It may be objected that the principle of selection has been reduced to methodical practice for scarcely more than three-quarters of a century; it has certainly been more attended to of late years, and many treatises have been published on the subject; and the result has been, in a corresponding degree, rapid and important. But it is very far from true that the principle is a modern discovery. I could give several references to works of high antiquity, in which the full importance of the principle is acknowledged. In rude and barbarous periods of English history choice animals were often imported, and laws were passed to prevent their exportation: the destruction of horses under a certain size was ordered, and this may be compared to the "roguing" of plants by nurserymen. The principle of selection I find distinctly given in an ancient Chinese encyclopaedia. Explicit rules are laid down by some of the Roman classical writers. From passages in Genesis, it is clear that the colour of domestic animals was at that early period attended to. Savages now sometimes cross their dogs with wild canine animals, to improve the breed, and they formerly did so, as is attested by passages in Pliny. The savages in South Africa match their draught cattle by colour, as do some of the Esquimaux their teams of dogs. Livingstone states that good domestic breeds are highly valued by the negroes in the interior of Africa who have not associated with Europeans. Some of these facts do not show actual selection, but they show that the breeding of domestic animals was carefully attended to in ancient times, and is now attended to by the lowest savages. It would, indeed, have been a strange fact, had attention not been paid to breeding, for the inheritance of good and bad qualities is so obvious.

UNCONSCIOUS SELECTION.

At the present time, eminent breeders try by methodical selection, with a distinct object in view, to make a new strain or sub-breed, superior to anything of the kind in the country. But, for our purpose, a form of selection, which may be called unconscious, and which results from every one trying to possess and breed from the best individual animals, is more important. Thus, a man who intends keeping pointers naturally tries to get as good dogs as he can, and afterwards breeds from his own best dogs, but he has no wish or expectation of permanently altering the breed. Nevertheless we may infer that this process, continued during centuries, would improve and modify any breed, in the same way as Bakewell, Collins, etc., by this very same process, only carried on more methodically, did greatly modify, even during their lifetimes, the forms and qualities of their cattle. Slow and insensible changes of this kind could never be recognised unless actual measurements or careful drawings of the breeds in question have been made long ago, which may serve for comparison. In some cases, however, unchanged, or but little changed, individuals of the same breed exist in less civilised districts, where the breed has been less improved. There is reason to believe that King Charles' spaniel has been unconsciously modified to a large extent since the time of that monarch. Some highly competent authorities are convinced that the setter is directly derived from the spaniel, and has probably been slowly altered from it. It is known that the English pointer has been greatly changed within the last century, and in this case the change has, it is believed, been chiefly effected by crosses with the foxhound; but what concerns us is, that the change has been effected unconsciously and gradually, and yet so effectually that, though the old Spanish pointer certainly came from Spain, Mr. Borrow has not seen, as I am informed by him, any native dog in Spain like our pointer.

By a similar process of selection, and by careful training, English race-horses have come to surpass in fleetness and size the parent Arabs, so that the latter, by the regulations for the Goodwood Races, are favoured in the weights which they carry. Lord Spencer and others have shown how the cattle of England have increased in weight and in early maturity, compared with the stock formerly kept in this country. By comparing the accounts given in various old treatises of the former and present state of carrier and tumbler pigeons in Britain, India, and Persia, we can trace the stages through which they have insensibly passed, and come to differ so greatly from the rock-pigeon.

Youatt gives an excellent illustration of the effects of a course of selection which may be considered as unconscious, in so far that the breeders could never have expected, or even wished, to produce the result which ensued—namely, the production of the distinct strains. The two flocks of Leicester sheep kept by Mr. Buckley and Mr. Burgess, as Mr. Youatt remarks, "Have been purely bred from the original stock of Mr. Bakewell for upwards of fifty years. There is not a suspicion existing in the mind of any one at all acquainted with the subject that the owner of either of them has deviated in any one instance from the pure blood of Mr. Bakewell's flock, and yet the difference between the sheep possessed by these two gentlemen is so great that they have the appearance of being quite different varieties."

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.

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 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 chanced to appear, selecting it, and so onwards. But the gardeners of the classical period, who cultivated the best pears which 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, thus slowly and unconsciously accumulated, explains, as I believe, the well-known fact, that in a 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 acquired by the plants in countries anciently civilised.

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

On the view here given of the important part which selection by man has played, it becomes at once obvious, how it is that our domestic races show adaptation in their structure or in their habits to man's wants or fancies. We can, I think, further understand the frequently abnormal character of our domestic races, and likewise their differences being so great in external characters, and relatively so slight in internal parts or organs. Man can hardly select, or only with much difficulty, any deviation of structure excepting such as is externally visible; and indeed he rarely cares for what is internal. He can never act by selection, excepting on variations which are first given to him in some slight degree by nature. No man would ever try to make a fantail till he saw a pigeon with a tail developed in some slight degree in an unusual manner, or a pouter till he saw a pigeon with a crop of somewhat unusual size; and the more abnormal or unusual any character was when it first appeared, the more likely it would be to catch his attention. But to use such an expression as trying to make a fantail is, I have no doubt, in most cases, utterly incorrect. The man who first selected a pigeon with a slightly larger tail, never dreamed what the descendants of that pigeon would become through long-continued, partly unconscious and partly methodical, selection. Perhaps the parent bird of all fantails had only fourteen tail-feathers somewhat expanded, like the present Java fantail, or like individuals of other and distinct breeds, in which as many as seventeen tail-feathers have been counted. Perhaps the first pouter-pigeon did not inflate its crop much more than the turbit now does the upper part of its oesophagus—a habit which is disregarded by all fanciers, as it is not one of the points of the breed.

Nor let it be thought that some great deviation of structure would be necessary to catch the fancier's eye: he perceives extremely small differences, and it is in human nature to value any novelty, however slight, in one's own possession. Nor must the value which would formerly have been set on any slight differences in the individuals of the same species, be judged of by the value which is now set on them, after several breeds have fairly been established. It is known that with pigeons many slight variations now occasionally appear, but these are rejected as faults or deviations from the standard of perfection in each breed. The common goose has not given rise to any marked varieties; hence the Toulouse and the common breed, which differ only in colour, that most fleeting of characters, have lately been exhibited as distinct at our poultry-shows.

These views appear to explain what has sometimes been noticed, namely, that we know hardly anything about the origin or history of any of our domestic breeds. But, in fact, a breed, like a dialect of a language, can hardly be said to have a distinct origin. A man preserves and breeds from an individual with some slight deviation of structure, or takes more care than usual in matching his best animals, and thus improves them, and the improved animals slowly spread in the immediate neighbourhood. But they will as yet hardly have a distinct name, and from being only slightly valued, their history will have been disregarded. When further improved by the same slow and gradual process, they will spread more widely, and will be recognised as something distinct and valuable, and will then probably first receive a provincial name. In semi-civilised countries, with little free communication, the spreading of a new sub-breed will be a slow process. As soon as the points of value are once acknowledged, the principle, as I have called it, of unconscious selection will always tend—perhaps more at one period than at another, as the breed rises or falls in fashion—perhaps more in one district than in another, according to the state of civilisation of the inhabitants—slowly to add to the characteristic features of the breed, whatever they may be. But the chance will be infinitely small of any record having been preserved of such slow, varying, and insensible changes.

CIRCUMSTANCES FAVOURABLE TO MAN'S POWER OF SELECTION.

I will now say a few words on the circumstances, favourable or the reverse, to man's power of selection. A high degree of variability is obviously favourable, as freely giving the materials for selection to work on; not that mere individual differences are not amply sufficient, with extreme care, to allow of the accumulation of a large amount of modification in almost any desired direction. But as variations manifestly useful or pleasing to man appear only occasionally, the chance of their appearance will be much increased by a large number of individuals being kept. Hence number is of the highest importance for success. On this principle Marshall formerly remarked, with respect to the sheep of part of Yorkshire, "As they generally belong to poor people, and are mostly IN SMALL LOTS, they never can be improved." On the other hand, nurserymen, from keeping large stocks of the same plant, are generally far more successful than amateurs in raising new and valuable varieties. A large number of individuals of an animal or plant can be reared only where the conditions for its propagation are favourable. When the individuals are scanty all will be allowed to breed, whatever their quality may be, and this will effectually prevent selection. But probably the most important element is that the animal or plant should be so highly valued by man, that the closest attention is paid to even the slightest deviations in its qualities or structure. Unless such attention be paid nothing can be effected. I have seen it gravely remarked, that it was most fortunate that the strawberry began to vary just when gardeners began to attend to this plant. No doubt the strawberry had always varied since it was cultivated, but the slight varieties had been neglected. As soon, however, as gardeners picked out individual plants with slightly larger, earlier, or better fruit, and raised seedlings from them, and again picked out the best seedlings and bred from them, then (with some aid by crossing distinct species) those many admirable varieties of the strawberry were raised which have appeared during the last half-century.

With animals, facility in preventing crosses is an important element in the formation of new races—at least, in a country which is already stocked with other races. In this respect enclosure of the land plays a part. Wandering savages or the inhabitants of open plains rarely possess more than one breed of the same species. Pigeons can be mated for life, and this is a great convenience to the fancier, for thus many races may be improved and kept true, though mingled in the same aviary; and this circumstance must have largely favoured the formation of new breeds. Pigeons, I may add, can be propagated in great numbers and at a very quick rate, and inferior birds may be freely rejected, as when killed they serve for food. On the other hand, cats, from their nocturnal rambling habits, can not be easily matched, and, although so much valued by women and children, we rarely see a distinct breed long kept up; such breeds as we do sometimes see are almost always imported from some other country. Although I do not doubt that some domestic animals vary less than others, yet the rarity or absence of distinct breeds of the cat, the donkey, peacock, goose, etc., may be attributed in main part to selection not having been brought into play: in cats, from the difficulty in pairing them; in donkeys, from only a few being kept by poor people, and little attention paid to their breeding; for recently in certain parts of Spain and of the United States this animal has been surprisingly modified and improved by careful selection; in peacocks, from not being very easily reared and a large stock not kept; in geese, from being valuable only for two purposes, food and feathers, and more especially from no pleasure having been felt in the display of distinct breeds; but the goose, under the conditions to which it is exposed when domesticated, seems to have a singularly inflexible organisation, though it has varied to a slight extent, as I have elsewhere described.

Some authors have maintained that the amount of variation in our domestic productions is soon reached, and can never afterward be exceeded. It would be somewhat rash to assert that the limit has been attained in any one case; for almost all our animals and plants have been greatly improved in many ways within a recent period; and this implies variation. It would be equally rash to assert that characters now increased to their utmost limit, could not, after remaining fixed for many centuries, again vary under new conditions of life. No doubt, as Mr. Wallace has remarked with much truth, a limit will be at last reached. For instance, there must be a limit to the fleetness of any terrestrial animal, as this will be determined by the friction to be overcome, the weight of the body to be carried, and the power of contraction in the muscular fibres. But what concerns us is that the domestic varieties of the same species differ from each other in almost every character, which man has attended to and selected, more than do the distinct species of the same genera. Isidore Geoffroy St. Hilaire has proved this in regard to size, and so it is with colour, and probably with the length of hair. With respect to fleetness, which depends on many bodily characters, Eclipse was far fleeter, and a dray-horse is comparably stronger, than any two natural species belonging to the same genus. So with plants, the seeds of the different varieties of the bean or maize probably differ more in size than do the seeds of the distinct species in any one genus in the same two families. The same remark holds good in regard to the fruit of the several varieties of the plum, and still more strongly with the melon, as well as in many other analogous cases.

To sum up on the origin of our domestic races of animals and plants. Changed conditions of life are of the highest importance in causing variability, both by acting directly on the organisation, and indirectly by affecting the reproductive system. It is not probable that variability is an inherent and necessary contingent, under all circumstances. The greater or less force of inheritance and reversion determine whether variations shall endure. Variability is governed by many unknown laws, of which correlated growth is probably the most important. Something, but how much we do not know, may be attributed to the definite action of the conditions of life. Some, perhaps a great, effect may be attributed to the increased use or disuse of parts. The final result is thus rendered infinitely complex. In some cases the intercrossing of aboriginally distinct species appears to have played an important part in the origin of our breeds. When several breeds have once been formed in any country, their occasional intercrossing, with the aid of selection, has, no doubt, largely aided in the formation of new sub-breeds; but the importance of crossing has been much exaggerated, both in regard to animals and to those plants which are propagated by seed. With plants which are temporarily propagated by cuttings, buds, etc., the importance of crossing is immense; for the cultivator may here disregard the extreme variability both of hybrids and of mongrels, and the sterility of hybrids; but plants not propagated by seed are of little importance to us, for their endurance is only temporary. Over all these causes of change, the accumulative action of selection, whether applied methodically and quickly, or unconsciously and slowly, but more efficiently, seems to have been the predominant power.



CHAPTER II. VARIATION UNDER NATURE.

Variability—Individual differences—Doubtful species—Wide ranging, much diffused, and common species, vary most—Species of the larger genera in each country vary more frequently than the species of the smaller genera—Many of the species of the larger genera resemble varieties in being very closely, but unequally, related to each other, and in having restricted ranges.

Before applying the principles arrived at in the last chapter to organic beings in a state of nature, we must briefly discuss whether these latter are subject to any variation. To treat this subject properly, a long catalogue of dry facts ought to be given; but these I shall reserve for a future work. Nor shall I here discuss the various definitions which have been given of the term species. No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species. Generally the term includes the unknown element of a distinct act of creation. The term "variety" is almost equally difficult to define; but here community of descent is almost universally implied, though it can rarely be proved. We have also what are called monstrosities; but they graduate into varieties. By a monstrosity I presume is meant some considerable deviation of structure, generally injurious, or not useful to the species. Some authors use the term "variation" in a technical sense, as implying a modification directly due to the physical conditions of life; and "variations" in this sense are supposed not to be inherited; but who can say that the dwarfed condition of shells in the brackish waters of the Baltic, or dwarfed plants on Alpine summits, or the thicker fur of an animal from far northwards, would not in some cases be inherited for at least a few generations? And in this case I presume that the form would be called a variety.

It may be doubted whether sudden and considerable deviations of structure, such as we occasionally see in our domestic productions, more especially with plants, are ever permanently propagated in a state of nature. Almost every part of every organic being is so beautifully related to its complex conditions of life that it seems as improbable that any part should have been suddenly produced perfect, as that a complex machine should have been invented by man in a perfect state. Under domestication monstrosities sometimes occur which resemble normal structures in widely different animals. Thus pigs have occasionally been born with a sort of proboscis, and if any wild species of the same genus had naturally possessed a proboscis, it might have been argued that this had appeared as a monstrosity; but I have as yet failed to find, after diligent search, cases of monstrosities resembling normal structures in nearly allied forms, and these alone bear on the question. If monstrous forms of this kind ever do appear in a state of nature and are capable of reproduction (which is not always the case), as they occur rarely and singly, their preservation would depend on unusually favourable circumstances. They would, also, during the first and succeeding generations cross with the ordinary form, and thus their abnormal character would almost inevitably be lost. But I shall have to return in a future chapter to the preservation and perpetuation of single or occasional variations.

INDIVIDUAL DIFFERENCES.

The many slight differences which appear in the offspring from the same parents, or which it may be presumed have thus arisen, from being observed in the individuals of the same species inhabiting the same confined locality, may be called individual differences. No one supposes that all the individuals of the same species are cast in the same actual mould. These individual differences are of the highest importance for us, for they are often inherited, as must be familiar to every one; and they thus afford materials for natural selection to act on and accumulate, in the same manner as man accumulates in any given direction individual differences in his domesticated productions. These individual differences generally affect what naturalists consider unimportant parts; but I could show, by a long catalogue of facts, that parts which must be called important, whether viewed under a physiological or classificatory point of view, sometimes vary in the individuals of the same species. I am convinced that the most experienced naturalist would be surprised at the number of the cases of variability, even in important parts of structure, which he could collect on good authority, as I have collected, during a course of years. It should be remembered that systematists are far from being pleased at finding variability in important characters, and that there are not many men who will laboriously examine internal and important organs, and compare them in many specimens of the same species. It would never have been expected that the branching of the main nerves close to the great central ganglion of an insect would have been variable in the same species; it might have been thought that changes of this nature could have been effected only by slow degrees; yet Sir J. Lubbock has shown a degree of variability in these main nerves in Coccus, which may almost be compared to the irregular branching of the stem of a tree. This philosophical naturalist, I may add, has also shown that the muscles in the larvae of certain insects are far from uniform. Authors sometimes argue in a circle when they state that important organs never vary; for these same authors practically rank those parts as important (as some few naturalists have honestly confessed) which do not vary; and, under this point of view, no instance will ever be found of an important part varying; but under any other point of view many instances assuredly can be given.

There is one point connected with individual differences which is extremely perplexing: I refer to those genera which have been called "protean" or "polymorphic," in which species present an inordinate amount of variation. With respect to many of these forms, hardly two naturalists agree whether to rank them as species or as varieties. We may instance Rubus, Rosa, and Hieracium among plants, several genera of insects, and of Brachiopod shells. In most polymorphic genera some of the species have fixed and definite characters. Genera which are polymorphic in one country seem to be, with a few exceptions, polymorphic in other countries, and likewise, judging from Brachiopod shells, at former periods of time. These facts are very perplexing, for they seem to show that this kind of variability is independent of the conditions of life. I am inclined to suspect that we see, at least in some of these polymorphic genera, variations which are of no service or disservice to the species, and which consequently have not been seized on and rendered definite by natural selection, as hereafter to be explained.

Individuals of the same species often present, as is known to every one, great differences of structure, independently of variation, as in the two sexes of various animals, in the two or three castes of sterile females or workers among insects, and in the immature and larval states of many of the lower animals. There are, also, cases of dimorphism and trimorphism, both with animals and plants. Thus, Mr. Wallace, who has lately called attention to the subject, has shown that the females of certain species of butterflies, in the Malayan Archipelago, regularly appear under two or even three conspicuously distinct forms, not connected by intermediate varieties. Fritz Muller has described analogous but more extraordinary cases with the males of certain Brazilian Crustaceans: thus, the male of a Tanais regularly occurs under two distinct forms; one of these has strong and differently shaped pincers, and the other has antennae much more abundantly furnished with smelling-hairs. Although in most of these cases, the two or three forms, both with animals and plants, are not now connected by intermediate gradations, it is possible that they were once thus connected. Mr. Wallace, for instance, describes a certain butterfly which presents in the same island a great range of varieties connected by intermediate links, and the extreme links of the chain closely resemble the two forms of an allied dimorphic species inhabiting another part of the Malay Archipelago. Thus also with ants, the several worker-castes are generally quite distinct; but in some cases, as we shall hereafter see, the castes are connected together by finely graduated varieties. So it is, as I have myself observed, with some dimorphic plants. It certainly at first appears a highly remarkable fact that the same female butterfly should have the power of producing at the same time three distinct female forms and a male; and that an hermaphrodite plant should produce from the same seed-capsule three distinct hermaphrodite forms, bearing three different kinds of females and three or even six different kinds of males. Nevertheless these cases are only exaggerations of the common fact that the female produces offspring of two sexes which sometimes differ from each other in a wonderful manner.

DOUBTFUL SPECIES.

The forms which possess in some considerable degree the character of species, but which are so closely similar to other forms, or are so closely linked to them by intermediate gradations, that naturalists do not like to rank them as distinct species, are in several respects the most important for us. We have every reason to believe that many of these doubtful and closely allied forms have permanently retained their characters for a long time; for as long, as far as we know, as have good and true species. Practically, when a naturalist can unite by means of intermediate links any two forms, he treats the one as a variety of the other, ranking the most common, but sometimes the one first described as the species, and the other as the variety. But cases of great difficulty, which I will not here enumerate, sometimes arise in deciding whether or not to rank one form as a variety of another, even when they are closely connected by intermediate links; nor will the commonly assumed hybrid nature of the intermediate forms always remove the difficulty. In very many cases, however, one form is ranked as a variety of another, not because the intermediate links have actually been found, but because analogy leads the observer to suppose either that they do now somewhere exist, or may formerly have existed; and here a wide door for the entry of doubt and conjecture is opened.

Hence, in determining whether a form should be ranked as a species or a variety, the opinion of naturalists having sound judgment and wide experience seems the only guide to follow. We must, however, in many cases, decide by a majority of naturalists, for few well-marked and well-known varieties can be named which have not been ranked as species by at least some competent judges.

That varieties of this doubtful nature are far from uncommon cannot be disputed. Compare the several floras of Great Britain, of France, or of the United States, drawn up by different botanists, and see what a surprising number of forms have been ranked by one botanist as good species, and by another as mere varieties. Mr. H.C. Watson, to whom I lie under deep obligation for assistance of all kinds, has marked for me 182 British plants, which are generally considered as varieties, but which have all been ranked by botanists as species; and in making this list he has omitted many trifling varieties, but which nevertheless have been ranked by some botanists as species, and he has entirely omitted several highly polymorphic genera. Under genera, including the most polymorphic forms, Mr. Babington gives 251 species, whereas Mr. Bentham gives only 112—a difference of 139 doubtful forms! Among animals which unite for each birth, and which are highly locomotive, doubtful forms, ranked by one zoologist as a species and by another as a variety, can rarely be found within the same country, but are common in separated areas. How many of the birds and insects in North America and Europe, which differ very slightly from each other, have been ranked by one eminent naturalist as undoubted species, and by another as varieties, or, as they are often called, geographical races! Mr. Wallace, in several valuable papers on the various animals, especially on the Lepidoptera, inhabiting the islands of the great Malayan Archipelago, shows that they may be classed under four heads, namely, as variable forms, as local forms, as geographical races or sub-species, and as true representative species. The first or variable forms vary much within the limits of the same island. The local forms are moderately constant and distinct in each separate island; but when all from the several islands are compared together, the differences are seen to be so slight and graduated that it is impossible to define or describe them, though at the same time the extreme forms are sufficiently distinct. The geographical races or sub-species are local forms completely fixed and isolated; but as they do not differ from each other by strongly marked and important characters, "There is no possible test but individual opinion to determine which of them shall be considered as species and which as varieties." Lastly, representative species fill the same place in the natural economy of each island as do the local forms and sub-species; but as they are distinguished from each other by a greater amount of difference than that between the local forms and sub-species, they are almost universally ranked by naturalists as true species. Nevertheless, no certain criterion can possibly be given by which variable forms, local forms, sub species and representative species can be recognised.

Many years ago, when comparing, and seeing others compare, the birds from the closely neighbouring islands of the Galapagos Archipelago, one with another, and with those from the American mainland, I was much struck how entirely vague and arbitrary is the distinction between species and varieties. On the islets of the little Madeira group there are many insects which are characterized as varieties in Mr. Wollaston's admirable work, but which would certainly be ranked as distinct species by many entomologists. Even Ireland has a few animals, now generally regarded as varieties, but which have been ranked as species by some zoologists. Several experienced ornithologists consider our British red grouse as only a strongly marked race of a Norwegian species, whereas the greater number rank it as an undoubted species peculiar to Great Britain. A wide distance between the homes of two doubtful forms leads many naturalists to rank them as distinct species; but what distance, it has been well asked, will suffice if that between America and Europe is ample, will that between Europe and the Azores, or Madeira, or the Canaries, or between the several islets of these small archipelagos, be sufficient?

Mr. B.D. Walsh, a distinguished entomologist of the United States, has described what he calls Phytophagic varieties and Phytophagic species. Most vegetable-feeding insects live on one kind of plant or on one group of plants; some feed indiscriminately on many kinds, but do not in consequence vary. In several cases, however, insects found living on different plants, have been observed by Mr. Walsh to present in their larval or mature state, or in both states, slight, though constant differences in colour, size, or in the nature of their secretions. In some instances the males alone, in other instances, both males and females, have been observed thus to differ in a slight degree. When the differences are rather more strongly marked, and when both sexes and all ages are affected, the forms are ranked by all entomologists as good species. But no observer can determine for another, even if he can do so for himself, which of these Phytophagic forms ought to be called species and which varieties. Mr. Walsh ranks the forms which it may be supposed would freely intercross, as varieties; and those which appear to have lost this power, as species. As the differences depend on the insects having long fed on distinct plants, it cannot be expected that intermediate links connecting the several forms should now be found. The naturalist thus loses his best guide in determining whether to rank doubtful forms as varieties or species. This likewise necessarily occurs with closely allied organisms, which inhabit distinct continents or islands. When, on the other hand, an animal or plant ranges over the same continent, or inhabits many islands in the same archipelago, and presents different forms in the different areas, there is always a good chance that intermediate forms will be discovered which will link together the extreme states; and these are then degraded to the rank of varieties.

Some few naturalists maintain that animals never present varieties; but then these same naturalists rank the slightest difference as of specific value; and when the same identical form is met with in two distant countries, or in two geological formations, they believe that two distinct species are hidden under the same dress. The term species thus comes to be a mere useless abstraction, implying and assuming a separate act of creation. It is certain that many forms, considered by highly competent judges to be varieties, resemble species so completely in character that they have been thus ranked by other highly competent judges. But to discuss whether they ought to be called species or varieties, before any definition of these terms has been generally accepted, is vainly to beat the air.

Many of the cases of strongly marked varieties or doubtful species well deserve consideration; for several interesting lines of argument, from geographical distribution, analogical variation, hybridism, etc., have been brought to bear in the attempt to determine their rank; but space does not here permit me to discuss them. Close investigation, in many cases, will no doubt bring naturalists to agree how to rank doubtful forms. Yet it must be confessed that it is in the best known countries that we find the greatest number of them. I have been struck with the fact that if any animal or plant in a state of nature be highly useful to man, or from any cause closely attracts his attention, varieties of it will almost universally be found recorded. These varieties, moreover, will often be ranked by some authors as species. Look at the common oak, how closely it has been studied; yet a German author makes more than a dozen species out of forms, which are almost universally considered by other botanists to be varieties; and in this country the highest botanical authorities and practical men can be quoted to show that the sessile and pedunculated oaks are either good and distinct species or mere varieties.

I may here allude to a remarkable memoir lately published by A. de Candolle, on the oaks of the whole world. No one ever had more ample materials for the discrimination of the species, or could have worked on them with more zeal and sagacity. He first gives in detail all the many points of structure which vary in the several species, and estimates numerically the relative frequency of the variations. He specifies above a dozen characters which may be found varying even on the same branch, sometimes according to age or development, sometimes without any assignable reason. Such characters are not of course of specific value, but they are, as Asa Gray has remarked in commenting on this memoir, such as generally enter into specific definitions. De Candolle then goes on to say that he gives the rank of species to the forms that differ by characters never varying on the same tree, and never found connected by intermediate states. After this discussion, the result of so much labour, he emphatically remarks: "They are mistaken, who repeat that the greater part of our species are clearly limited, and that the doubtful species are in a feeble minority. This seemed to be true, so long as a genus was imperfectly known, and its species were founded upon a few specimens, that is to say, were provisional. Just as we come to know them better, intermediate forms flow in, and doubts as to specific limits augment." He also adds that it is the best known species which present the greatest number of spontaneous varieties and sub-varieties. Thus Quercus robur has twenty-eight varieties, all of which, excepting six, are clustered round three sub-species, namely Q. pedunculata, sessiliflora and pubescens. The forms which connect these three sub-species are comparatively rare; and, as Asa Gray again remarks, if these connecting forms which are now rare were to become totally extinct the three sub-species would hold exactly the same relation to each other as do the four or five provisionally admitted species which closely surround the typical Quercus robur. Finally, De Candolle admits that out of the 300 species, which will be enumerated in his Prodromus as belonging to the oak family, at least two-thirds are provisional species, that is, are not known strictly to fulfil the definition above given of a true species. It should be added that De Candolle no longer believes that species are immutable creations, but concludes that the derivative theory is the most natural one, "and the most accordant with the known facts in palaeontology, geographical botany and zoology, of anatomical structure and classification."

When a young naturalist commences the study of a group of organisms quite unknown to him he is at first much perplexed in determining what differences to consider as specific and what as varietal; 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 able to make up his own mind; 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 he comes to study allied forms brought from countries not now continuous, in which case he cannot hope to find intermediate links, he will be compelled 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 by an insensible series; and a series impresses the mind with the idea of an actual passage.

Hence I look at individual differences, though of small interest to the systematist, as of the highest importance for us, as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. And I look at varieties which are in any degree more distinct and permanent, as steps toward more strongly marked and permanent varieties; and at the latter, as leading to sub-species, and then to species. The passage from one stage of difference to another may, in many cases, be the simple result of the nature of the organism and of the different physical conditions to which it has long been exposed; but with respect to the more important and adaptive characters, the passage from one stage of difference to another may be safely attributed to the cumulative action of natural selection, hereafter to be explained, and to the effects of the increased use or disuse of parts. A well-marked variety may therefore be called an incipient species; but whether this belief is justifiable must be judged by the weight of the various facts and considerations to be given throughout this work.

It need not be supposed that all varieties or incipient species attain the rank of species. They may become extinct, or they may endure as varieties for very long periods, as has been shown to be the case by Mr. Wollaston with the varieties of certain fossil land-shells in Madeira, and with plants by Gaston de Saporta. If a variety were to flourish so as to exceed in numbers the parent species, it would then rank as the species, and the species as the variety; or it might come to supplant and exterminate the parent species; or both might co-exist, and both rank as independent species. But we shall hereafter return to this subject.

From these remarks it will be seen that I look at the term species as one arbitrarily given, for the sake of convenience, to a set of individuals closely resembling each other, and that it does not essentially differ from the term variety, which is given to less distinct and more fluctuating forms. The term variety, again, in comparison with mere individual differences, is also applied arbitrarily, for convenience sake.

WIDE-RANGING, MUCH DIFFUSED, AND COMMON SPECIES VARY MOST.

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 a future work the discussion of these difficulties, and the tables 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.

Alphonse de Candolle and others have shown that plants which have very wide ranges generally present varieties; and this might have been expected, as they are 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 the 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), oftenest 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, 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, still inherit those advantages that enabled their parents to become dominant over their compatriots. In these remarks on predominence, it should be understood that reference is made only to the forms which come into competition with each other, and more especially to the members of the same genus or class having nearly similar habits of life. With respect to the number of individuals or commonness of species, the comparison of course relates only to the members of the same group. One of the higher plants may be said to be dominant if it be more numerous in individuals and more widely diffused than the other plants of the same country, which live under nearly the same conditions. A plant of this kind is not the less dominant because some conferva inhabiting the water or some parasitic fungus is infinitely more numerous in individuals, and more widely diffused. But if the conferva or parasitic fungus exceeds its allies in the above respects, it will then be dominant within its own class.

SPECIES OF THE LARGER GENERA IN EACH COUNTRY VARY MORE FREQUENTLY THAN THE SPECIES OF THE SMALLER GENERA.

If the plants inhabiting a country as described in any Flora, be divided into two equal masses, all those in the larger genera (i.e., those including many species) being placed on one side, and all those in the smaller genera on the other side, the former will be found to include a somewhat larger number of the very common and much diffused or dominant species. This 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 larger 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 generally have 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 still be 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.

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 presented 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 least genera, with from only one to four species, are altogether excluded from the tables. These facts are of plain signification on the view that species are only strongly marked and permanent varieties; for wherever 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 holds true 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 certainly holds good.

MANY OF THE SPECIES INCLUDED WITHIN THE LARGER GENERA RESEMBLE VARIETIES IN BEING VERY CLOSELY, BUT UNEQUALLY, RELATED TO EACH OTHER, AND IN HAVING RESTRICTED RANGES.

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 when 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 confirm the view. I have also consulted some sagacious and 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 the usual amount of difference.

Moreover, the species of the larger 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 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 tend to increase into the greater differences between species.

There is one other point which is 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 would be reversed. But there is reason to believe that the 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) sixty-three 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 sixty-three 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, fifty-three 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 the 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.

SUMMARY.

Finally, varieties cannot be distinguished from species—except, first, by the discovery of intermediate linking forms; and, secondly, by a certain indefinite amount of difference between them; for two forms, if differing very little, are generally ranked as varieties, notwithstanding that they cannot be closely connected; but the amount of difference considered necessary to give to any two forms the rank of species cannot be defined. In genera having more than the average number of species in any country, the species of these genera have more than the average number of varieties. In large genera the species are apt to be closely but unequally allied together, forming little clusters round other species. Species very closely allied to other species apparently have restricted ranges. In all these respects the species of large genera present a strong analogy with varieties. And we can clearly understand these analogies, if species once existed as varieties, and thus originated; whereas, these analogies are utterly inexplicable if species are independent creations.

We have also seen that it is the most flourishing or dominant species of the larger genera within each class which on an average yield the greatest number of varieties, and varieties, as we shall hereafter see, tend to become converted into new and distinct species. Thus the larger genera tend to become larger; and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. But, by steps hereafter to be explained, the larger genera also tend to break up into smaller genera. And thus, the forms of life throughout the universe become divided into groups subordinate to groups.



CHAPTER III. STRUGGLE FOR EXISTENCE.

Its bearing on natural selection—The term used in a wide sense—Geometrical ratio of increase—Rapid increase of naturalised animals and plants—Nature of the checks to increase—Competition universal—Effects of climate—Protection from the number of individuals—Complex relations of all animals and plants throughout nature—Struggle for life most severe between individuals and varieties of the same species: often severe between species of the same genus—The relation of organism to organism the most important of all relations.

Before entering on the subject of this chapter I must make a few preliminary remarks to show how the struggle for existence bears on natural selection. It has been seen in the last chapter that among organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life and of one organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.

Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow from the struggle for life. Owing to this struggle, variations, however slight and from whatever cause proceeding, if they be in any degree profitable to the individuals of a species, in their infinitely complex relations to other organic beings and to their physical conditions of life, will tend to the preservation of such individuals, and will generally be inherited by the offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term natural selection, in order to mark its relation to man's power of selection. But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man's feeble efforts, as the works of Nature are to those of Art.

We will now discuss in a little more detail the struggle for existence. In my future work this subject will be treated, as it well deserves, at greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult—at least I found it so—than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that, though food may be now superabundant, it is not so at all seasons of each recurring year.

THE TERM, STRUGGLE FOR EXISTENCE, USED IN A LARGE SENSE.

I should premise that I use this term in a large and metaphorical sense, including dependence of one being on another, and including (which is more important) not only the life of the individual, but success in leaving progeny. Two canine animals, in a time of dearth, may be truly said to struggle with each other which shall get food and live. But a plant on the edge of a desert is said to struggle for life against the drought, though more properly it should be said to be dependent on the moisture. A plant which annually produces a thousand seeds, of which only one of an average comes to maturity, may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. The mistletoe is dependent on the apple and a few other trees, but can only in a far-fetched sense be said to struggle with these trees, for, if too many of these parasites grow on the same tree, it languishes and dies. But several seedling mistletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the mistletoe is disseminated by birds, its existence depends on them; and it may metaphorically be said to struggle with other fruit-bearing plants, in tempting the birds to devour and thus disseminate its seeds. In these several senses, which pass into each other, I use for convenience sake the general term of Struggle for Existence.

GEOMETRICAL RATIO OF INCREASE.

A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season or occasional year, otherwise, on the principle of geometrical increase, its numbers would quickly become so inordinately great that no country could support the product. Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence, either one individual with another of the same species, or with the individuals of distinct species, or with the physical conditions of life. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms; for in this case there can be no artificial increase of food, and no prudential restraint from marriage. Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them.

There is no exception to the rule that every organic being naturally increases at so high a rate, that, if not destroyed, the earth would soon be covered by the progeny of a single pair. Even slow-breeding man has doubled in twenty-five years, and at this rate, in less than a thousand years, there would literally not be standing room for his progeny. Linnaeus has calculated that if an annual plant produced only two seeds—and there is no plant so unproductive as this—and their seedlings next year produced two, and so on, then in twenty years there would be a million plants. The elephant is reckoned the slowest breeder of all known animals, and I have taken some pains to estimate its probable minimum rate of natural increase; it will be safest to assume that it begins breeding when thirty years old, and goes on breeding till ninety years old, bringing forth six young in the interval, and surviving till one hundred years old; if this be so, after a period of from 740 to 750 years there would be nearly nineteen million elephants alive descended from the first pair.

But we have better evidence on this subject than mere theoretical calculations, namely, the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature, when circumstances have been favourable to them during two or three following seasons. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world; if the statements of the rate of increase of slow-breeding cattle and horses in South America, and latterly in Australia, had not been well authenticated, they would have been incredible. So it is with plants; cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. Several of the plants, such as the cardoon and a tall thistle, which are now the commonest over the wide plains of La Plata, clothing square leagues of surface almost to the exclusion of every other plant, have been introduced from Europe; and there are plants which now range in India, as I hear from Dr. Falconer, from Cape Comorin to the Himalaya, which have been imported from America since its discovery. In such cases, and endless others could be given, no one supposes that the fertility of the animals or plants has been suddenly and temporarily increased in any sensible degree. The obvious explanation is that the conditions of life have been highly favourable, and that there has consequently been less destruction of the old and young and that nearly all the young have been enabled to breed. Their geometrical ratio of increase, the result of which never fails to be surprising, simply explains their extraordinarily rapid increase and wide diffusion in their new homes.

In a state of nature almost every full-grown plant annually produces seed, and among animals there are very few which do not annually pair. Hence we may confidently assert that all plants and animals are tending to increase at a geometrical ratio—that all would rapidly stock every station in which they could any how exist, and that this geometrical tendency to increase must be checked by destruction at some period of life. Our familiarity with the larger domestic animals tends, I think, to mislead us; we see no great destruction falling on them, and we do not keep in mind that thousands are annually slaughtered for food, and that in a state of nature an equal number would have somehow to be disposed of.

The only difference between organisms which annually produce eggs or seeds by the thousand, and those which produce extremely few, is, that the slow breeders would require a few more years to people, under favourable conditions, a whole district, let it be ever so large. The condor lays a couple of eggs and the ostrich a score, and yet in the same country the condor may be the more numerous of the two. The Fulmar petrel lays but one egg, yet it is believed to be the most numerous bird in the world. One fly deposits hundreds of eggs, and another, like the hippobosca, a single one. But this difference does not determine how many individuals of the two species can be supported in a district. A large number of eggs is of some importance to those species which depend on a fluctuating amount of food, for it allows them rapidly to increase in number. But the real importance of a large number of eggs or seeds is to make up for much destruction at some period of life; and this period in the great majority of cases is an early one. If an animal can in any way protect its own eggs or young, a small number may be produced, and yet the average stock be fully kept up; but if many eggs or young are destroyed, many must be produced or the species will become extinct. It would suffice to keep up the full number of a tree, which lived on an average for a thousand years, if a single seed were produced once in a thousand years, supposing that this seed were never destroyed and could be ensured to germinate in a fitting place; so that, in all cases, the average number of any animal or plant depends only indirectly on the number of its eggs or seeds.

In looking at Nature, it is most necessary to keep the foregoing considerations always in mind—never to forget that every single organic being may be said to be striving to the utmost to increase in numbers; that each lives by a struggle at some period of its life; that heavy destruction inevitably falls either on the young or old during each generation or at recurrent intervals. Lighten any check, mitigate the destruction ever so little, and the number of the species will almost instantaneously increase to any amount.

NATURE OF THE CHECKS TO INCREASE.

The causes which check the natural tendency of each species to increase are most obscure. Look at the most vigorous species; by as much as it swarms in numbers, by so much will it tend to increase still further. We know not exactly what the checks are even in a single instance. Nor will this surprise any one who reflects how ignorant we are on this head, even in regard to mankind, although so incomparably better known than any other animal. This subject of the checks to increase has been ably treated by several authors, and I hope in a future work to discuss it at considerable length, more especially in regard to the feral animals of South America. Here I will make only a few remarks, just to recall to the reader's mind some of the chief points. Eggs or very young animals seem generally to suffer most, but this is not invariably the case. With plants there is a vast destruction of seeds, but from some observations which I have made it appears that the seedlings suffer most from germinating in ground already thickly stocked with other plants. Seedlings, also, are destroyed in vast numbers by various enemies; for instance, on a piece of ground three feet long and two wide, dug and cleared, and where there could be no choking from other plants, I marked all the seedlings of our native weeds as they came up, and out of 357 no less than 295 were destroyed, chiefly by slugs and insects. If turf which has long been mown, and the case would be the same with turf closely browsed by quadrupeds, be let to grow, the more vigorous plants gradually kill the less vigorous, though fully grown plants; thus out of twenty species grown on a little plot of mown turf (three feet by four) nine species perished, from the other species being allowed to grow up freely.

The amount of food for each species, of course, gives the extreme limit to which each can increase; but very frequently it is not the obtaining food, but the serving as prey to other animals, which determines the average number of a species. Thus, there seems to be little doubt that the stock of partridges, grouse, and hares on any large estate depends chiefly on the destruction of vermin. If not one head of game were shot during the next twenty years in England, and, at the same time, if no vermin were destroyed, there would, in all probability, be less game than at present, although hundreds of thousands of game animals are now annually shot. On the other hand, in some cases, as with the elephant, none are destroyed by beasts of prey; for even the tiger in India most rarely dares to attack a young elephant protected by its dam.

Climate plays an important part in determining the average numbers of a species, and periodical seasons of extreme cold or drought seem to be the most effective of all checks. I estimated (chiefly from the greatly reduced numbers of nests in the spring) that the winter of 1854-5 destroyed four-fifths of the birds in my own grounds; and this is a tremendous destruction, when we remember that ten per cent. is an extraordinarily severe mortality from epidemics with man. The action of climate seems at first sight to be quite independent of the struggle for existence; but in so far as climate chiefly acts in reducing food, it brings on the most severe struggle between the individuals, whether of the same or of distinct species, which subsist on the same kind of food. Even when climate, for instance, extreme cold, acts directly, it will be the least vigorous individuals, or those which have got least food through the advancing winter, which will suffer the most. When we travel from south to north, or from a damp region to a dry, we invariably see some species gradually getting rarer and rarer, and finally disappearing; and the change of climate being conspicuous, we are tempted to attribute the whole effect to its direct action. But this is a false view; we forget that each species, even where it most abounds, is constantly suffering enormous destruction at some period of its life, from enemies or from competitors for the same place and food; and if these enemies or competitors be in the least degree favoured by any slight change of climate, they will increase in numbers; and as each area is already fully stocked with inhabitants, the other species must decrease. When we travel southward and see a species decreasing in numbers, we may feel sure that the cause lies quite as much in other species being favoured, as in this one being hurt. So it is when we travel northward, but in a somewhat lesser degree, for the number of species of all kinds, and therefore of competitors, decreases northward; hence in going northward, or in ascending a mountain, we far oftener meet with stunted forms, due to the DIRECTLY injurious action of climate, than we do in proceeding southward or in descending a mountain. When we reach the Arctic regions, or snow-capped summits, or absolute deserts, the struggle for life is almost exclusively with the elements.

That climate acts in main part indirectly by favouring other species we clearly see in the prodigious number of plants which in our gardens can perfectly well endure our climate, but which never become naturalised, for they cannot compete with our native plants nor resist destruction by our native animals.

When a species, owing to highly favourable circumstances, increases inordinately in numbers in a small tract, epidemics—at least, this seems generally to occur with our game animals—often ensue; and here we have a limiting check independent of the struggle for life. But even some of these so-called epidemics appear to be due to parasitic worms, which have from some cause, possibly in part through facility of diffusion among the crowded animals, been disproportionally favoured: and here comes in a sort of struggle between the parasite and its prey.

On the other hand, in many cases, a large stock of individuals of the same species, relatively to the numbers of its enemies, is absolutely necessary for its preservation. Thus we can easily raise plenty of corn and rape-seed, etc., in our fields, because the seeds are in great excess compared with the number of birds which feed on them; nor can the birds, though having a superabundance of food at this one season, increase in number proportionally to the supply of seed, as their numbers are checked during the 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 exist; and that of some social plants being social, that is abounding in individuals, even on the extreme verge 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 the species from utter destruction. I should add that the good effects of intercrossing, and the ill effects of close interbreeding, no doubt come into play in many of these cases; but I will not here enlarge on this subject.

COMPLEX RELATIONS OF ALL ANIMALS AND PLANTS TO EACH OTHER IN THE STRUGGLE FOR EXISTENCE.

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, 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 of the land having been enclosed, so that cattle could not enter. But how important an element enclosure is, I plainly saw near Farnham, in Surrey. Here there are extensive heaths, with a few clumps of old Scotch firs on the distant hill-tops: within the last ten years large spaces have been enclosed, and self-sown firs are now springing up in multitudes, so close together that all cannot live. When I ascertained that these young trees had not been sown or planted I was so much surprised at their numbers that I went to several points of view, whence I could examine hundreds of acres of the unenclosed heath, and literally I could not see a single Scotch fir, except the old planted clumps. But on looking closely between the stems of the heath, I found a multitude of seedlings and little trees, which had been perpetually browsed down by the cattle. In one square yard, at a point some hundred yards distant from one of the old clumps, I counted thirty-two little trees; and one of them, with twenty-six rings of growth, had, during many 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 other parasitic insects. Hence, if certain insectivorous birds were to decrease in Paraguay, the parasitic insects would probably increase; and this would lessen the number of the navel-frequenting flies—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 have just seen in Staffordshire, the insectivorous birds, and so onwards in ever-increasing circles of complexity. Not that under nature the relations will ever be as simple as this. Battle within battle must be continually recurring with varying success; and yet in the long-run the forces are so nicely balanced that the face of nature remains for long periods of time uniform, though assuredly the merest trifle would 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, 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 is never visited in my garden by insects, and consequently, from its peculiar structure, never sets a seed. Nearly all our orchidaceous plants absolutely require the visits of insects to remove their pollen-masses and thus to fertilise them. I find from experiments that humble-bees are almost indispensable to the fertilisation of the heartsease (Viola tricolor), for other bees do not visit this flower. I have also found that the visits of bees are necessary for the fertilisation of some kinds of clover; for instance twenty heads of Dutch clover (Trifolium repens) yielded 2,290 seeds, but twenty other heads, protected from bees, produced not one. Again, 100 heads of red clover (T. pratense) produced 2,700 seeds, but the same number of protected heads produced not a single seed. Humble bees alone visit red clover, as other bees cannot reach the nectar. It has been suggested that moths may fertilise the clovers; but I doubt whether they could do so in the case of the red clover, from their weight not being sufficient to depress the wing petals. Hence we may infer as highly probable 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 measure upon the number of field-mice, which destroy their combs and nests; and Colonel 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 Colonel 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 will concur 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 ancient Indian ruins in the Southern United States, which must formerly have been cleared of trees, now display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. What a struggle must have gone on during long centuries between the several kinds of trees, 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, all feeding on each other, or on the trees, 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 fall to the ground according to definite laws; but how simple is the problem where each shall fall compared to that of 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 likewise sometimes 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 will almost invariably 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 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 number 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 of our domestic plants or animals have so exactly the same strength, habits, and constitution, that the original proportions of a mixed stock (crossing being prevented) could be kept up for half-a-dozen generations, if they were allowed to struggle together, in the same manner as beings in a state of nature, and if the seed or young were not annually preserved in due proportion.

STRUGGLE FOR LIFE MOST SEVERE BETWEEN INDIVIDUALS AND VARIETIES OF THE SAME SPECIES.

As the species of the same genus usually have, though by no means invariably, much similarity in habits and constitution, and always in structure, the struggle will generally be more severe between them, if they come into competition with each other, than between the 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. In Australia the imported hive-bee is rapidly exterminating the small, stingless native bee. One species of charlock has been known to supplant another species; 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 the plumed seeds no doubt stands in the closest relation to the land being already thickly clothed with 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, it may be suspected that the chief use of the nutriment in the seed is to favour the growth of the seedlings, whilst struggling with other plants growing vigorously all around.

Look at a plant in the midst of its range! Why does it not double or quadruple its numbers? We know that it can perfectly well withstand a little more heat or cold, dampness or dryness, for elsewhere it ranges into slightly hotter or colder, damper or drier districts. In this case we can clearly see that if we wish in imagination to give the plant the power of increasing in numbers, we should have to give it some advantage over its competitors, or over the animals which prey on it. On the confines of its geographical range, a change of constitution with respect to climate would clearly be an advantage to our plant; but we have reason to believe that only a few plants or animals range so far, that they are destroyed exclusively by the rigour of the climate. Not until we reach the extreme confines of life, in the Arctic regions or on the borders of an utter desert, will competition cease. The land may be extremely cold or dry, yet there will be competition between some few species, or between the individuals of the same species, for the warmest or dampest spots.

Hence we can see that when a plant or animal is placed in a new country, among new competitors, the conditions of its life will generally be changed in an essential manner, although the climate may be exactly the same as in its former home. If its average numbers are to increase in its new home, we should have to modify it in a different way to what we should have had to do in its native country; for we should have to give it some advantage over a different set of competitors or enemies.

It is good thus to try in imagination to give any one species an advantage over another. Probably in no single instance should we know what to do. This ought to convince us of our ignorance on the mutual relations of all organic beings; a conviction as necessary, as it is difficult to acquire. All that we can do is to keep steadily in mind that each organic being is striving to increase in 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.



CHAPTER IV. NATURAL SELECTION; OR THE SURVIVAL OF THE FITTEST.

Natural Selection—its power compared with man's selection—its power on characters of trifling importance—its power at all ages and on both sexes—Sexual Selection—On the generality of intercrosses between individuals of the same species—Circumstances favourable and unfavourable to the results of Natural Selection, namely, intercrossing, isolation, number of individuals—Slow action—Extinction caused by Natural Selection—Divergence of Character, related to the diversity of inhabitants of any small area and to naturalisation—Action of Natural Selection, through Divergence of Character and Extinction, on the descendants from a common parent—Explains the Grouping of all organic beings—Advance in organisation—Low forms preserved—Convergence of character—Indefinite multiplication of species—Summary.

How will the struggle for existence, briefly discussed in the last chapter, act in regard to variation? Can the principle of selection, which we have seen is so potent in the hands of man, apply under nature? I think we shall see that it can act most efficiently. Let the endless number of slight variations and individual differences occurring in our domestic productions, and, in a lesser degree, in those under nature, be borne in mind; as well as the strength of the hereditary tendency. Under domestication, it may truly be said that the whole organisation becomes in some degree plastic. But the variability, which we almost universally meet with in our domestic productions is not directly produced, as Hooker and Asa Gray have well remarked, by man; he can neither originate varieties nor prevent their occurrence; he can only preserve and accumulate such as do occur. Unintentionally he exposes organic beings to new and changing conditions of life, and variability ensues; but similar changes of conditions might and do occur under nature. Let it also be borne in mind how infinitely complex and close-fitting are the mutual relations of all organic beings to each other and to their physical conditions of life; and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life. Can it then be thought improbable, seeing that variations useful to man have undoubtedly occurred, that other variations useful in some way to each being in the great and complex battle of life, should occur in the course of many successive generations? If such do occur, can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage, however slight, over others, would have the best chance of surviving and procreating their kind? On the other hand, we may feel sure that any variation in the least degree injurious would be rigidly destroyed. This preservation of favourable individual differences and variations, and the destruction of those which are injurious, I have called Natural Selection, or the Survival of the Fittest. Variations neither useful nor injurious would not be affected by natural selection, and would be left either a fluctuating element, as perhaps we see in certain polymorphic species, or would ultimately become fixed, owing to the nature of the organism and the nature of the conditions.