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1.【址:a g 9 559⒐ v i p】1  Effects of Use and Disuse
2.  BEF0RE 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.
3.  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, &c., 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 hundreds 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.
4.  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.
5.  These difficulties and objections may be classed under the following heads:-Firstly, why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion instead of the species being, as we see them, well defined?
6.  It has often been assumed that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary, and likewise to withstand diverse climates. I do not dispute that these capacities have added largely to the value of most of our domesticated productions; but how could a savage possibly know, when he first tamed an animal, whether it would vary in succeeding generations, and whether it would endure other climates? Has the little variability of the ass or guinea-fowl, or the small power of endurance of warmth by the reindeer, or of cold by the common camel, prevented their domestication? I cannot doubt that if other animals and plants, equal in number to our domesticated productions, and belonging to equally diverse classes and countries, were taken from a state of nature, and could be made to breed for an equal number of generations under domestication, they would vary on an average as largely as the parent species of our existing domesticated productions have varied.

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1.  Habit is hereditary with plants, as in the period of flowering, in the amount of rain requisite for seeds to germinate, in the time of sleep, &c., and this leads me to say a few words on acclimatisation. As it is extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe that all the species of the same genus have descended from a single parent, if this view be correct, acclimatisation must be readily effected during long-continued descent. It is notorious that each species is adapted to the climate of its own home: species from an arctic or even from a temperate region cannot endure a tropical climate, or conversely. So again, many succulent plants cannot endure a damp climate. But the degree of adaptation of species to the climates under which they live is often overrated. We may infer this from our frequent inability to predict whether or not an imported plant will endure our climate, and from the number of plants and animals brought from warmer countries which here enjoy good health. We have reason to believe that species in a state of nature are limited in their ranges by the competition of other organic beings quite as much as, or more than, by adaptation to particular climates. But whether or not the adaptation be generally very close, we have evidence, in the case of some few plants, of their becoming, to a certain extent, naturally habituated to different temperatures, or becoming acclimatised: thus the pines and rhododendrons, raised from seed collected by Dr Hooker from trees growing at different heights on the Himalaya were found in this country to possess different constitutional powers of resisting cold. Mr Thwaites informs me that he has observed similar facts in Ceylon, and analogous observations have been made by Mr H. C. Watson on European species of plants brought from the Azores to England. In regard to animals, several authentic cases could be given of species within historical times having largely extended their range from warmer to cooler latitudes, and conversely; but we do not positively know that these animals were strictly adapted to their native climate, but in all ordinary cases we assume such to be the case; nor do we know that they have subsequently become acclimatised to their new homes.As I believe that our domestic animals were originally chosen by uncivilised man because they were useful and bred readily under confinement, and not because they were subsequently found capable of far-extended transportation, I think the common and extraordinary capacity in our domestic animals of not only withstanding the most different climates but of being perfectly fertile (a far severer test) under them, may be used as an argument that a large proportion of other animals, now in a state of nature, could easily be brought to bear widely different climates. We must not, however, push the foregoing argument too far, on account of the probable origin of some of our domestic animals from several wild stocks: the blood, for instance, of a tropical and arctic wolf or wild dog may perhaps be mingled in our domestic breeds. The rat and mouse cannot be considered as domestic animals, but they have been transported by man to many parts of the world, and now have a far wider range than any other rodent, living free under the cold climate of Faroe in the north and of the Falklands in the south, and on many islands in the torrid zones. Hence I am inclined to look at adaptation to any special climate as a quality readily grafted on an innate wide flexibility of constitution, which is common to most animals. On this view, the capacity of enduring the most different climates by man himself and by his domestic animals, and such facts as that former species of the elephant and rhinoceros were capable of enduring a glacial climate, whereas the living species are now all tropical or sub-tropical in their habits, ought not to be looked at as anomalies, but merely as examples of a very common flexibility of constitution, brought, under peculiar circumstances, into play.How much of the acclimatisation of species to any peculiar climate is due to mere habit, and how much to the natural selection of varieties having different innate constitutions, and how much to means combined, is a very obscure question. That habit or custom has some influence I must believe, both from analogy, and from the incessant advice given in agricultural works, even in the ancient Encyclopaedias of China, to be very cautious in transposing animals from one district to another; for it is not likely that man should have succeeded in selecting so many breeds and sub-breeds with constitutions specially fitted for their own districts: the result must, I think, be due to habit. On the other hand, I can see no reason to doubt that natural selection will continually tend to preserve those individuals which are born with constitutions best adapted to their native countries. In treatises on many kinds of cultivated plants, certain varieties are said to withstand certain climates better than others: this is very strikingly shown in works on fruit trees published in the United States, in which certain varieties are habitually recommended for the northern, and others for the southern States; and as most of these varieties are of recent origin, they cannot owe their constitutional differences to habit. The case of the Jerusalem artichoke, which is never propagated by seed, and of which consequently new varieties have not been produced, has even been advanced for it is now as tender as ever it was -- as proving that acclimatisation cannot be effected! The case, also, of the kidney-bean has been often cited for a similar purpose, and with much greater weight; but until some one will sow, during a score of generations, his kidney-beans so early that a very large proportion are destroyed by frost, and then collect seed from the few survivors, with care to prevent accidental crosses, and then again get seed from these seedlings, with the same precautions, the experiment cannot be said to have been even tried. Nor let it be supposed that no differences in the constitution of seedling kidney-beans ever appear, for an account has been published how much more hardy some seedlings appeared to be than others.On the whole, I think we may conclude that habit, use, and disuse, have, in some cases, played a considerable part in the modification of the constitution, and of the structure of various organs; but that the effects of use and disuse have often been largely combined with, and sometimes overmastered by, the natural selection of innate differences.
2.  It is worth while to reflect for a moment on the character of the new species F14, which is supposed not to have diverged much in character, but to have retained the form of (F), either unaltered or altered only in a slight degree. In this case, its affinities to the other fourteen new species will be of a curious and circuitous nature. Having descended from a form which stood between the two parent-species (A) and (I), now supposed to be extinct and unknown, it will be in some degree intermediate in character between the two groups descended from these species. But as these two groups have gone on diverging in character from the type of their parents, the new species (F14) will not be directly intermediate between them, but rather between types of the two groups; and every naturalist will be able to bring some such case before his mind.
3.  Any variation which is not inherited is unimportant for us. But the number and diversity of inheritable deviations of structure, both those of slight and those of considerable physiological importance, is endless. Dr Prosper Lucas's treatise, in two large volumes, is the fullest and the best on this subject. No breeder doubts how strong is the tendency to inheritance: like produces like is his fundamental belief: doubts have been thrown on this principle by theoretical writers alone. When a deviation appears not unfrequently, and we see it in the father and child, we cannot tell whether it may not be due to the same original cause acting on both; but when amongst individuals, apparently exposed to the same conditions, any very rare deviation, due to some extraordinary combination of circumstances, appears in the parent say, once amongst several million individuals and it reappears in the child, the mere doctrine of chances almost compels us to attribute its reappearance to inheritance. Every one must have heard of cases of albinism, prickly skin, hairy bodies, &c. appearing in several members of the same family. If strange and rare deviations of structure are truly inherited, less strange and commoner deviations may be freely admitted to be inheritable. Perhaps the correct way of viewing the whole subject, would be, to look at the inheritance of every character whatever as the rule, and non-inheritance as the anomaly.The laws governing inheritance are quite unknown; no one can say why the same peculiarity in different individuals of the same species, and in individuals of different species, is sometimes inherited and sometimes not so; why the child often reverts in certain characters to its grandfather or grandmother or other much more remote ancestor; why a peculiarity is often transmitted from one sex to both sexes or to one sex alone, more commonly but not exclusively to the like sex. It is a fact of some little importance to us, that peculiarities appearing in the males of our domestic breeds are often transmitted either exclusively, or in a much greater degree, to males alone. A much more important rule, which I think may be trusted, is that, at whatever period of life a peculiarity first appears, it tends to appear in the offspring at a corresponding age, though sometimes earlier. In many cases this could not be otherwise: thus the inherited peculiarities in the horns of cattle could appear only in the offspring when nearly mature; peculiarities in the silkworm are known to appear at the corresponding caterpillar or cocoon stage. But hereditary diseases and some other facts make me believe that the rule has a wider extension, and that when there is no apparent reason why a peculiarity should appear at any particular age, yet that it does tend to appear in the offspring at the same period at which it first appeared in the parent. I believe this rule to be of the highest importance in explaining the laws of embryology. These remarks are of course confined to the first appearance of the peculiarity, and not to its primary cause, which may have acted on the ovules or male element; in nearly the same manner as in the crossed offspring from a short-horned cow by a long-horned bull, the greater length of horn, though appearing late in life, is clearly due to the male element.Having alluded to the subject of reversion, I may here refer to a statement often made by naturalists namely, that our domestic varieties, when run wild, gradually but certainly revert in character to their aboriginal stocks. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature. I have in vain endeavoured to discover on what decisive facts the above statement has so often and so boldly been made. There would be great difficulty in proving its truth: we may safely conclude that very many of the most strongly-marked domestic varieties could not possibly live in a wild state. In many cases we do not know what the aboriginal stock was, and so could not tell whether or not nearly perfect reversion had ensued. It would be quite necessary, in order to prevent the effects of intercrossing, that only a single variety should be turned loose in its new home. Nevertheless, as our varieties certainly do occasionally revert in some of their characters to ancestral forms, it seems to me not improbable, that if we could succeed in naturalising, or were to cultivate, during many generations, the several races, for instance, of the cabbage, in very poor soil (in which case, however, some effect would have to be attributed to the direct action of the poor soil), that they would to a large extent, or even wholly, revert to the wild aboriginal stock. Whether or not the experiment would succeed, is not of great importance for our line of argument; for by the experiment itself the conditions of life are changed. If it could be shown that our domestic varieties manifested a strong tendency to reversion, that is, to lose their acquired characters, whilst kept under unchanged conditions, and whilst kept in a considerable body, so that free intercrossing might check, by blending together, any slight deviations of structure, in such case, I grant that we could deduce nothing from domestic varieties in regard to species. But there is not a shadow of evidence in favour of this view: to assert that we could not breed our cart and race-horses, long and short-horned cattle and poultry of various breeds, and esculent vegetables, for an almost infinite number of generations, would be opposed to all experience. I may add, that when under nature the conditions of life do change, variations and reversions of character probably do occur; but natural selection, as will hereafter be explained, will determine how far the new characters thus arising shall be preserved.When we look to the hereditary varieties or races of our domestic animals and plants, and compare them with species closely allied together, we generally perceive in each domestic race, as already remarked, less uniformity of character than in true species. Domestic races of the same species, also, often have a somewhat monstrous character; by which I mean, that, although differing from each other, and from the other species of the same genus, in several trifling respects, they often differ in an extreme degree in some one part, both when compared one with another, and more especially when compared with all the species in nature to which they are nearest allied. With these exceptions (and with that of the perfect fertility of varieties when crossed, a subject hereafter to be discussed), domestic races of the same species differ from each other in the same manner as, only in most cases in a lesser degree than, do closely-allied species of the same genus in a state of nature. I think this must be admitted, when we find that there are hardly any domestic races, either amongst animals or plants, which have not been ranked by some competent judges as mere varieties, and by other competent judges as the descendants of aboriginally distinct species. If any marked distinction existed between domestic races and species, this source of doubt could not so perpetually recur. It has often been stated that domestic races do not differ from each other in characters of generic value. I think it could be shown that this statement is hardly correct; but naturalists differ most widely in determining what characters are of generic value; all such valuations being at present empirical. Moreover, on the view of the origin of genera which I shall presently give, we have no right to expect often to meet with generic differences in our domesticated productions.When we attempt to estimate the amount of structural difference between the domestic races of the same species, we are soon involved in doubt, from not knowing whether they have descended from one or several parent-species. This point, if could be cleared up, would be interesting; if, for instance, it could be shown that the greyhound, bloodhound, terrier, spaniel, and bull-dog, which we all know propagate their kind so truly, were the offspring of any single species, then such facts would have great weight in making us doubt about the immutability of the many very closely allied and natural species for instance, of the many foxes inhabiting different quarters of the world. I do not believe, as we shall presently see, that all our dogs have descended from any one wild species; but, in the case of some other domestic races, there is presumptive, or even strong, evidence in favour of this view.
4.  Alph. De Candolle and others have shown that plants which have very wide ranges generally present varieties; and this might have been expected, as they become exposed to diverse physical conditions, and as they come into competition (which, as we shall hereafter see, is a far more important circumstance) with different sets of organic beings. But my tables further show that, in any limited country, the species which are most common, that is abound most in individuals, and the species which are most widely diffused within their own country (and this is a different consideration from wide range, and to a certain extent from commonness), often give rise to varieties sufficiently well-marked to have been recorded in botanical works. Hence it is the most flourishing, or, as they may be called, the dominant species, those which range widely over the world, are the most diffused in their own country, and are the most numerous in individuals, which oftenest produce well-marked varieties, or, as I consider them, incipient species. And this, perhaps, might have been anticipated; for, as varieties, in order to become in any degree permanent, necessarily have to struggle with the other inhabitants of the country, the species which are already dominant will be the most likely to yield offspring which, though in some slight degree modified, will still inherit those advantages that enabled their parents to become dominant over their compatriots.If the plants inhabiting a country and described in any Flora be divided into two equal masses, all those in the larger genera being placed on one side, and all those in the smaller genera on the other side, a somewhat larger number of the very common and much diffused or dominant species will be found on the side of the larger genera. This, again, might have been anticipated; for the mere fact of many species of the same genus inhabiting any country, shows that there is something in the organic or inorganic conditions of that country favourable to the genus; and, consequently, we might have expected to have found in the larger genera, or those including many species, a large proportional number of dominant species. But so many causes tend to obscure this result, that I am surprised that my tables show even a small majority on the side of the larger genera. I will here allude to only two causes of obscurity. Fresh-water and salt-loving plants have generally very wide ranges and are much diffused, but this seems to be connected with the nature of the stations inhabited by them, and has little or no relation to the size of the genera to which the species belong. Again, plants low in the scale of organisation are generally much more widely diffused than plants higher in the scale; and here again there is no close relation to the size of the genera. The cause of lowly-organised plants ranging widely will be discussed in our chapter on geographical distribution.From looking at species as only strongly-marked and well-defined varieties, I was led to anticipate that the species of the larger genera in each country would oftener present varieties, than the species of the smaller genera; for wherever many closely related species (i.e. species of the same genus) have been formed, many varieties or incipient species ought, as a general rule, to be now forming. Where many large trees grow, we expect to find saplings. Where many species of a genus have been formed through variation, circumstances have been favourable for variation; and hence we might expect that the circumstances would generally be still favourable to variation. On the other hand, if we look at each species as a special act of creation, there is no apparent reason why more varieties should occur in a group having many species, than in one having few.
5.  Circumstances favourable to Natural Selection
6.  Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some little effect may, perhaps, be attributed to the direct action of the external conditions of life, and some little to habit; but he would be a bold man who would account by such agencies for the differences of a dray and race horse, a greyhound and bloodhound, a carrier and tumbler pigeon. One of the most remarkable features in our domesticated races is that we see in them adaptation, not indeed to the animal's or plant's own good, but to man's use or fancy. Some variations useful to him have probably arisen suddenly, or by one step; many botanists, for instance, believe that the fuller's teazle, with its hooks, which cannot be rivalled by any mechanical contrivance, is only a variety of the wild Dipsacus; and this amount of change may have suddenly arisen in a seedling. So it has probably been with the turnspit dog; and this is known to have been the case with the ancon sheep. But when we compare the dray-horse and race-horse, the dromedary and camel, the various breeds of sheep fitted either for cultivated land or mountain pasture, with the wool of one breed good for one purpose, and that of another breed for another purpose; when we compare the many breeds of dogs, each good for man in very different ways; when we compare the gamecock, so pertinacious in battle, with other breeds so little quarrelsome, with 'everlasting layers' which never desire to sit, and with the bantam so small and elegant; when we compare the host of agricultural, culinary, orchard, and flower-garden races of plants, most useful to man at different seasons and for different purposes, or so beautiful in his eyes, we must, I think, look further than to mere variability. We cannot suppose that all the breeds were suddenly produced as perfect and as useful as we now see them; indeed, in several cases, we know that this has not been their history. The key is man's power of accumulative selection: nature gives successive variations; man adds them up in certain directions useful to him. In this sense he may be said to make for himself useful breeds.The great power of this principle of selection is not hypothetical. It is certain that several of our eminent breeders have, even within a single lifetime, modified to a large extent some breeds of cattle and sheep. In order fully to realise what they have done, it is almost necessary to read several of the many treatises devoted to this subject, and to inspect the animals. Breeders habitually speak of an animal's organisation as something quite plastic, which they can model almost as they please. If I had space I could quote numerous passages to this effect from highly competent authorities. Youatt, who was probably better acquainted with the works of agriculturalists than almost any other individual, and who was himself a very good judge of an animal, speaks of the principle of selection as 'that which enables the agriculturist, not only to modify the character of his flock, but to change it altogether. It is the magician's wand, by means of which he may summon into life whatever form and mould he pleases.' Lord Somerville, speaking of what breeders have done for sheep, says: 'It would seem as if they had chalked out upon a wall a form perfect in itself, and then had given it existence.' That most skilful breeder, Sir John Sebright, used to say, with respect to pigeons, that 'he would produce any given feather in three years, but it would take him six years to obtain head and beak.' In Saxony the importance of the principle of selection in regard to merino sheep is so fully recognised, that men follow it as a trade: the sheep are placed on a table and are studied, like a picture by a connoisseur; this is done three times at intervals of months, and the sheep are each time marked and classed, so that the very best may ultimately be selected for breeding.What English breeders have actually effected is proved by the enormous prices given for animals with a good pedigree; and these have now been exported to almost every quarter of the world. The improvement is by no means generally due to crossing different breeds; all the best breeders are strongly opposed to this practice, except sometimes amongst closely allied sub-breeds. And when a cross has been made, the closest selection is far more indispensable even than in ordinary cases. If selection consisted merely in separating some very distinct variety, and breeding from it, the principle would be so obvious as hardly to be worth notice; but its importance consists in the great effect produced by the accumulation in one direction, during successive generations, of differences absolutely inappreciable by an uneducated eye differences which I for one have vainly attempted to appreciate. Not one man in a thousand has accuracy of eye and judgement sufficient to become an eminent breeder. If gifted with these qualities, and he studies his subject for years, and devotes his lifetime to it with indomitable perseverance, he will succeed, and may make great improvements; if he wants any of these qualities, he will assuredly fail. Few would readily believe in the natural capacity and years of practice requisite to become even a skilful pigeon-fancier.The same principles are followed by horticulturists; but the variations are here often more abrupt. No one supposes that our choicest productions have been produced by a single variation from the aboriginal stock. We have proofs that this is not so in some cases, in which exact records have been kept; thus, to give a very trifling instance, the steadily-increasing size of the common gooseberry may be quoted. We see an astonishing improvement in many florists' flowers, when the flowers of the present day are compared with drawings made only twenty or thirty years ago. When a race of plants is once pretty well established, the seed-raisers do not pick out the best plants, but merely go over their seed-beds, and pull up the 'rogues,' as they call the plants that deviate from the proper standard. With animals this kind of selection is, in fact, also followed; for hardly any one is so careless as to allow his worst animals to breed.

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1.  Thirdly, can instincts be acquired and modified through natural selection? What shall we say to so marvellous an instinct as that which leads the bee to make cells, which have practically anticipated the discoveries of profound mathematicians?
2.  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 pairs 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.
3.  LONG before having arrived at this part of my work, a crowd of difficulties will have occurred to the reader. Some of them are so grave that to this day I can never reflect on them without being staggered; but, to the best of my judgment, the greater number are only apparent, and those that are real are not, I think, fatal to my theory.
4.  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.
5.   The accompanying diagram will aid us in understanding this rather perplexing subject. Let A to L represent the species of a genus large in its own country; these species are supposed to resemble each other in unequal degrees, as is so generally the case in nature, and as is represented in the diagram by the letters standing at unequal distances. I have said a large genus, because we have seen in the second chapter, that on an average more of the species of large genera vary than of small genera; and the varying species of the large genera present a greater number of varieties. We have, also, seen that the species, which are the commonest and the most widely-diffused, vary more than rare species with restricted ranges. Let (A) be a common, widely-diffused, and varying species, belonging to a genus large in its own country. The little fan of diverging dotted lines of unequal lengths proceeding from (A), may represent its varying offspring. The variations are supposed to be extremely slight, but of the most diversified nature; they are not supposed all to appear simultaneously, but often after long intervals of time; nor are they all supposed to endure for equal periods. Only those variations which are in some way profitable will be preserved or naturally selected. And here the importance of the principle of benefit being derived from divergence of character comes in; for this will generally lead to the most different or divergent variations (represented by the outer dotted lines) being preserved and accumulated by natural selection. When a dotted line reaches one of the horizontal lines, and is there marked by a small numbered letter, a sufficient amount of variation is supposed to have been accumulated to have formed a fairly well-marked variety, such as would be thought worthy of record in a systematic work.The intervals between the horizontal lines in the diagram, may represent each a thousand generations; but it would have been better if each had represented ten thousand generations. After a thousand generations, species (A) is supposed to have produced two fairly well-marked varieties, namely a1 and m1. These two varieties will generally continue to be exposed to the same conditions which made their parents variable, and the tendency to variability is in itself hereditary, consequently they will tend to vary, and generally to vary in nearly the same manner as their parents varied. Moreover, these two varieties, being only slightly modified forms, will tend to inherit those advantages which made their common parent (A) more numerous than most of the other inhabitants of the same country; they will likewise partake of those more general advantages which made the genus to which the parent-species belonged, a large genus in its own country. And these circumstances we know to be favourable to the production of new varieties.
6.  Intercrossing plays a very important part in nature in keeping the individuals of the same species, or of the same variety, true and uniform in character. It will obviously thus act far more efficiently with those animals which unite for each birth; but I have already attempted to show that we have reason to believe that occasional intercrosses take place with all animals and with all plants. Even if these take place only at long intervals, I am convinced that the young thus produced will gain so much in vigour and fertility over the offspring from long-continued self-fertilisation, that they will have a better chance of surviving and propagating their kind; and thus, in the long run, the influence of intercrosses, even at rare intervals, will be great. If there exist organic beings which never intercross, uniformity of character can be retained amongst them, as long as their conditions of life remain the same, only through the principle of inheritance, and through natural selection destroying any which depart from the proper type; but if their conditions of life change and they undergo modification, uniformity of character can be given to their modified offspring, solely by natural selection preserving the same favourable variations.Isolation, also, is an important element in the process of natural selection. In a confined or isolated area, if not very large, the organic and inorganic conditions of life will generally be in a great degree uniform; so that natural selection will tend to modify all the individuals of a varying species throughout the area in the same manner in relation to the same conditions. Intercrosses, also, with the individuals of the same species, which otherwise would have inhabited the surrounding and differently circumstanced districts, will be prevented. But isolation probably acts more efficiently in checking the immigration of better adapted organisms, after any physical change, such as of climate or elevation of the land, &c.; and thus new places in the natural economy of the country are left open for the old inhabitants to struggle for, and become adapted to, through modifications in their structure and constitution. Lastly, isolation, by checking immigration and consequently competition, will give time for any new variety to be slowly improved; and this may sometimes be of importance in the production of new species. If, however, an isolated area be very small, either from being surrounded by barriers, or from having very peculiar physical conditions, the total number of the individuals supported on it will necessarily be very small; and fewness of individuals will greatly retard the production of new species through natural selection, by decreasing the chance of the appearance of favourable variations.If we turn to nature to test the truth of these remarks, and look at any small isolated area, such as an oceanic island, although the total number of the species inhabiting it, will be found to be small, as we shall see in our chapter on geographical distribution; yet of these species a very large proportion are endemic, that is, have been produced there, and nowhere else. Hence an oceanic island at first sight seems to have been highly favourable for the production of new species. But we may thus greatly deceive ourselves, for to ascertain whether a small isolated area, or a large open area like a continent, has been most favourable for the production of new organic forms, we ought to make the comparison within equal times; and this we are incapable of doing.

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1.  It is well known that several animals, belonging to the most different classes, which inhabit the caves of Styria and of Kentucky, are blind. In some of the crabs the foot-stalk for the eye remains, though the eye is gone; the stand for the telescope is there, though the telescope with its glasses has been lost. As it is difficult to imagine that eyes, though useless, could be in any way injurious to animals living in darkness, I attribute their loss wholly to disuse. In one of the blind animals, namely, the cave-rat, the eyes are of immense size; and Professor Silliman thought that it regained, after living some days in the light, some slight power of vision. In the same manner as in Madeira the wings of some of the insects have been enlarged, and the wings of others have been reduced by natural selection aided by use and disuse, so in the case of the cave-rat natural selection seems to have struggled with the loss of light and to have increased the size of the eyes; whereas with all the other inhabitants of the caves, disuse by itself seems to have done its work.It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar climate; so that on the common view of the blind animals having been separately created for the American and European caverns, close similarity in their organisation and affinities might have been expected; but, as Schi?dte and others have remarked, this is not the case, and the cave-insects of the two continents are not more closely allied than might have been anticipated from the general resemblance of the other inhabitants of North America and Europe. On my view we must suppose that American animals, having ordinary powers of vision, slowly migrated by successive generations from the outer world into the deeper and deeper recesses of the Kentucky caves, as did European animals into the caves of Europe. We have some evidence of this gradation of habit; for, as Schi?dte remarks, 'animals not far remote from ordinary forms, prepare the transition from light to darkness. Next follow those that are constructed for twilight; and, last of all, those destined for total darkness.' By the time that an animal had reached, after numberless generations, the deepest recesses, disuse will on this view have more or less perfectly obliterated its eyes, and natural selection will often have effected other changes, such as an increase in the length of the antennae or palpi, as a compensation for blindness. Notwithstanding such modifications, we might expect still to see in the cave-animals of America, affinities to the other inhabitants of that continent, and in those of Europe, to the inhabitants of the European continent. And this is the case with some of the American cave-animals, as I hear from Professor Dana; and some of the European cave-insects are very closely allied to those of the surrounding country. It would be most difficult to give any rational explanation of the affinities of the blind cave-animals to the other inhabitants of the two continents on the ordinary view of their independent creation. That several of the inhabitants of the caves of the Old and New Worlds should be closely related, we might expect from the well-known relationship of most of their other productions. Far from feeling any surprise that some of the cave-animals should be very anomalous, as Agassiz has remarked in regard to the blind fish, the Amblyopsis, and as is the case with the blind Proteus with reference to the reptiles of Europe, I am only surprised that more wrecks of ancient life have not been preserved, owing to the less severe competition to which the inhabitants of these dark abodes will probably have been exposed.Acclimatisation
2.  A long list could easily be given of 'sporting plants;' by this term gardeners mean a single bud or offset, which suddenly assumes a new and sometimes very different character from that of the rest of the plant. Such buds can be propagated by grafting, &c., and sometimes by seed. These 'sports' are extremely rare under nature, but far from rare under cultivation; and in this case we see that the treatment of the parent has affected a bud or offset, and not the ovules or pollen. But it is the opinion of most physiologists that there is no essential difference between a bud and an ovule in their earliest stages of formation; so that, in fact,'sports' support my view, that variability may be largely attributed to the ovules or pollen, or to both, having been affected by the treatment of the parent prior to the act of conception. These cases anyhow show that variation is not necessarily connected, as some authors have supposed, with the act of generation.
3.WHEN we look to the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us, is, that they generally differ much more from each other, than do the individuals of any one species or variety in a state of nature. When we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, I think we are driven to conclude that this greater variability is simply due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent-species have been exposed under nature. There is, also, I think, some probability in the view propounded by Andrew Knight, that this variability may be partly connected with excess of food. It seems pretty clear that organic beings must be exposed during several generations to the new conditions of life to cause any appreciable amount of variation; and that when the organisation has once begun to vary, it generally continues to vary for many generations. No case is on record of a variable being ceasing to be variable under cultivation. Our oldest cultivated plants, such as wheat, still often yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification.It has been disputed at what period of time the causes of variability, whatever they may be, generally act; whether during the early or late period of development of the embryo, or at the instant of conception. Geoffroy St Hilaire's experiments show that unnatural treatment of the embryo causes monstrosities; and monstrosities cannot be separated by any clear line of distinction from mere variations. But I am strongly inclined to suspect that the most frequent cause of variability may be attributed to the male and female reproductive elements having been affected prior to the act of conception. Several reasons make me believe in this; but the chief one is the remarkable effect which confinement or cultivation has on the functions of the reproductive system; this system appearing to be far more susceptible than any other part of the organization, to the action of any change in the conditions of life. Nothing is more easy than to tame an animal, and few things more difficult than to get it to breed freely under confinement, even in the many cases when the male and female unite. How many animals there are which will not breed, though living long under not very close confinement in their native country! This is generally attributed to vitiated instincts; but how many cultivated plants display the utmost vigour, and yet rarely or never seed! In some few such cases it has been found out that very trifling changes, such as a little more or less water at some particular period of growth, will determine whether or not the plant sets a seed. I cannot here enter on the copious details which I have collected on this curious subject; but to show how singular the laws are which determine the reproduction of animals under confinement, I may just mention that carnivorous animals, even from the tropics, breed in this country pretty freely under confinement, with the exception of the plantigrades or bear family; whereas, carnivorous birds, with the rarest exceptions, hardly ever lay fertile eggs. Many exotic plants have pollen utterly worthless, in the same exact condition as in the most sterile hybrids. When, on the one hand, we see domesticated animals and plants, though often weak and sickly, yet breeding quite freely under confinement; and when, on the other hand, we see individuals, though taken young from a state of nature, perfectly tamed, long-lived, and healthy (of which I could give numerous instances), yet having their reproductive system so seriously affected by unperceived causes as to fail in acting, we need not be surprised at this system, when it does act under confinement, acting not quite regularly, and producing offspring not perfectly like their parents or variable.Sterility has been said to be the bane of horticulture; but on this view we owe variability to the same cause which produces sterility; and variability is the source of all the choicest productions of the garden. I may add, that as some organisms will breed most freely under the most unnatural conditions (for instance, the rabbit and ferret kept in hutches), showing that their reproductive system has not been thus affected; so will some animals and plants withstand domestication or cultivation, and vary very slightly perhaps hardly more than in a state of nature.
4、  Alph. De Candolle and others have shown that plants which have very wide ranges generally present varieties; and this might have been expected, as they become exposed to diverse physical conditions, and as they come into competition (which, as we shall hereafter see, is a far more important circumstance) with different sets of organic beings. But my tables further show that, in any limited country, the species which are most common, that is abound most in individuals, and the species which are most widely diffused within their own country (and this is a different consideration from wide range, and to a certain extent from commonness), often give rise to varieties sufficiently well-marked to have been recorded in botanical works. Hence it is the most flourishing, or, as they may be called, the dominant species, those which range widely over the world, are the most diffused in their own country, and are the most numerous in individuals, which oftenest produce well-marked varieties, or, as I consider them, incipient species. And this, perhaps, might have been anticipated; for, as varieties, in order to become in any degree permanent, necessarily have to struggle with the other inhabitants of the country, the species which are already dominant will be the most likely to yield offspring which, though in some slight degree modified, will still inherit those advantages that enabled their parents to become dominant over their compatriots.If the plants inhabiting a country and described in any Flora be divided into two equal masses, all those in the larger genera being placed on one side, and all those in the smaller genera on the other side, a somewhat larger number of the very common and much diffused or dominant species will be found on the side of the larger genera. This, again, might have been anticipated; for the mere fact of many species of the same genus inhabiting any country, shows that there is something in the organic or inorganic conditions of that country favourable to the genus; and, consequently, we might have expected to have found in the larger genera, or those including many species, a large proportional number of dominant species. But so many causes tend to obscure this result, that I am surprised that my tables show even a small majority on the side of the larger genera. I will here allude to only two causes of obscurity. Fresh-water and salt-loving plants have generally very wide ranges and are much diffused, but this seems to be connected with the nature of the stations inhabited by them, and has little or no relation to the size of the genera to which the species belong. Again, plants low in the scale of organisation are generally much more widely diffused than plants higher in the scale; and here again there is no close relation to the size of the genera. The cause of lowly-organised plants ranging widely will be discussed in our chapter on geographical distribution.From looking at species as only strongly-marked and well-defined varieties, I was led to anticipate that the species of the larger genera in each country would oftener present varieties, than the species of the smaller genera; for wherever many closely related species (i.e. species of the same genus) have been formed, many varieties or incipient species ought, as a general rule, to be now forming. Where many large trees grow, we expect to find saplings. Where many species of a genus have been formed through variation, circumstances have been favourable for variation; and hence we might expect that the circumstances would generally be still favourable to variation. On the other hand, if we look at each species as a special act of creation, there is no apparent reason why more varieties should occur in a group having many species, than in one having few.
5、  Chapter 2 - Variation Under Nature

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  • 郝玉 08-08

      Habit is hereditary with plants, as in the period of flowering, in the amount of rain requisite for seeds to germinate, in the time of sleep, &c., and this leads me to say a few words on acclimatisation. As it is extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe that all the species of the same genus have descended from a single parent, if this view be correct, acclimatisation must be readily effected during long-continued descent. It is notorious that each species is adapted to the climate of its own home: species from an arctic or even from a temperate region cannot endure a tropical climate, or conversely. So again, many succulent plants cannot endure a damp climate. But the degree of adaptation of species to the climates under which they live is often overrated. We may infer this from our frequent inability to predict whether or not an imported plant will endure our climate, and from the number of plants and animals brought from warmer countries which here enjoy good health. We have reason to believe that species in a state of nature are limited in their ranges by the competition of other organic beings quite as much as, or more than, by adaptation to particular climates. But whether or not the adaptation be generally very close, we have evidence, in the case of some few plants, of their becoming, to a certain extent, naturally habituated to different temperatures, or becoming acclimatised: thus the pines and rhododendrons, raised from seed collected by Dr Hooker from trees growing at different heights on the Himalaya were found in this country to possess different constitutional powers of resisting cold. Mr Thwaites informs me that he has observed similar facts in Ceylon, and analogous observations have been made by Mr H. C. Watson on European species of plants brought from the Azores to England. In regard to animals, several authentic cases could be given of species within historical times having largely extended their range from warmer to cooler latitudes, and conversely; but we do not positively know that these animals were strictly adapted to their native climate, but in all ordinary cases we assume such to be the case; nor do we know that they have subsequently become acclimatised to their new homes.As I believe that our domestic animals were originally chosen by uncivilised man because they were useful and bred readily under confinement, and not because they were subsequently found capable of far-extended transportation, I think the common and extraordinary capacity in our domestic animals of not only withstanding the most different climates but of being perfectly fertile (a far severer test) under them, may be used as an argument that a large proportion of other animals, now in a state of nature, could easily be brought to bear widely different climates. We must not, however, push the foregoing argument too far, on account of the probable origin of some of our domestic animals from several wild stocks: the blood, for instance, of a tropical and arctic wolf or wild dog may perhaps be mingled in our domestic breeds. The rat and mouse cannot be considered as domestic animals, but they have been transported by man to many parts of the world, and now have a far wider range than any other rodent, living free under the cold climate of Faroe in the north and of the Falklands in the south, and on many islands in the torrid zones. Hence I am inclined to look at adaptation to any special climate as a quality readily grafted on an innate wide flexibility of constitution, which is common to most animals. On this view, the capacity of enduring the most different climates by man himself and by his domestic animals, and such facts as that former species of the elephant and rhinoceros were capable of enduring a glacial climate, whereas the living species are now all tropical or sub-tropical in their habits, ought not to be looked at as anomalies, but merely as examples of a very common flexibility of constitution, brought, under peculiar circumstances, into play.How much of the acclimatisation of species to any peculiar climate is due to mere habit, and how much to the natural selection of varieties having different innate constitutions, and how much to means combined, is a very obscure question. That habit or custom has some influence I must believe, both from analogy, and from the incessant advice given in agricultural works, even in the ancient Encyclopaedias of China, to be very cautious in transposing animals from one district to another; for it is not likely that man should have succeeded in selecting so many breeds and sub-breeds with constitutions specially fitted for their own districts: the result must, I think, be due to habit. On the other hand, I can see no reason to doubt that natural selection will continually tend to preserve those individuals which are born with constitutions best adapted to their native countries. In treatises on many kinds of cultivated plants, certain varieties are said to withstand certain climates better than others: this is very strikingly shown in works on fruit trees published in the United States, in which certain varieties are habitually recommended for the northern, and others for the southern States; and as most of these varieties are of recent origin, they cannot owe their constitutional differences to habit. The case of the Jerusalem artichoke, which is never propagated by seed, and of which consequently new varieties have not been produced, has even been advanced for it is now as tender as ever it was -- as proving that acclimatisation cannot be effected! The case, also, of the kidney-bean has been often cited for a similar purpose, and with much greater weight; but until some one will sow, during a score of generations, his kidney-beans so early that a very large proportion are destroyed by frost, and then collect seed from the few survivors, with care to prevent accidental crosses, and then again get seed from these seedlings, with the same precautions, the experiment cannot be said to have been even tried. Nor let it be supposed that no differences in the constitution of seedling kidney-beans ever appear, for an account has been published how much more hardy some seedlings appeared to be than others.On the whole, I think we may conclude that habit, use, and disuse, have, in some cases, played a considerable part in the modification of the constitution, and of the structure of various organs; but that the effects of use and disuse have often been largely combined with, and sometimes overmastered by, the natural selection of innate differences.

  • 狄德罗 08-08

      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.

  • 王政 08-08

     WHEN we look to the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us, is, that they generally differ much more from each other, than do the individuals of any one species or variety in a state of nature. When we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, I think we are driven to conclude that this greater variability is simply due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent-species have been exposed under nature. There is, also, I think, some probability in the view propounded by Andrew Knight, that this variability may be partly connected with excess of food. It seems pretty clear that organic beings must be exposed during several generations to the new conditions of life to cause any appreciable amount of variation; and that when the organisation has once begun to vary, it generally continues to vary for many generations. No case is on record of a variable being ceasing to be variable under cultivation. Our oldest cultivated plants, such as wheat, still often yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification.It has been disputed at what period of time the causes of variability, whatever they may be, generally act; whether during the early or late period of development of the embryo, or at the instant of conception. Geoffroy St Hilaire's experiments show that unnatural treatment of the embryo causes monstrosities; and monstrosities cannot be separated by any clear line of distinction from mere variations. But I am strongly inclined to suspect that the most frequent cause of variability may be attributed to the male and female reproductive elements having been affected prior to the act of conception. Several reasons make me believe in this; but the chief one is the remarkable effect which confinement or cultivation has on the functions of the reproductive system; this system appearing to be far more susceptible than any other part of the organization, to the action of any change in the conditions of life. Nothing is more easy than to tame an animal, and few things more difficult than to get it to breed freely under confinement, even in the many cases when the male and female unite. How many animals there are which will not breed, though living long under not very close confinement in their native country! This is generally attributed to vitiated instincts; but how many cultivated plants display the utmost vigour, and yet rarely or never seed! In some few such cases it has been found out that very trifling changes, such as a little more or less water at some particular period of growth, will determine whether or not the plant sets a seed. I cannot here enter on the copious details which I have collected on this curious subject; but to show how singular the laws are which determine the reproduction of animals under confinement, I may just mention that carnivorous animals, even from the tropics, breed in this country pretty freely under confinement, with the exception of the plantigrades or bear family; whereas, carnivorous birds, with the rarest exceptions, hardly ever lay fertile eggs. Many exotic plants have pollen utterly worthless, in the same exact condition as in the most sterile hybrids. When, on the one hand, we see domesticated animals and plants, though often weak and sickly, yet breeding quite freely under confinement; and when, on the other hand, we see individuals, though taken young from a state of nature, perfectly tamed, long-lived, and healthy (of which I could give numerous instances), yet having their reproductive system so seriously affected by unperceived causes as to fail in acting, we need not be surprised at this system, when it does act under confinement, acting not quite regularly, and producing offspring not perfectly like their parents or variable.Sterility has been said to be the bane of horticulture; but on this view we owe variability to the same cause which produces sterility; and variability is the source of all the choicest productions of the garden. I may add, that as some organisms will breed most freely under the most unnatural conditions (for instance, the rabbit and ferret kept in hutches), showing that their reproductive system has not been thus affected; so will some animals and plants withstand domestication or cultivation, and vary very slightly perhaps hardly more than in a state of nature.

  • 陈沸宇 08-08

      In regard to the domestic animals kept by uncivilised man, it should not be overlooked that they almost always have to struggle for their own food, at least during certain seasons. And in two countries very differently circumstanced, individuals of the same species, having slightly different constitutions or structure, would often succeed better in the one country than in the other, and thus by a process of 'natural selection,' as will hereafter be more fully explained, two sub-breeds might be formed. This, perhaps, partly explains what has been remarked by some authors, namely, that the varieties kept by savages have more of the character of species than the varieties kept in civilised countries.

  • 罗兰安腾 08-07

    {  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 pairs 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.

  • 孟斌 08-06

      Illustrations of the action of Natural Selection}

  • 罗集乡 08-06

      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.

  • 克萨拉 08-06

      We can clearly see this in the case of animals with simple habits. Take the case of a carnivorous quadruped, of which the number that can be supported in any country has long ago arrived at its full average. If its natural powers of increase be allowed to act, it can succeed in increasing (the country not undergoing any change in its conditions) only by its varying descendants seizing on places at present occupied by other animals: some of them, for instance, being enabled to feed on new kinds of prey, either dead or alive; some inhabiting new stations, climbing trees, frequenting water, and some perhaps becoming less carnivorous. The more diversified in habits and structure the descendants of our carnivorous animal became, the more places they would be enabled to occupy. What applies to one animal will apply throughout all time to all animals that is, if they vary for otherwise natural selection can do nothing. So it will be with plants. It has been experimentally proved, that if a plot of ground be sown with several distinct genera of grasses, a greater number of plants and a greater weight of dry herbage can thus be raised. The same has been found to hold good when first one variety and then several mixed varieties of wheat have been sown on equal spaces of ground. Hence, if any one species of grass were to go on varying, and those varieties were continually selected which differed from each other in at all the same manner as distinct species and genera of grasses differ from each other, a greater number of individual plants of this species of grass, including its modified descendants, would succeed in living on the same piece of ground. And we well know that each species and each variety of grass is annually sowing almost countless seeds; and thus, as it may be said, is striving its utmost to increase its numbers. Consequently, I cannot doubt that in the course of many thousands of generations, the most distinct varieties of any one species of grass would always have the best chance of succeeding and of increasing in numbers, and thus of supplanting the less distinct varieties; and varieties, when rendered very distinct from each other, take the rank of species.The truth of the principle, that the greatest amount of life can be supported by great diversification of structure, is seen under many natural circumstances. In an extremely small area, especially if freely open to immigration, and where the contest between individual and individual must be severe, we always find great diversity in its inhabitants. For instance, I found that a piece of turf, three feet by four in size, which had been exposed for many years to exactly the same conditions, supported twenty species of plants, and these belonged to eighteen genera and to eight orders, which shows how much these plants differed from each other. So it is with the plants and insects on small and uniform islets; and so in small ponds of fresh water. Farmers find that they can raise most food by a rotation of plants belonging to the most different orders: nature follows what may be called a simultaneous rotation. Most of the animals and plants which live close round any small piece of ground, could live on it (supposing it not to be in any way peculiar in its nature), and may be said to be striving to the utmost to live there; but, it is seen, that where they come into the closest competition with each other, the advantages of diversification of structure, with the accompanying differences of habit and constitution, determine that the inhabitants, which thus jostle each other most closely, shall, as a general rule, belong to what we call different genera and orders.The same principle is seen in the naturalisation of plants through man's agency in foreign lands. It might have been expected that the plants which have succeeded in becoming naturalised in any land would generally have been closely allied to the indigenes; for these are commonly looked at as specially created and adapted for their own country. It might, also, perhaps have been expected that naturalised plants would have belonged to a few groups more especially adapted to certain stations in their new homes. But the case is very different; and Alph. De Candolle has well remarked in his great and admirable work, that floras gain by naturalisation, proportionally with the number of the native genera and species, far more in new genera than in new species. To give a single instance: in the last edition of Dr Asa Gray's 'Manual of the Flora of the Northern United States,' 260 naturalised plants are enumerated, and these belong to 162 genera. We thus see that these naturalised plants are of a highly diversified nature. They differ, moreover, to a large extent from the indigenes, for out of the 162 genera, no less than 100 genera are not there indigenous, and thus a large proportional addition is made to the genera of these States.By considering the nature of the plants or animals which have struggled successfully with the indigenes of any country, and have there become naturalised, we can gain some crude idea in what manner some of the natives would have had to be modified, in order to have gained an advantage over the other natives; and we may, I think, at least safely infer that diversification of structure, amounting to new generic differences, would have been profitable to them.

  • 古正举 08-05

       The eyes of moles and of some burrowing rodents are rudimentary in size, and in some cases are quite covered up by skin and fur. This state of the eyes is probably due to gradual reduction from disuse, but aided perhaps by natural selection. In South America, a burrowing rodent, the tuco-tuco, or Ctenomys, is even more subterranean in its habits than the mole; and I was assured by a Spaniard, who had often caught them, that they were frequently blind; one which I kept alive was certainly in this condition, the cause, as appeared on dissection, having been inflammation of the nictitating membrane. As frequent inflammation of the eyes must be injurious to any animal, and as eyes are certainly not indispensable to animals with subterranean habits, a reduction in their size with the adhesion of the eyelids and growth of fur over them, might in such case be an advantage; and if so, natural selection would constantly aid the effects of disuse.

  • 宋太平 08-03

    {  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.

  • 欧建明 08-03

      The two first heads shall be here d

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