被復(fù)雜的關(guān)系網(wǎng)聯(lián)結(jié)在一起。以后還有機(jī)會(huì)闡明,英格蘭這個(gè)地區(qū)的外來植物亮毛半邊蓮(Lobelia fulgens)從來沒有昆蟲光顧,結(jié)果由于它的特殊構(gòu)造,從不結(jié)籽。許多蘭科植物都絕對(duì)需要蛾子的光顧,帶走花粉塊,從而使其受精。我還有理由相信,大黃蜂是三色堇(Viola tricolor)受精所不可缺少的,因?yàn)閯e的蜂類都不來光顧這種花。我從試驗(yàn)里發(fā)現(xiàn),蜂類的光顧對(duì)于三葉草(clover)受精,哪怕不是不可或缺,也至少是高度有益。而只有大黃蜂才光顧紅三葉草(苜蓿Trifolium pratense),因?yàn)閯e的蜂類都不能接觸到它的花蜜。因此,我不懷疑,如果英格蘭的整個(gè)大黃蜂屬都滅絕了或變得稀少,三色堇和紅三葉草也會(huì)變得稀少,或全部消失。任何地方的大黃蜂數(shù)量大都是由鼠的多少來決定的,田鼠毀滅蜂房蜂群。紐曼(H. Newman)先生長期研究過大黃蜂的習(xí)性,認(rèn)為“全英格蘭三分之二以上的大黃蜂都是這樣消滅的”。眾所周知,鼠的數(shù)量大多取決于貓的數(shù)量;紐曼先生說:“在村莊和小鎮(zhèn)的附近,我看見大黃蜂窩比別的地方多,我把這一點(diǎn)歸因于有大量的貓?jiān)诓妒蟮木壒??!币虼丝梢韵嘈?,一處地方有大量的貓科?dòng)物,先干預(yù)鼠,再干預(yù)蜂,就可以決定該地區(qū)內(nèi)某些花的多少!
針對(duì)每一個(gè)物種,在不同的生命時(shí)期、不同的季節(jié)和年份,有多種不同的抑制因素會(huì)出現(xiàn),對(duì)其發(fā)生作用;其中某一種或者某少數(shù)幾種抑制作用一般最有力量,但在決定物種的平均數(shù),乃至它的生存上,則需要共同發(fā)揮作用。有時(shí)候可以闡明,同一物種在不同地區(qū)所受到的抑制作用大相徑庭。當(dāng)我們看到糾纏在岸邊的植物和灌木時(shí),易于把它們的比例數(shù)和種類歸因于所謂的偶然機(jī)會(huì)。但這是大錯(cuò)特錯(cuò)的!誰都聽說過,美洲森林砍伐以后,便有很不同的植物群落生長起來;但已經(jīng)有人談到,美國南方的印第安古冢上現(xiàn)在生長的樹木同周圍的處女林相似,呈現(xiàn)了同樣美麗的多樣性和同樣比例的各類植物。千百年來,在每年各自成千上萬散播種子的若干樹類之間,想必進(jìn)行了十分激烈的斗爭;昆蟲和昆蟲之間——昆蟲、蝸牛等動(dòng)物與猛禽、猛獸之間——進(jìn)行了何等戰(zhàn)爭啊,它們都努力增殖,彼此相食,或者吃樹、吃樹的種子和幼苗,或者吃最初密布于地面而抑制這些樹木生長的其他植物!將一把羽毛拋出,都必須依照一定的法則落到地面上;但是這個(gè)問題比起無數(shù)動(dòng)植物之間的關(guān)系,就顯得非常簡單了,動(dòng)植物的作用和反作用在千百年里決定了現(xiàn)今生長在印第安廢墟上各類樹木的比例數(shù)和樹木的種類!
生物彼此的依存關(guān)系,有如寄生物之于寄主,一般是在性狀級(jí)別遠(yuǎn)的生物之間發(fā)生的。嚴(yán)格意義上,彼此進(jìn)行著生存斗爭的生物往往如此,例如飛蝗類和食草獸。不過同種個(gè)體之間的斗爭幾乎都是你死我活的,因?yàn)樽⊥粎^(qū)域,需要同樣的食物,還遭遇同樣的危險(xiǎn)。同種的變種之間的斗爭一般差不多是同等劇烈的,而且我們有時(shí)看到爭奪很快就見分曉。例如幾個(gè)小麥變種混播,然后把混雜的種子再播種,那些最適于土壤氣候的,或者天生最能育的變種,便會(huì)打敗別的變種,結(jié)籽更多,幾年之后就會(huì)將其淘汰。哪怕極度相近的變種,如顏色不同的香豌豆,混合種植時(shí),必須每年分別采收種子,播種時(shí)再照適當(dāng)?shù)谋壤旌?,否則弱種類的數(shù)量會(huì)不斷減少而終于消滅。綿羊的變種也是這樣:有人斷言,某些山地綿羊變種能使另外一些變種餓死,所以不能混養(yǎng)。不同變種的醫(yī)用蛭混養(yǎng),結(jié)果也這樣。讓任何一種家養(yǎng)植物或家畜的一些變種,像在自然狀況下那樣相互進(jìn)行斗爭,假如不每年選種或揀選幼畜,那么甚至可以懷疑這些變種有沒有一模一樣的體力、習(xí)性和體質(zhì),足以讓一個(gè)混合群的原始比例維持六代之久。
由于同屬的物種通常在習(xí)性和體質(zhì)方面是相似的,并且在構(gòu)造方面總是相似(雖然不是絕對(duì)如此),所以之間如發(fā)生競爭,斗爭一般要比異屬的物種之間更劇烈。近來有一種燕子在美國局部地區(qū)拓展了,致使另一種燕子減量,可見這一點(diǎn)不謬。近來蘇格蘭一些地方槲鶇(missel-thrush)增量,導(dǎo)致歌鶇(song-thrush)的減量。我們不是每每聽說,在千差萬別的氣候下一個(gè)鼠種代替了另一鼠種!在俄羅斯,小型的亞洲種蟑螂(Asiatic cockroach)入境之后,趕著大型蟑螂到處跑。一種田芥菜(charlock)將淘汰另一種,如此種種,不一而足。我們能夠隱約看到,大自然系統(tǒng)中填補(bǔ)近乎相同地位的近似類型之間的競爭為什么最為劇烈;但我們大概怎么也說不確切,在偉大的生存斗爭中一個(gè)物種為什么戰(zhàn)勝了另一個(gè)物種。
從上述可以演繹出高度重要的結(jié)論,即每一種生物的構(gòu)造,以最基本然而往往是隱蔽的方式和一切其他生物的構(gòu)造相關(guān)聯(lián),競爭食物或住所,要么被迫躲避它們,要么捕殺它們。在虎牙虎爪的構(gòu)造上這一點(diǎn)很明顯,攀附在虎毛上的寄生蟲的腿和爪的構(gòu)造也這樣。但是蒲公英美麗的羽毛種子和水生甲蟲扁平而生有排毛的腿,乍看似乎僅僅和空氣和水有關(guān)系,但羽毛種子的優(yōu)點(diǎn),無疑在于和密布他種植物的地面最密切相關(guān);這樣,種子才能廣泛散布,落在空地上。水生甲蟲的腿的構(gòu)造,非常適于潛水,以便和其他水棲昆蟲競爭,捕食食物,并逃避被捕食。
許多植物種子里貯藏養(yǎng)料,乍看似乎和其他植物沒有任何關(guān)系。但是這樣的種子(例如豌豆和蠶豆)播種在大草中間時(shí),萌發(fā)的幼小植株就能茁壯生長,由此可以推知,種子中養(yǎng)料的主要用途是有利于幼苗生長,以便和四周繁茂的其他植物作斗爭。
看一看生長在分布范圍中央的植物吧,為什么其數(shù)量沒有翻一番、翻兩番呢?我們知道它對(duì)于稍熱或稍冷,稍潮濕或稍干燥的環(huán)境都能完全抵御,因?yàn)樗芊植嫉缴詿嵘岳?、稍濕稍干的其他地區(qū)。在此可以清楚看出,如果我們指望這種植物有能力增殖,就必須使它對(duì)競爭者、對(duì)于吃它的動(dòng)物占些優(yōu)勢(shì)。在它的地理分布范圍邊緣,如果體質(zhì)針對(duì)氣候而發(fā)生變化,這顯然有利于該植物;但有理由相信,只有少數(shù)的動(dòng)植物能分布到僅僅嚴(yán)酷的氣候就可加以消滅的遠(yuǎn)方。除非到達(dá)生活范圍的極限,如北極地方或荒漠的邊緣,競爭是不會(huì)停止的。有些地面可能極冷、極干,然而仍有少數(shù)幾個(gè)物種或同種的個(gè)體之間為著爭取最暖濕的地點(diǎn)而進(jìn)行斗爭。
由此可見,當(dāng)一種動(dòng)植物放置在新的地方而處于新的競爭者之中時(shí),雖然氣候可能和原產(chǎn)地一模一樣,但生活條件一般已發(fā)生了質(zhì)變。如果要它在新地方增加平均數(shù),就得放棄在其原產(chǎn)地的做法,而使用不同的方法來改變它;必須使它對(duì)一批不同的競爭者和敵害占些優(yōu)勢(shì)。
因此,我們不妨去設(shè)想使任何類型對(duì)其他類型占有優(yōu)勢(shì)。也許事到臨頭,我們根本不知道應(yīng)該如何下手才能如愿以償。這使我們確信,我們對(duì)于一切生物之間的相互關(guān)系實(shí)在無知;此種信念似乎難以獲得,所以是必要的。我們所能做到的,就是牢牢記住,每一種生物都努力按照幾何級(jí)數(shù)增殖;每一種都在生命的某一時(shí)期,一年中的某一季節(jié),每一世代或隔代,必須進(jìn)行生存斗爭,并且遭受大量毀滅。想到這種斗爭,我們可以安慰自己,堅(jiān)信自然界的戰(zhàn)爭不是無休無止的,恐懼是感覺不到的,死亡一般是瞬間發(fā)生的,而強(qiáng)壯的、健康的和幸運(yùn)的個(gè)體則生存并繁殖下去。
瘋狂英語 英語語法 新概念英語 走遍美國 四級(jí)聽力 英語音標(biāo) 英語入門 發(fā)音 美語 四級(jí) 新東方 七年級(jí) 賴世雄 zero是什么意思大連市新生巷小區(qū)英語學(xué)習(xí)交流群