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1.【址:a g 9 559⒐ v i p】1  This subject will be more fully discussed in our chapter on Geology; but it must be here alluded to from being intimately connected with natural selection. Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure. But as from the high geometrical powers of increase of all organic beings, each area is already fully stocked with inhabitants, it follows that as each selected and favoured form increases in number, so will the less favoured forms decrease and become rare. Rarity, as geology tells us, is the precursor to extinction. We can, also, see that any form represented by few individuals will, during fluctuations in the seasons or in the number of its enemies, run a good chance of utter extinction. But we may go further than this; for as new forms are continually and slowly being produced, unless we believe that the number of specific forms goes on perpetually and almost indefinitely increasing, numbers inevitably must become extinct. That the number of specific forms has not indefinitely increased, geology shows us plainly; and indeed we can see reason why they should not have thus increased, for the number of places in the polity of nature is not indefinitely great, not that we have any means of knowing that any one region has as yet got its maximum of species. probably no region is as yet fully stocked, for at the Cape of Good Hope, where more species of plants are crowded together than in any other quarter of the world, some foreign plants have become naturalised, without causing, as far as we know, the extinction of any natives.Furthermore, the species which are most numerous in individuals will have the best chance of producing within any given period favourable variations. We have evidence of this, in the facts given in the second chapter, showing that it is the common species which afford the greatest number of recorded varieties, or incipient species. Hence, rare species will be less quickly modified or improved within any given period, and they will consequently be beaten in the race for life by the modified descendants of the commoner species.
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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.  The Origin of Species
5.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.
6.  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.

计划指导

1.  Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the Hon. W. Elliot from India, and by the Hon. C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerably antiquity. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs. The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head, and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular and strictly inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels. The runt is a bird of great size, with long, massive beak and large feet; some of the sub-breeds of runts have very long necks, others very long wings and tails, others singularly short tails. The barb is allied to the carrier, but, instead of a very long beak, has a very short and very broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter. The turbit has a very short and conical beak, with a line of reversed feathers down the breast; and it has the habit of continually expanding slightly the upper part of the oesophagus. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood, and it has, proportionally to its size, much elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds. The fantail has thirty or even forty tail-feathers, instead of twelve or fourteen, the normal number in all members of the great pigeon family; and these feathers are kept expanded, and are carried so erect that in good birds the head and tail touch; the oil-gland is quite aborted. Several other less distinct breeds might have been specified.In the skeletons of the several breeds, the development of the bones of the face in length and breadth and curvature differs enormously. The shape, as well as the breadth and length of the ramus of the lower jaw, varies in a highly remarkable manner. The number of the caudal and sacral vertebrae vary; as does the number of the ribs, together with their relative breadth and the presence of processes. The size and shape of the apertures in the sternum are highly variable; so is the degree of divergence and relative size of the two arms of the furcula. The proportional width of the gape of mouth, the proportional length of the eyelids, of the orifice of the nostrils, of the tongue (not always in strict correlation with the length of beak), the size of the crop and of the upper part of the oesophagus; the development and abortion of the oil-gland; the number of the primary wing and caudal feathers; the relative length of wing and tail to each other and to the body; the relative length of leg and of the feet; the number of scutellae on the toes, the development of skin between the toes, are all points of structure which are variable. The period at which the perfect plumage is acquired varies, as does the state of the down with which the nestling birds are clothed when hatched. The shape and size of the eggs vary. The manner of flight differs remarkably; as does in some breeds the voice and disposition. Lastly, in certain breeds, the males and females have come to differ to a slight degree from each other.Altogether at least a score of pigeons might be chosen, which if shown to an ornithologist, and he were told that they were wild birds, would certainly, I think, be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail in the same genus; more especially as in each of these breeds several truly-inherited sub-breeds, or species as he might have called them, could be shown him.
2.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.
3.  To test the truth of this anticipation I have arranged the plants of twelve countries, and the coleopterous insects of two districts, into two nearly equal masses, the species of the larger genera on one side, and those of the smaller genera on the other side, and it has invariably proved to be the case that a larger proportion of the species on the side of the larger genera present varieties, than on the side of the smaller genera. Moreover, the species of the large genera which present any varieties, invariably present a larger average number of varieties than do the species of the small genera. Both these results follow when another division is made, and when all the smallest genera, with from only one to four species, are absolutely excluded from the tables. These facts are of plain signification on the view that species are only strongly marked and permanent varieties; for whenever many species of the same genus have been formed, or where, if we may use the expression, the manufactory of species has been active, we ought generally to find the manufactory still in action, more especially as we have every reason to believe the process of manufacturing new species to be a slow one. And this certainly is the case, if varieties be looked at as incipient species; for my tables clearly show as a general rule that, wherever many species of a genus have been formed, the species of that genus present a number of varieties, that is of incipient species, beyond the average. It is not that all large genera are now varying much, and are thus increasing in the number of their species, or that no small genera are now varying and increasing; for if this had been so, it would have been fatal to my theory; inasmuch as geology plainly tells us that small genera have in the lapse of time often increased greatly in size; and that large genera have often come to their maxima, declined, and disappeared. All that we want to show is, that where many species of a genus have been formed, on an average many are still forming; and this holds good.There are other relations between the species of large genera and their recorded varieties which deserve notice. We have seen that there is no infallible criterion by which to distinguish species and well-marked varieties; and in those cases in which intermediate links have not been found between doubtful forms, naturalists are compelled to come to a determination by the amount of difference between them, judging by analogy whether or not the amount suffices to raise one or both to the rank of species. Hence the amount of difference is one very important criterion in settling whether two forms should be ranked as species or varieties. Now Fries has remarked in regard to plants, and Westwood in regard to insects, that in large genera the amount of difference between the species is often exceedingly small. I have endeavoured to test this numerically by averages, and, as far as my imperfect results go, they always confirm the view. I have also consulted some sagacious and most experienced observers, and, after deliberation, they concur in this view. In this respect, therefore, the species of the larger genera resemble varieties, more than do the species of the smaller genera. Or the case may be put in another way, and it may be said, that in the larger genera, in which a number of varieties or incipient species greater than the average are now manufacturing, many of the species already manufactured still to a certain extent resemble varieties, for they differ from each other by a less than usual amount of difference.Moreover, the species of the large genera are related to each other, in the same manner as the varieties of any one species are related to each other. No naturalist pretends that all the species of a genus are equally distinct from each other; they may generally be divided into sub-genera, or sections, or lesser groups. As Fries has well remarked, little groups of species are generally clustered like satellites around certain other species. And what are varieties but groups of forms, unequally related to each other, and clustered round certain forms that is, round their parent-species? Undoubtedly there is one most important point of difference between varieties and species; namely, that the amount of difference between varieties, when compared with each other or with their parent-species, is much less than that between the species of the same genus. But when we come to discuss the principle, as I call it, of Divergence of Character, we shall see how this may be explained, and how the lesser differences between varieties will tend to increase into the greater differences between species.There is one other point which seems to me worth notice. Varieties generally have much restricted ranges: this statement is indeed scarcely more than a truism, for if a variety were found to have a wider range than that of its supposed parent-species, their denominations ought to be reversed. But there is also reason to believe, that those species which are very closely allied to other species, and in so far resemble varieties, often have much restricted ranges. For instance, Mr H. C. Watson has marked for me in the well-sifted London Catalogue of plants (4th edition) 63 plants which are therein ranked as species, but which he considers as so closely allied to other species as to be of doubtful value: these 63 reputed species range on an average over 6.9 of the provinces into which Mr Watson has divided Great Britain. Now, in this same catalogue, 53 acknowledged varieties are recorded, and these range over 7.7 provinces; whereas, the species to which these varieties belong range over 14.3 provinces. So that the acknowledged varieties have very nearly the same restricted average range, as have those very closely allied forms, marked for me by Mr Watson as doubtful species, but which are almost universally ranked by British botanists as good and true species.Finally, then, varieties have the same general characters as species, for they cannot be distinguished from species, except, firstly, by the discovery of intermediate linking forms, and the occurrence of such links cannot affect the actual characters of the forms which they connect; and except, secondly, by a certain amount of difference, for two forms, if differing very little, are generally ranked as varieties, notwithstanding that intermediate linking forms have not been discovered; but the amount of difference considered necessary to give to two forms the rank of species is quite indefinite. In genera having more than the average number of species in any country, the species of these genera have more than the average number of varieties. In large genera the species are apt to be closely, but unequally, allied together, forming little clusters round certain species. Species very closely allied to other species apparently have restricted ranges. In all these several respects the species of large genera present a strong analogy with varieties. And we can clearly understand these analogies, if species have once existed as varieties, and have thus originated: whereas, these analogies are utterly inexplicable if each species has been independently created.We have, also, seen that it is the most flourishing and dominant species of the larger genera which on an average vary most; and varieties, as we shall hereafter see, tend to become converted into new and distinct species. The larger genera thus tend to become larger; and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. But by steps hereafter to be explained, the larger genera also tend to break up into smaller genera. And thus, the forms of life throughout the universe become divided into groups subordinate to groups.
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.  I mean by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood. The most obvious case is, that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded, affect the structure of the adult; in the same manner as any malconformation affecting the early embryo, seriously affects the whole organisation of the adult. The several parts of the body which are homologous, and which, at an early embryonic period, are alike, seem liable to vary in an allied manner: we see this in the right and left sides of the body varying in the same manner; in the front and hind legs, and even in the jaws and limbs, varying together, for the lower jaw is believed to be homologous with the limbs. These tendencies, I do not doubt, may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side; and if this had been of any great use to the breed it might probably have been rendered permanent by natural selection.Homologous parts, as has been remarked by some authors, tend to cohere; this is often seen in monstrous plants; and nothing is more common than the union of homologous parts in normal structures, as the union of the petals of the corolla into a tube. Hard parts seem to affect the form of adjoining soft parts; it is believed by some authors that the diversity in the shape of the pelvis in birds causes the remarkable diversity in the shape of their kidneys. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. In snakes, according to Schlegel, the shape of the body and the manner of swallowing determine the position of several of the most important viscera.
6.  by Charles Darwin

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1.  This is an extremely intricate subject. A large amount of inheritable and diversified variability is favourable, but I believe mere individual differences suffice for the work. A large number of individuals, by giving a better chance for the appearance within any given period of profitable variations, will compensate for a lesser amount of variability in each individual, and is, I believe, an extremely important element of success. Though nature grants vast periods of time for the work of natural selection, she does not grant an indefinite period; for as all organic beings are striving, it may be said, to seize on each place in the economy of nature, if any one species does not become modified and improved in a corresponding degree with its competitors, it will soon be exterminated.
2.  I see no reason to limit the process of modification, as now explained, to the formation of genera alone. If, in our diagram, we suppose the amount of change represented by each successive group of diverging dotted lines to be very great, the forms marked a214 to p14, those marked b14 and f14, and those marked o14 to m14, will form three very distinct genera. We shall also have two very distinct genera descended from (I) and as these latter two genera, both from continued divergence of character and from inheritance from a different parent, will differ widely from the three genera descended from (A), the two little groups of genera will form two distinct families, or even orders, according to the amount of divergent modification supposed to be represented in the diagram. And the two new families, or orders, will have descended from two species of the original genus; and these two species are supposed to have descended from one species of a still more ancient and unknown genus.
3.  Previous Chapter
4.  When a young naturalist commences the study of a group of organisms quite unknown to him, he is at first much perplexed to determine what differences to consider as specific, and what as varieties; for he knows nothing of the amount and kind of variation to which the group is subject; and this shows, at least, how very generally there is some variation. But if he confine his attention to one class within one country, he will soon make up his mind how to rank most of the doubtful forms. His general tendency will be to make many species, for he will become impressed, just like the pigeon or poultry-fancier before alluded to, with the amount of difference in the forms which he is continually studying; and he has little general knowledge of analogical variation in other groups and in other countries, by which to correct his first impressions. As he extends the range of his observations, he will meet with more cases of difficulty; for he will encounter a greater number of closely-allied forms. But if his observations be widely extended, he will in the end generally be enabled to make up his own mind which to call varieties and which species; but he will succeed in this at the expense of admitting much variation, and the truth of this admission will often be disputed by other naturalists. When, moreover, he comes to study allied forms brought from countries not now continuous, in which case he can hardly hope to find the intermediate links between his doubtful forms, he will have to trust almost entirely to analogy, and his difficulties will rise to a climax.Certainly no clear line of demarcation has as yet been drawn between species and sub-species that is, the forms which in the opinion of some naturalists come very near to, but do not quite arrive at the rank of species; or, again, between sub-species and well-marked varieties, or between lesser varieties and individual differences. These differences blend into each other in an insensible series; and a series impresses the mind with the idea of an actual passage.
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.  These propositions will be most readily understood by looking to our domestic races. The most distinct breeds of pigeons, in countries most widely apart, present sub-varieties with reversed feathers on the head and feathers on the feet, characters not possessed by the aboriginal rock-pigeon; these then are analogous variations in two or more distinct races. The frequent presence of fourteen or even sixteen tail-feathers in the pouter, may be considered as a variation representing the normal structure of another race, the fantail. I presume that no one will doubt that all such analogous variations are due to the several races of the pigeon having inherited from a common parent the same constitution and tendency to variation, when acted on by similar unknown influences. In the vegetable kingdom we have a case of analogous variation, in the enlarged stems, or roots as commonly called, of the Swedish turnip and Ruta baga, plants which several botanists rank as varieties produced by cultivation from a common parent: if this be not so, the case will then be one of analogous variation in two so-called distinct species; and to these a third may be added, namely, the common turnip. According to the ordinary view of each species having been independently created, we should have to attribute this similarity in the enlarged stems of these three plants, not to the vera causa of community of descent, and a consequent tendency to vary in a like manner, but to three separate yet closely related acts of creation.With pigeons, however, we have another case, namely, the occasional appearance in all the breeds, of slaty-blue birds with two black bars on the wings, a white rump, a bar at the end of the tail, with the outer feathers externally edged near their bases with white. As all these marks are characteristic of the parent rock-pigeon, I presume that no one will doubt that this is a case of reversion, and not of a new yet analogous variation appearing in the several breeds. We may I think confidently come to this conclusion, because, as we have seen, these coloured marks are eminently liable to appear in the crossed offspring of two distinct and differently coloured breeds; and in this case there is nothing in the external conditions of life to cause the reappearance of the slaty-blue, with the several marks, beyond the influence of the mere act of crossing on the laws of inheritance.

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1.  If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many variations have occurred useful to man. But if variations useful to any organic being do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection. Natural selection, on the principle of qualities being inherited at corresponding ages, can modify the egg, seed, or young, as easily as the adult. Amongst many animals, sexual selection will give its aid to ordinary selection, by assuring to the most vigorous and best adapted males the greatest number of offspring. Sexual selection will also give characters useful to the males alone, in their struggles with other males.Whether natural selection has really thus acted in nature, in modifying and adapting the various forms of life to their several conditions and stations, must be judged of by the general tenour and balance of evidence given in the following chapters. But we already see how it entails extinction; and how largely extinction has acted in the world's history, geology plainly declares. Natural selection, also, leads to divergence of character; for more living beings can be supported on the same area the more they diverge in structure, habits, and constitution, of which we see proof by looking at the inhabitants of any small spot or at naturalised productions. Therefore during the modification of the descendants of any one species, and during the incessant struggle of all species to increase in numbers, the more diversified these descendants become, the better will be their chance of succeeding in the battle of life. Thus the small differences distinguishing varieties of the same species, will steadily tend to increase till they come to equal the greater differences between species of the same genus, or even of distinct genera.We have seen that it is the common, the widely-diffused, and widely-ranging species, belonging to the larger genera, which vary most; and these will tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. Natural selection, as has just been remarked, leads to divergence of character and to much extinction of the less improved and intermediate forms of life. On these principles, I believe, the nature of the affinities of all organic beings may be explained. It is a truly wonderful fact the wonder of which we are apt to overlook from familiarity that all animals and all plants throughout all time and space should be related to each other in group subordinate to group, in the manner which we everywhere behold namely, varieties of the same species most closely related together, species of the same genus less closely and unequally related together, forming sections and sub-genera, species of distinct genera much less closely related, and genera related in different degrees, forming sub-families, families, orders, sub-classes, and classes. The several subordinate groups in any class cannot be ranked in a single file, but seem rather to be clustered round points, and these round other points, and so on in almost endless cycles. On the view that each species has been independently created, I can see no explanation of this great fact in the classification of all organic beings; but, to the best of my judgment, it is explained through inheritance and the complex action of natural selection, entailing extinction and divergence of character, as we have seen illustrated in the diagram.The affinities of all the beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth. The green and budding twigs may represent existing species; and those produced during each former year may represent the long succession of extinct species. At each period of growth all the growing twigs have tried to branch out on all sides, and to overtop and kill the surrounding twigs and branches, in the same manner as species and groups of species have tried to overmaster other species in the great battle for life. The limbs divided into great branches, and these into lesser and lesser branches, were themselves once, when the tree was small, budding twigs; and this connexion of the former and present buds by ramifying branches may well represent the classification of all extinct and living species in groups subordinate to groups. Of the many twigs which flourished when the tree was a mere bush, only two or three, now grown into great branches, yet survive and bear all the other branches; so with the species which lived during long-past geological periods, very few now have living and modified descendants. From the first growth of the tree, many a limb and branch has decayed and dropped off; and these lost branches of various sizes may represent those whole orders, families, and genera which have now no living representatives, and which are known to us only from having been found in a fossil state. As we here and there see a thin straggling branch springing from a fork low down in a tree, and which by some chance has been favoured and is still alive on its summit, so we occasionally see an animal like the Ornithorhynchus or Lepidosiren, which in some small degree connects by its affinities two large branches of life, and which has apparently been saved from fatal competition by having inhabited a protected station. As buds give rise by growth to fresh buds, and these, if vigorous, branch out and overtop on all sides many a feebler branch, so by generation I believe it has been with the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever branching and beautiful ramifications.
2.  In plants the same gradual process of improvement, through the occasional preservation of the best individuals, whether or not sufficiently distinct to be ranked at their first appearance as distinct varieties, and whether or not two or more species or races have become blended together by crossing, may plainly be recognised in the increased size and beauty which we now see in the varieties of the heartsease, rose, pelargonium, dahlia, and other plants, when compared with the older varieties or with their parent-stocks. No one would ever expect to get a first-rate heartsease or dahlia from the seed of a wild plant. No one would expect to raise a first-rate melting pear from the seed of a wild pear, though he might succeed from a poor seedling growing wild, if it had come from a garden-stock. The pear, though cultivated in classical times, appears, from Pliny's description, to have been a fruit of very inferior quality. I have seen great surprise expressed in horticultural works at the wonderful skill of gardeners, in having produced such splendid results from such poor materials; but the art, I cannot doubt, has been simple, and, as far as the final result is concerned, has been followed almost unconsciously. It has consisted in always cultivating the best known variety, sowing its seeds, and, when a slightly better variety has chanced to appear, selecting it, and so onwards. But the gardeners of the classical period, who cultivated the best pear they could procure, never thought what splendid fruit we should eat; though we owe our excellent fruit, in some small degree, to their having naturally chosen and preserved the best varieties they could anywhere find.A large amount of change in our cultivated plants, thus slowly and unconsciously accumulated, explains, as I believe, the well-known fact, that in a vast number of cases we cannot recognise, and therefore do not know, the wild parent-stocks of the plants which have been longest cultivated in our flower and kitchen gardens. If it has taken centuries or thousands of years to improve or modify most of our plants up to their present standard of usefulness to man, we can understand how it is that neither Australia, the Cape of Good Hope, nor any other region inhabited by quite uncivilised man, has afforded us a single plant worth culture. It is not that these countries, so rich in species, do not by a strange chance possess the aboriginal stocks of any useful plants, but that the native plants have not been improved by continued selection up to a standard of perfection comparable with that given to the plants in countries anciently civilised.
3.  In the case of most of our anciently domesticated animals and plants, I do not think it is possible to come to any definite conclusion, whether they have descended from one or several species. The argument mainly relied on by those who believe in the multiple origin of our domestic animals is, that we find in the most ancient records, more especially on the monuments of Egypt, much diversity in the breeds; and that some of the breeds closely resemble, perhaps are identical with, those still existing. Even if this latter fact were found more strictly and generally true than seems to me to be the case, what does it show, but that some of our breeds originated there, four or five thousand years ago? But Mr Horner's researches have rendered it in some degree probable that man sufficiently civilized to have manufactured pottery existed in the valley of the Nile thirteen or fourteen thousand years ago; and who will pretend to say how long before these ancient periods, savages, like those of Tierra del Fuego or Australia, who possess a semi-domestic dog, may not have existed in Egypt?The whole subject must, I think, remain vague; nevertheless, I may, without here entering on any details, state that, from geographical and other considerations, I think it highly probable that our domestic dogs have descended from several wild species. In regard to sheep and goats I can form no opinion. I should think, from facts communicated to me by Mr Blyth, on the habits, voice, and constitution, &c., of the humped Indian cattle, that these had descended from a different aboriginal stock from our European cattle; and several competent judges believe that these latter have had more than one wild parent. With respect to horses, from reasons which I cannot give here, I am doubtfully inclined to believe, in opposition to several authors, that all the races have descended from one wild stock. Mr Blyth, whose opinion, from his large and varied stores of knowledge, I should value more than that of almost any one, thinks that all the breeds of poultry have proceeded from the common wild Indian fowl (Gallus bankiva). In regard to ducks and rabbits, the breeds of which differ considerably from each other in structure, I do not doubt that they all have descended from the common wild duck and rabbit.The doctrine of the origin of our several domestic races from several aboriginal stocks, has been carried to an absurd extreme by some authors. They believe that every race which breeds true, let the distinctive characters be ever so slight, has had its wild prototype. At this rate there must have existed at least a score of species of wild cattle, as many sheep, and several goats in Europe alone, and several even within Great Britain. One author believes that there formerly existed in Great Britain eleven wild species of sheep peculiar to it! When we bear in mind that Britain has now hardly one peculiar mammal, and France but few distinct from those of Germany and conversely, and so with Hungary, Spain, &c., but that each of these kingdoms possesses several peculiar breeds of cattle, sheep, &c., we must admit that many domestic breeds have originated in Europe; for whence could they have been derived, as these several countries do not possess a number of peculiar species as distinct parent-stocks? So it is in India. Even in the case of the domestic dogs of the whole world, which I fully admit have probably descended from several wild species, I cannot doubt that there has been an immense amount of inherited variation. Who can believe that animals closely resembling the Italian greyhound, the bloodhound, the bull-dog, or Blenheim spaniel, &c. so unlike all wild Canidae ever existed freely in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species; but by crossing we can get only forms in some degree intermediate between their parents; and if we account for our several domestic races by this process, we must admit the former existence of the most extreme forms, as the Italian greyhound, bloodhound, bull-dog, &c., in the wild state. Moreover, the possibility of making distinct races by crossing has been greatly exaggerated. There can be no doubt that a race may be modified by occasional crosses, if aided by the careful selection of those individual mongrels, which present any desired character; but that a race could be obtained nearly intermediate between two extremely different races or species, I can hardly believe. Sir J. Sebright expressly experimentised for this object, and failed. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) extremely uniform, and everything seems simple enough; but when these mongrels are crossed one with another for several generations, hardly two of them will be alike, and then the extreme difficulty, or rather utter hopelessness, of the task becomes apparent. Certainly, a breed intermediate between two very distinct breeds could not be got without extreme care and long-continued selection; nor can I find a single case on record of a permanent race having been thus formed.On the Breeds of the Domestic pigeon.
4、  When a variation is of the slightest use to a being, we cannot tell how much of it to attribute to the accumulative action of natural selection, and how much to the conditions of life. Thus, it is well known to furriers that animals of the same species have thicker and better fur the more severe the climate is under which they have lived; but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations, and how much to the direct action of the severe climate? for it would appear that climate has some direct action on the hair of our domestic quadrupeds.
5、  Some facts in regard to the colouring of pigeons well deserve consideration. The rock-pigeon is of a slaty-blue, and has a white rump (the Indian sub-species, C. intermedia of Strickland, having it bluish); the tail has a terminal dark bar, with the bases of the outer feathers externally edged with white; the wings have two black bars: some semi-domestic breeds and some apparently truly wild breeds have, besides the two black bars, the wings chequered with black. These several marks do not occur together in any other species of the whole family. Now, in every one of the domestic breeds, taking thoroughly well-bred birds, all the above marks, even to the white edging of the outer tail-feathers, sometimes concur perfectly developed. Moreover, when two birds belonging to two distinct breeds are crossed, neither of which is blue or has any of the above-specified marks, the mongrel offspring are very apt suddenly to acquire these characters; for instance, I crossed some uniformly white fantails with some uniformly black barbs, and they produced mottled brown and black birds; these I again crossed together, and one grandchild of the pure white fantail and pure black barb was of as beautiful a blue colour, with the white rump, double black wing-bar, and barred and white-edged tail-feathers, as any wild rock-pigeon! We can understand these facts, on the well-known principle of reversion to ancestral characters, if all the domestic breeds have descended from the rock-pigeon. But if we deny this, we must make one of the two following highly improbable suppositions. Either, firstly, that all the several imagined aboriginal stocks were coloured and marked like the rock-pigeon, although no other existing species is thus coloured and marked, so that in each separate breed there might be a tendency to revert to the very same colours and markings. Or, secondly, that each breed, even the purest, has within a dozen or, at most, within a score of generations, been crossed by the rock-pigeon: I say within a dozen or twenty generations, for we know of no fact countenancing the belief that the child ever reverts to some one ancestor, removed by a greater number of generations. In a breed which has been crossed only once with some distinct breed, the tendency to reversion to any character derived from such cross will naturally become less and less, as in each succeeding generation there will be less of the foreign blood; but when there has been no cross with a distinct breed, and there is a tendency in both parents to revert to a character, which has been lost during some former generation, this tendency, for all that we can see to the contrary, may be transmitted undiminished for an indefinite number of generations. These two distinct cases are often confounded in treatises on inheritance.Lastly, the hybrids or mongrels from between all the domestic breeds of pigeons are perfectly fertile. I can state this from my own observations, purposely made on the most distinct breeds. Now, it is difficult, perhaps impossible, to bring forward one case of the hybrid offspring of two animals clearly distinct being themselves perfectly fertile. Some authors believe that long-continued domestication eliminates this strong tendency to sterility: from the history of the dog I think there is some probability in this hypothesis, if applied to species closely related together, though it is unsupported by a single experiment. But to extend the hypothesis so far as to suppose that species, aboriginally as distinct as carriers, tumblers, pouters, and fantails now are, should yield offspring perfectly fertile, inter se, seems to me rash in the extreme.

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  • 石军慧 08-08

      Now let us turn to the effects of crossing the several species of the horse-genus. Rollin asserts, that the common mule from the ass and horse is particularly apt to have bars on its legs. I once saw a mule with its legs so much striped that any one at first would have thought that it must have been the product of a zebra; and Mr. W. C. Martin, in his excellent treatise on the horse, has given a figure of a similar mule. In four coloured drawings, which I have seen, of hybrids between the ass and zebra, the legs were much more plainly barred than the rest of the body; and in one of them there was a double shoulder-stripe. In Lord Moreton's famous hybrid from a chestnut mare and male quagga, the hybrid, and even the pure offspring subsequently produced from the mare by a black Arabian sire, were much more plainly barred across the legs than is even the pure quagga. Lastly, and this is another most remarkable case, a hybrid has been figured by Dr Gray (and he informs me that he knows of a second case) from the ass and the hemionus; and this hybrid, though the ass seldom has stripes on its legs and the hemionus has none and has not even a shoulder-stripe, nevertheless had all four legs barred, and had three short shoulder-stripes, like those on the dun Welch pony, and even had some zebra-like stripes on the sides of its face. With respect to this last fact, I was so convinced that not even a stripe of colour appears from what would commonly be called an accident, that I was led solely from the occurrence of the face-stripes on this hybrid from the ass and hemionus, to ask Colonel Poole whether such face-stripes ever occur in the eminently striped Kattywar breed of horses, and was, as we have seen, answered in the affirmative.What now are we to say to these several facts? We see several very distinct species of the horse-genus becoming, by simple variation, striped on the legs like a zebra, or striped on the shoulders like an ass. In the horse we see this tendency strong whenever a dun tint appears a tint which approaches to that of the general colouring of the other species of the genus. The appearance of the stripes is not accompanied by any change of form or by any other new character. We see this tendency to become striped most strongly displayed in hybrids from between several of the most distinct species. Now observe the case of the several breeds of pigeons: they are descended from a pigeon (including two or three sub-species or geographical races) of a bluish colour, with certain bars and other marks; and when any breed assumes by simple variation a bluish tint, these bars and other marks invariably reappear; but without any other change of form or character. When the oldest and truest breeds of various colours are crossed, we see a strong tendency for the blue tint and bars and marks to reappear in the mongrels. I have stated that the most probable hypothesis to account for the reappearance of very ancient characters, is that there is a tendency in the young of each successive generation to produce the long-lost character, and that this tendency, from unknown causes, sometimes prevails. And we have just seen that in several species of the horse-genus the stripes are either plainer or appear more commonly in the young than in the old. Call the breeds of pigeons, some of which have bred true for centuries, species; and how exactly parallel is the case with that of the species of the horse-genus! For myself, I venture confidently to look back thousands on thousands of generations, and I see an animal striped like a zebra, but perhaps otherwise very differently constructed, the common parent of our domestic horse, whether or not it be descended from one or more wild stocks, of the ass, the hemionus, quagga, and zebra.He who believes that each equine species was independently created, will, I presume, assert that each species has been created with a tendency to vary, both under nature and under domestication, in this particular manner, so as often to become striped like other species of the genus; and that each has been created with a strong tendency, when crossed with species inhabiting distant quarters of the world, to produce hybrids resembling in their stripes, not their own parents, but other species of the genus. To admit this view is, as it seems to me, to reject a real for an unreal, or at least for an unknown, cause. It makes the works of God a mere mockery and deception; I would almost as soon believe with the old and ignorant cosmogonists, that fossil shells had never lived, but had been created in stone so as to mock the shells now living on the sea-shore.

  • 柳重言 08-08

      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.

  • 萨维德斯 08-08

       Chapter 2 - Variation Under Nature

  • 伊丽莎白·阿普尔亚德 08-08

      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.

  • 闫慧荣 08-07

    {  Chapter 3 - Struggle for Existence

  • 潘迎紫 08-06

      Fourthly, how can we account for species, when crossed, being sterile and producing sterile offspring, whereas, when varieties are crossed, their fertility is unimpaired?}

  • 郑宪景 08-06

      Chapter 6 - Difficulties on Theory

  • 克托亚 08-06

      I HAVE hitherto sometimes spoken as if the variations so common and multiform in organic beings under domestication, and in a lesser degree in those in a state of nature had been due to chance. This, of course, is a wholly incorrect expression, but it serves to acknowledge plainly our ignorance of the cause of each particular variation. Some authors believe it to be as much the function of the reproductive system to produce individual differences, or very slight deviations of structure, as to make the child like its parents. But the much greater variability, as well as the greater frequency of monstrosities, under domestication or cultivation, than under nature, leads me to believe that deviations of structure are in some way due to the nature of the conditions of life, to which the parents and their more remote ancestors have been exposed during several generations. I have remarked in the first chapter but a long catalogue of facts which cannot be here given would be necessary to show the truth of the remark that the reproductive system is eminently susceptible to changes in the conditions of life; and to this system being functionally disturbed in the parents, I chiefly attribute the varying or plastic condition of the offspring. The male and female sexual elements seem to be affected before that union takes place which is to form a new being. In the case of 'sporting' plants, the bud, which in its earliest condition does not apparently differ essentially from an ovule, is alone affected. But why, because the reproductive system is disturbed, this or that part should vary more or less, we are profoundly ignorant. Nevertheless, we can here and there dimly catch a faint ray of light, and we may feel sure that there must be some cause for each deviation of structure, however slight.How much direct effect difference of climate, food, &c., produces on any being is extremely doubtful. My impression is, that the effect is extremely small in the case of animals, but perhaps rather more in that of plants. We may, at least, safely conclude that such influences cannot have produced the many striking and complex co-adaptations of structure between one organic being and another, which we see everywhere throughout nature. Some little influence may be attributed to climate, food, &c.: thus, E. Forbes speaks confidently that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species further north or from greater depths. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere, than when living on islands or near the coast. So with insects, Wollaston is convinced that residence near the sea affects their colours. Moquin-Tandon gives a list of plants which when growing near the sea-shore have their leaves in some degree fleshy, though not elsewhere fleshy. Several other such cases could be given.The fact of varieties of one species, when they range into the zone of habitation of other species, often acquiring in a very slight degree some of the characters of such species, accords with our view that species of all kinds are only well-marked and permanent varieties. Thus the species of shells which are confined to tropical and shallow seas are generally brighter-coloured than those confined to cold and deeper seas. The birds which are confined to continents are, according to Mr Gould, brighter-coloured than those of islands. The insect-species confined to sea-coasts, as every collector knows, are often brassy or lurid. Plants which live exclusively on the sea-side are very apt to have fleshy leaves. He who believes in the creation of each species, will have to say that this shell, for instance, was created with bright colours for a warm sea; but that this other shell became bright-coloured by variation when it ranged into warmer or shallower waters.

  • 苗剑 08-05

       Guided by theoretical considerations, I thought that some interesting results might be obtained in regard to the nature and relations of the species which vary most, by tabulating all the varieties in several well-worked floras. At first this seemed a simple task; but Mr H. C. Watson, to whom I am much indebted for valuable advice and assistance on this subject, soon convinced me that there were many difficulties, as did subsequently Dr Hooker, even in stronger terms. I shall reserve for my future work the discussion of these difficulties, and the tables themselves of the proportional numbers of the varying species. Dr Hooker permits me to add, that after having carefully read my manuscript, and examined the tables, he thinks that the following statements are fairly well established. The whole subject, however, treated as it necessarily here is with much brevity, is rather perplexing, and allusions cannot be avoided to the 'struggle for existence,' 'divergence of character,' and other questions, hereafter to be discussed.

  • 吴树强 08-03

    {  If there exist savages so barbarous as never to think of the inherited character of the offspring of their domestic animals, yet any one animal particularly useful to them, for any special purpose, would be carefully preserved during famines and other accidents, to which savages are so liable, and such choice animals would thus generally leave more offspring than the inferior ones; so that in this case there would be a kind of unconscious selection going on. We see the value set on animals even by the barbarians of Tierra del Fuego, by their killing and devouring their old women, in times of dearth, as of less value than their dogs.

  • 张珮华 08-03

      It is good thus to try in our imagination to give any form some advantage over another. Probably in no single instance should we know what to do, so as to succeed. It will convince us of our ignorance on the mutual relations of all organic beings; a conviction as necessary, as it seems to be difficult to acquire. All that we can do, is to keep steadily in mind that each organic being is striving to increase at a geometrical ratio; that each at some period of its life, during some season of the year, during each generation or at intervals, has to struggle for life, and to suffer great destruction. When we reflect on this struggle, we may console ourselves with the full belief, that the war of nature is not incessant, that no fear is felt, that death is generally prompt, and that the vigorous, the healthy, and the happy survive and multiply.

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