The Origin of Species by Means of Natural Selection Part 3

PRINCIPLES OF SELECTION ANCIENTLY FOLLOWED, AND THEIR EFFECTS.

Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some effect may be attributed to the direct and definite action of the external conditions of life, and some to habit; but he would be a bold man who would account by such agencies for the differences between a dray and race-horse, a greyhound and bloodhound, a carrier and tumbler pigeon.

One of the most remarkable features in our domesticated races is that we see in them adaptation, not indeed to the animal's or plant's own good, but to man's use or fancy. Some variations useful to him have probably arisen suddenly, or by one step; many botanists, for instance, believe that the fuller's teasel, with its hooks, which can not be rivalled by any mechanical contrivance, is only a variety of the wild Dipsacus; and this amount of change may have suddenly arisen in a seedling. So it has probably been with the turnspit dog; and this is known to have been the case with the ancon sheep. But when we compare the dray-horse and race-horse, the dromedary and camel, the various breeds of sheep fitted either for cultivated land or mountain pasture, with the wool of one breed good for one purpose, and that of another breed for another purpose; when we compare the many breeds of dogs, each good for man in different ways; when we compare the game-c.o.c.k, so pertinacious in battle, with other breeds so little quarrelsome, with "everlasting layers" which never desire to sit, and with the bantam so small and elegant; when we compare the host of agricultural, culinary, orchard, and flower-garden races of plants, most useful to man at different seasons and for different purposes, or so beautiful in his eyes, we must, I think, look further than to mere variability. We can not suppose that all the breeds were suddenly produced as perfect and as useful as we now see them; indeed, in many cases, we know that this has not been their history. The key is man's power of acc.u.mulative selection: nature gives successive variations; man adds them up in certain directions useful to him. In this sense he may be said to have made for himself useful breeds.

The great power of this principle of selection is not hypothetical.

It is certain that several of our eminent breeders have, even within a single lifetime, modified to a large extent their breeds of cattle and sheep. In order fully to realise what they have done it is almost necessary to read several of the many treatises devoted to this subject, and to inspect the animals. Breeders habitually speak of an animal's organisation as something plastic, which they can model almost as they please. If I had s.p.a.ce I could quote numerous pa.s.sages to this effect from highly competent authorities. Youatt, who was probably better acquainted with the works of agriculturalists than almost any other individual, and who was himself a very good judge of animals, speaks of the principle of selection as "that which enables the agriculturist, not only to modify the character of his flock, but to change it altogether.

It is the magician's wand, by means of which he may summon into life whatever form and mould he pleases." Lord Somerville, speaking of what breeders have done for sheep, says: "It would seem as if they had chalked out upon a wall a form perfect in itself, and then had given it existence." In Saxony the importance of the principle of selection in regard to merino sheep is so fully recognised, that men follow it as a trade: the sheep are placed on a table and are studied, like a picture by a connoisseur; this is done three times at intervals of months, and the sheep are each time marked and cla.s.sed, so that the very best may ultimately be selected for breeding.

What English breeders have actually effected is proved by the enormous prices given for animals with a good pedigree; and these have been exported to almost every quarter of the world. The improvement is by no means generally due to crossing different breeds; all the best breeders are strongly opposed to this practice, except sometimes among closely allied sub-breeds. And when a cross has been made, the closest selection is far more indispensable even than in ordinary cases. If selection consisted merely in separating some very distinct variety and breeding from it, the principle would be so obvious as hardly to be worth notice; but its importance consists in the great effect produced by the acc.u.mulation in one direction, during successive generations, of differences absolutely inappreciable by an uneducated eye--differences which I for one have vainly attempted to appreciate. Not one man in a thousand has accuracy of eye and judgment sufficient to become an eminent breeder. If gifted with these qualities, and he studies his subject for years, and devotes his lifetime to it with indomitable perseverance, he will succeed, and may make great improvements; if he wants any of these qualities, he will a.s.suredly fail. Few would readily believe in the natural capacity and years of practice requisite to become even a skilful pigeon-fancier.

The same principles are followed by horticulturists; but the variations are here often more abrupt. No one supposes that our choicest productions have been produced by a single variation from the aboriginal stock. We have proofs that this is not so in several cases in which exact records have been kept; thus, to give a very trifling instance, the steadily increasing size of the common gooseberry may be quoted.

We see an astonis.h.i.+ng improvement in many florists' flowers, when the flowers of the present day are compared with drawings made only twenty or thirty years ago. When a race of plants is once pretty well established, the seed-raisers do not pick out the best plants, but merely go over their seed-beds, and pull up the "rogues," as they call the plants that deviate from the proper standard. With animals this kind of selection is, in fact, likewise followed; for hardly any one is so careless as to breed from his worst animals.

In regard to plants, there is another means of observing the acc.u.mulated effects of selection--namely, by comparing the diversity of flowers in the different varieties of the same species in the flower-garden; the diversity of leaves, pods, or tubers, or whatever part is valued, in the kitchen-garden, in comparison with the flowers of the same varieties; and the diversity of fruit of the same species in the orchard, in comparison with the leaves and flowers of the same set of varieties. See how different the leaves of the cabbage are, and how extremely alike the flowers; how unlike the flowers of the heartsease are, and how alike the leaves; how much the fruit of the different kinds of gooseberries differ in size, colour, shape, and hairiness, and yet the flowers present very slight differences. It is not that the varieties which differ largely in some one point do not differ at all in other points; this is hardly ever--I speak after careful observation--perhaps never, the case. The law of correlated variation, the importance of which should never be overlooked, will ensure some differences; but, as a general rule, it cannot be doubted that the continued selection of slight variations, either in the leaves, the flowers, or the fruit, will produce races differing from each other chiefly in these characters.

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 pa.s.sed 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 cla.s.sical writers. From pa.s.sages 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 pa.s.sages 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 a.s.sociated 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 surpa.s.s 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 pa.s.sed, and come to differ so greatly from the rock-pigeon.

Youatt gives an excellent ill.u.s.tration 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 cla.s.sical 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 cla.s.sical 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 acc.u.mulated, 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 circ.u.mstanced, individuals of the same species, having slightly different const.i.tutions 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 fas.h.i.+on--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.

CIRc.u.mSTANCES FAVOURABLE TO MAN'S POWER OF SELECTION.

I will now say a few words on the circ.u.mstances, 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 acc.u.mulation 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 Yorks.h.i.+re, "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 circ.u.mstance 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, peac.o.c.k, 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 peac.o.c.ks, 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 a.s.sert 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 a.s.sert 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 a.n.a.logous 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 circ.u.mstances. 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 acc.u.mulative 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 sh.e.l.ls 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 circ.u.mstances. 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 acc.u.mulate, in the same manner as man acc.u.mulates 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 cla.s.sificatory 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 a.s.suredly 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 sh.e.l.ls. 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 sh.e.l.ls, 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 s.e.xes 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 b.u.t.terflies, in the Malayan Archipelago, regularly appear under two or even three conspicuously distinct forms, not connected by intermediate varieties. Fritz Muller has described a.n.a.logous 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 b.u.t.terfly 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 b.u.t.terfly 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 s.e.xes 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 a.s.sumed 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 a.n.a.logy 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 a.s.sistance 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 cla.s.sed 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 s.e.xes 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 a.s.suming 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, a.n.a.logical variation, hybridism, etc., have been brought to bear in the attempt to determine their rank; but s.p.a.ce 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 a.s.signable 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 cl.u.s.tered round three sub-species, namely Q. pedunculata, sessiliflora and p.u.b.escens. 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 cla.s.sification."