THE LAWS OF VARIABILITY WITH RESPECT TO ANIMALS AND PLANTS

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THE LAWS OF VARIABILITY WITH RESPECT TO ANIMALS AND PLANTS.

I shall in this volume treat, as fully as my materials permit, the whole subject of variation under domestication. We may thus hope to obtain some light, little though it be, on the causes of variability, on the laws which govern it—such as the direct action of climate and food, the effects of use and disuse, and of correlation of growth—and on the amount of change to which domesticated organisms are liable.

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Although man does not cause variability and can not even prevent it, he can select, preserve, and accumulate the variations given to him by the hand of Nature almost in any way which he chooses; and thus he can certainly produce a great result. Selection may be followed either methodically and intentionally, or unconsciously and unintentionally. Man may select and preserve each successive variation, with the distinct intention of improving and altering a breed, in accordance with a preconceived idea; and by thus adding up variations, often so slight as to be imperceptible by an uneducated eye, he has effected wonderful changes and improvements. It can, also, be clearly shown that man, without any intention or thought of improving the breed, by preserving in each successive generation the individuals which he prizes most, and by destroying the worthless individuals, slowly, though surely, induces great changes. As the will of man thus comes into play, we can understand how it is that domesticated breeds show adaptation to his wants and pleasures. We can further understand how it is that domestic races of animals and cultivated races of plants often exhibit an abnormal character, as compared with natural species; for they have been modified not for their own benefit, but for that of man.

INHERITED EFFECT OF CHANGED HABITS.

When we compare the individuals of the same variety or subvariety of our older cultivated plants and animals, one of the first points which strikes us is, that they generally differ more from each other than do the individuals of any one species or variety in a state of nature. And if we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, we are driven to conclude that this great variability is due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent species had been exposed under nature.

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Changed habits produce an inherited effect, as in the period of the flowering of plants when transported from one climate to another. With animals the increased use or disuse of parts has had a more marked influence; thus I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more, in proportion to the whole skeleton, than do the same bones in the wild-duck; and this change may be safely attributed to the domestic duck flying much less, and walking more, than its wild parents. The great and inherited development of the udders in cows and goats in countries where they are habitually milked, in comparison with these organs in other countries, is probably another instance of the effects of use. Not one of our domestic animals can be named which has not in some country drooping ears; and the view which has been suggested that the drooping is due to the disease of the muscles of the ear, from the animals being seldom much alarmed, seems probable.

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From facts collected by Heusinger, it appears that white sheep and pigs are injured by certain plants, while dark-colored individuals escape, Professor Wyman has recently communicated to me a good illustration of this fact: on asking some farmers in Virginia how it was that all their pigs were black, they informed him that the pigs ate the paint-root (Lachnanthes), which colored their bones pink, and which caused the hoofs of all but the black varieties to drop off; and one of the “crackers” (i. e., Virginia squatters) added, “We select the black members of a litter for raising, as they alone have a good chance of living.” Hairless dogs have imperfect teeth; long-haired and coarse-haired animals are apt to have, as is asserted, long or many horns; pigeons with feathered feet have skin between their outer toes; pigeons with short beaks have small feet, and those with long beaks large feet. Hence, if man goes on selecting, and thus augmenting, any peculiarity, he will almost certainly modify unintentionally other parts of the structure, owing to the mysterious laws of correlation.

EFFECTS OF THE USE AND DISUSE OF PARTS.

From the facts alluded to in the first chapter, I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts, and disuse diminished them, and that such modifications are inherited. Under free nature we have no standard of comparison by which to judge of the effects of long-continued use or disuse, for we know not the parent forms; but many animals possess structures which can be best explained by the effects of disuse. As Professor Owen has remarked, there is no greater anomaly in nature than a bird that can not fly; yet there are several in this state. The logger-headed duck of South America can only flap along the surface of the water, and has its wings in nearly the same condition as the domestic Aylesbury duck: it is a remarkable fact that the young birds, according to Mr. Cunningham, can fly, while the adults have lost this power. As the larger ground-feeding birds seldom take flight, except to escape danger, it is probable that the nearly wingless condition of several birds, now inhabiting or which lately inhabited several oceanic islands, tenanted by no beast of prey, has been caused by disuse. The ostrich, indeed, inhabits continents, and is exposed to danger from which it can not escape by flight, but it can defend itself by kicking its enemies as efficiently as many quadrupeds. We may believe that the progenitor of the ostrich genus had habits like those of the bustard, and that, as the size and weight of its body were increased during successive generations, its legs were used more, and its wings less, until they became incapable of flight.

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The insects in Madeira which are not ground-feeders, and which, as certain flower-feeding Coleoptera and Lepidoptera, must habitually use their wings to gain their subsistence, have, as Mr. Wollaston suspects, their wings not at all reduced, but even enlarged. This is quite compatible with the action of natural selection. For, when a new insect first arrived on the island, the tendency of natural selection to enlarge or to reduce the wings would depend on whether a greater number of individuals were saved by successfully battling with the winds, or by giving up the attempt and rarely or never flying. As with mariners shipwrecked near a coast, it would have been better for the good swimmers if they had been able to swim still farther, whereas it would have been better for the bad swimmers if they had not been able to swim at all and had stuck to the wreck.

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

VAGUE ORIGIN OF OUR DOMESTIC ANIMALS.

In the case of most of our anciently domesticated animals and plants, it is not possible to come to any definite conclusion whether they are descended from one or several wild species. The argument mainly relied on by those who believe in the multiple origin of our domestic animals is, that we find in the most ancient times, on the monuments of Egypt, and in the lake-habitations of Switzerland, much diversity in the breeds; and that some of these ancient breeds closely resemble or are even identical with, those still existing. But this only throws far backward the history of civilization, and shows that animals were domesticated at a much earlier period than has hitherto been supposed. The lake-inhabitants of Switzerland cultivated several kinds of wheat and barley, the pea, the poppy for oil, and flax; and they possessed several domesticated animals. They also carried on commerce with other nations. All this clearly shows, as Heer has remarked, that they had at this early age progressed considerably in civilization; and this again implies a long-continued previous period of less advanced civilization, during which the domesticated animals, kept by different tribes in different districts, might have varied and given rise to distinct races. Since the discovery of flint tools in the superficial formations of many parts of the world, all geologists believe that barbarian man existed at an enormously remote period; and we know that at the present day there is hardly a tribe so barbarous as not to have domesticated at least the dog.

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The origin of most of our domestic animals will probably forever remain vague.

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In attempting to estimate the amount of structural difference between allied domestic races, we are soon involved in doubt, from not knowing whether they are descended from one or several parent species. This point, if it could be cleared up, would be interesting; if, for instance, it could be shown that the greyhound, bloodhound, terrier, spaniel, and bull-dog, which we all know propagate their kind truly, were the offspring of any single species. Then such facts would have great weight in making us doubt about the immutability of the many closely allied natural species—for instance, of the many foxes—inhabiting different quarters of the world.

DESCENT OF THE DOMESTIC PIGEON.

Great as are the differences between the breeds of the pigeon, I am fully convinced that the common opinion of naturalists is correct, namely, that all are descended from the rock-pigeon (Columba livia), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects. As several of the reasons which have led me to this belief are in some degree applicable in other cases, I will here briefly give them. If the several breeds are not varieties, and have not proceeded from the rock-pigeon, they must have descended from at least seven or eight aboriginal stocks; for it is impossible to make the present domestic breeds by the crossing of any lesser number: how, for instance, could a pouter be produced by crossing two breeds unless one of the parent-stocks possessed the characteristic enormous crop? The supposed aboriginal stocks must all have been rock-pigeons—that is, they did not breed or willingly perch on trees. But besides C. livia, with its geographical sub-species, only two or three other species of rock-pigeons are known, and these have not any of the characters of the domestic breeds. Hence the supposed aboriginal stocks must either still exist in the countries where they were originally domesticated, and yet be unknown to ornithologists—and this, considering their size, habits, and remarkable characters, seems improbable—or they must have become extinct in the wild state. But birds breeding on precipices, and good fliers, are unlikely to be exterminated; and the common rock-pigeon, which has the same habits with the domestic breeds, has not been exterminated even on several of the smaller British islets, or on the shores of the Mediterranean. Hence the supposed extermination of so many species having similar habits with the rock-pigeon seems a very rash assumption. Moreover, the several above-named domesticated breeds have been transported to all parts of the world, and therefore some of them must have been carried back again into their native country; but not one has become wild or feral, though the dovecot-pigeon, which is the rock-pigeon in a very slightly altered state, has become feral in several places. Again, all recent experience shows that it is difficult to get wild animals to breed freely under domestication; yet, on the hypothesis of the multiple origin of our pigeons, it must be assumed that at least seven or eight species were so thoroughly domesticated in ancient times by half-civilized man as to be quite prolific under confinement.

An argument of great weight, and applicable in several other cases, is, that the above-specified breeds, though agreeing generally with the wild rock-pigeon in constitution, habits, voice, coloring, and in most parts of their structure, yet are certainly highly abnormal in other parts; we may look in vain through the whole great family of Columbidæ for a beak like that of the English carrier, or that of the short-faced tumbler, or barb; for reversed feathers like those of the Jacobin; for a crop like that of the pouter; for tail-feathers like those of the fantail. Hence it must be assumed not only that half-civilized man succeeded in thoroughly domesticating several species, but that he intentionally or by chance picked out extraordinarily abnormal species; and, further, that these very species have since all become extinct or unknown. So many strange contingencies are improbable in the highest degree.

ORIGIN OF THE DOG.

The first and chief point of interest in this chapter is, whether the numerous domesticated varieties of the dog have descended from a single wild species, or from several. Some authors believe that all have descended from the wolf, or from the jackal, or from an unknown and extinct species. Others again believe, and this of late has been the favorite tenet, that they have descended from several species, extinct and recent, more or less commingled together. We shall probably never be able to ascertain their origin with certainty. Paleontology does not throw much light on the question, owing, on the one hand, to the close similarity of the skulls of extinct as well as living wolves and jackals, and owing, on the other hand, to the great dissimilarity of the skulls of the several breeds of the domestic dogs. It seems, however, that remains have been found in the later tertiary deposits more like those of a large dog than of a wolf, which favors the belief of De Blainville that our dogs are the descendants of a single extinct species. On the other hand, some authors go so far as to assert that every chief domestic breed must have had its wild prototype. This latter view is extremely improbable: it allows nothing for variation; it passes over the almost monstrous character of some of the breeds; and it almost necessarily assumes that a large number of species have become extinct since man domesticated the dog; whereas we plainly see that wild members of the dog-family are extirpated by human agency with much difficulty; even so recently as 1710 the wolf existed in so small an island as Ireland.

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At a period between four and five thousand years ago, various breeds—viz., pariah dogs, greyhounds, common hounds, mastiffs, house-dogs, lap-dogs, and turnspits—existed, more or less closely resembling our present breeds. But there is not sufficient evidence that any of these ancient dogs belonged to the same identical sub-varieties with our present dogs. As long as man was believed to have existed on this earth only about six thousand years, this fact of the great diversity of the breeds at so early a period was an argument of much weight that they had proceeded from several wild sources, for there would not have been sufficient time for their divergence and modification. But now that we know, from the discovery of flint tools imbedded with the remains of extinct animals, in districts which have since undergone great geographical changes, that man has existed for an incomparably longer period, and bearing in mind that the most barbarous nations possess domestic dogs, the argument from insufficient time falls away greatly in value.

From this resemblance of the half-domesticated dogs in several countries to the wild species still living there—from the facility with which they can often be crossed together—from even half-tamed animals being so much valued by savages—and from the other circumstances previously remarked on which favor their domestication, it is highly probable that the domestic dogs of the world are descended from two well-defined species of wolf (viz., C. lupus and C. latrans), and from two or three other doubtful species (namely, the European, Indian, and North African wolves); from at least one or two South American canine species; from several races or species of jackal; and perhaps from one or more extinct species.

ORIGIN OF THE HORSE.

The history of the horse is lost in antiquity. Remains of this animal in a domesticated condition have been found in the Swiss lake-dwellings, belonging to the Neolithic period. At the present time the number of breeds is great, as may be seen by consulting any treatise on the horse. Looking only to the native ponies of Great Britain, those of the Shetland Isles, Wales, the New Forest, and Devonshire are distinguishable; and so it is, among other instances, with each separate island in the great Malay Archipelago. Some of the breeds present great differences in size, shape of ears, length of mane, proportions of the body, form of the withers and hind-quarters, and especially in the head. Compare the race-horse, dray-horse, and a Shetland pony in size, configuration, and disposition; and see how much greater the difference is than between the seven or eight other living species of the genus Equus.

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Horses have often been observed, according to M. Gaudry, to possess a trapezium and a rudiment of a fifth metacarpal bone, so that “one sees appearing by monstrosity, in the foot of the horse, structures which normally exist in the foot of the hipparion”—an allied and extinct animal. In various countries horn-like projections have been observed on the frontal bones of the horse: in one case described by Mr. Percival they arose about two inches above the orbital processes, and were “very like those in a calf from five to six months old,” being from half to three quarters of an inch in length.

CAUSES OF MODIFICATIONS IN THE HORSE.

With respect to the causes of the modifications which horses have undergone, the conditions of life seem to produce a considerable direct effect. Mr. D. Forbes, who has had excellent opportunities of comparing the horses of Spain with those of South America, informs me that the horses of Chili, which have lived under nearly the same conditions as their progenitors in Andalusia, remain unaltered, while the Pampas horses and the Puno ponies are considerably modified. There can be no doubt that horses become greatly reduced in size and altered in appearance by living on mountains and islands; and this apparently is due to want of nutritious or varied food. Every one knows how small and rugged the ponies are on the northern islands and on the mountains of Europe. Corsica and Sardinia have their native ponies; and there were, or still are, on some islands on the coast of Virginia, ponies like those of the Shetland Islands, which are believed to have originated through exposure to unfavorable conditions. The Puno ponies, which inhabit the lofty regions of the Cordillera, are, as I hear from Mr. D. Forbes, strange little creatures, very unlike their Spanish progenitors. Farther south, in the Falkland Islands, the offspring of the horses imported in 1764 have already so much deteriorated in size and strength, that they are unfitted for catching wild cattle with the lasso; so that fresh horses have to be brought for this purpose from La Plata at a great expense. The reduced size of the horses bred on both southern and northern islands, and on several mountain-chains, can hardly have been caused by the cold, as a similar reduction has occurred on the Virginian and Mediterranean islands.

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It is scarcely possible to doubt that the long-continued selection of qualities serviceable to man has been the chief agent in the formation of the several breeds of the horse. Look at a dray-horse, and see how well adapted he is to draw heavy weights, and how unlike in appearance to any allied wild animal. The English race-horse is known to be derived from the commingled blood of Arabs, Turks, and Barbs; but selection, which was carried on during very early times in England, together with training, have made him a very different animal from his parent stocks.

“MAKING THE WORKS OF GOD A MERE MOCKERY.”

We see several distinct species of the horse-genus becoming, by simple variation, striped on the legs like a zebra, or striped on the shoulders like an ass. In the horse we see this tendency strong whenever a dun tint appears—a tint that approaches to that of the general coloring of the other species of the genus. The appearance of the stripes is not accompanied by any change of form or by any other new character. We see this tendency to become striped most strongly displayed in hybrids from between several of the most distinct species. Now observe the case of the several breeds of pigeons: they are descended from a pigeon (including two or three sub-species or geographical races) of a bluish color, with certain bars and other marks; and, when any breed assumes by simple variation a bluish tint, these bars and other marks invariably reappear; but without any other change of form or character. When the oldest and truest breeds of various colors are crossed, we see a strong tendency for the blue tint and bars and marks to reappear in the mongrels. I have stated that the most probable hypothesis to account for the reappearance of very ancient characters is—that there is a tendency in the young of each successive generation to produce the long-lost character, and that this tendency, from unknown causes, sometimes prevails. And we have just seen that in several species of the horse-genus the stripes are either plainer or appear more commonly in the young than in the old. Call the breeds of pigeons, some of which have bred true for centuries, species; and how exactly parallel is the case with that of the species of the horse-genus! For myself, I venture confidently to look back thousands on thousands of generations, and I see an animal striped like a zebra, but perhaps otherwise very differently constructed, the common parent of our domestic horse (whether or not it be descended from one or more wild stocks), of the ass, the hemionus, quagga, and zebra.

He who believes that each equine species was independently created, will, I presume, assert that each species has been created with a tendency to vary, both under nature and under domestication, in this particular manner, so as often to become striped like the other species of the genus; and that each has been created with a strong tendency, when crossed with species inhabiting distant quarters of the world, to produce hybrids resembling in their stripes, not their own parents, but other species of the genus. To admit this view is, as it seems to me, to reject a real for an unreal, or at least for an unknown, cause. It makes the works of God a mere mockery and deception; I would almost as soon believe with the old and ignorant cosmogonists, that fossil shells had never lived, but had been created in stone so as to mock the shells living on the sea-shore.

VARIABILITY OF CULTIVATED PLANTS.

I shall not enter into so much detail on the variability of cultivated plants as in the case of domesticated animals. The subject is involved in much difficulty. Botanists have generally neglected cultivated varieties, as beneath their notice. In several cases the wild prototype is unknown or doubtfully known; and in other cases it is hardly possible to distinguish between escaped seedlings and truly wild plants, so that there is no safe standard of comparison by which to judge of any supposed amount of change. Not a few botanists believe that several of our anciently cultivated plants have become so profoundly modified that it is not possible now to recognize their aboriginal parent-forms. Equally perplexing are the doubts whether some of them are descended from one species, or from several inextricably commingled by crossing and variation. Variations often pass into, and can not be distinguished from, monstrosities; and monstrosities are of little significance for our purpose. Many varieties are propagated solely by grafts, buds, layers, bulbs, etc., and frequently it is not known how far their peculiarities can be transmitted by seminal generation.

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From innumerable experiments made through dire necessity by the savages of every land, with the results handed down by tradition, the nutritious, stimulating, and medicinal properties of the most unpromising plants were probably first discovered. It appears, for instance, at first an inexplicable fact that untutored man, in three distant quarters of the world, should have discovered, among a host of native plants, that the leaves of the tea-plant and mattee, and the berries of the coffee, all included a stimulating and nutritious essence, now known to be chemically the same. We can also see that savages suffering from severe constipation would naturally observe whether any of the roots which they devoured acted as aperients. We probably owe our knowledge of the uses of almost all plants to man having originally existed in a barbarous state, and having been often compelled by severe want to try as food almost everything which he could chew and swallow.

SAVAGE WISDOM IN THE CULTIVATION OF PLANTS.

The savage inhabitants of each land, having found out by many and hard trials what plants were useful, or could be rendered useful by various cooking processes, would after a time take the first step in cultivation by planting them near their usual abodes. Livingstone states that the savage Batokas sometimes left wild fruit-trees standing in their gardens, and occasionally even planted them, “a practice seen nowhere else among the natives.” But Du Chaillu saw a palm and some other wild fruit-trees which had been planted; and these trees were considered private property. The next step in cultivation, and this would require but little forethought, would be to sow the seeds of useful plants; and, as the soil near the hovels of the natives would often be in some degree manured, improved varieties would sooner or later arise. Or a wild and unusually good variety of a native plant might attract the attention of some wise old savage; and he would transplant it, or sow its seed. That superior varieties of wild fruit-trees occasionally are found is certain, as in the case of the American species of hawthorns, plums, cherries, grapes, and hickories, specified by Professor Asa Gray.

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We now know that man was sufficiently civilized to cultivate the ground at an immensely remote period; so that wheat might have been improved long ago up to that standard of excellence which was possible under the then existing state of agriculture. One small class of facts supports this view of the slow and gradual improvement of our cereals. In the most ancient lake-habitations of Switzerland, when men employed only flint-tools, the most extensively cultivated wheat was a peculiar kind, with remarkably small ears and grains. “While the grains of the modern forms are in section from seven to eight millimetres in length, the larger grains from the lake-habitations are six, seldom seven, and the smaller ones only four. The ear is thus much narrower, and the spikelets stand out more horizontally, than in our present forms.” So again with barley, the most ancient and most extensively cultivated kind had small ears, and the grains were “smaller, shorter, and nearer to each other, than in that now grown; without the husk they were two and one half lines long, and scarcely one and one half broad, while those now grown have a length of three lines, and almost the same in breadth.” These small-grained varieties of wheat and barley are believed by Heer to be the parent-forms of certain existing allied varieties, which have supplanted their early progenitors.

UNKNOWN LAWS OF INHERITANCE.

The laws governing inheritance are for the most part unknown. No one can say why the same peculiarity in different individuals of the same species, or in different species, is sometimes inherited and sometimes not so; why the child often reverts in certain characters to its grandfather or grandmother or more remote ancestor; why a peculiarity is often transmitted from one sex to both sexes, or to one sex alone, more commonly but not exclusively to the like sex. It is a fact of some importance to us that peculiarities appearing in the males of our domestic breeds are often transmitted either exclusively, or in a much greater degree, to the males alone. A much more important rule, which I think may be trusted, is that, at whatever period of life a peculiarity first appears, it tends to reappear in the offspring at a corresponding age, though sometimes earlier. In many cases this could not be otherwise: thus the inherited peculiarities in the horns of cattle could appear only in the offspring when nearly mature; peculiarities in the silk-worm are known to appear at the corresponding caterpillar or cocoon stage. But hereditary diseases and some other facts make me believe that the rule has a wider extension, and that, when there is no apparent reason why a peculiarity should appear at any particular age, yet that it does tend to appear in the offspring at the same period at which it first appeared in the parent. I believe this rule to be of the highest importance in explaining the laws of embryology. These remarks are, of course, confined to the first appearance of the peculiarity, and not to the primary cause which may have acted on the ovules or on the male element; in nearly the same manner as the increased length of the horns in the offspring from a short-horned cow by a long-horned bull, though appearing late in life, is clearly due to the male element.

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If animals and plants had never been domesticated, and wild ones alone had been observed, we should probably never have heard the saying that “like begets like.” The proposition would have been as self-evident as that all the buds on the same tree are alike, though neither proposition is strictly true. For, as has often been remarked, probably no two individuals are identically the same. All wild animals recognize each other, which shows that there is some difference between them; and, when the eye is well practiced, the shepherd knows each sheep, and man can distinguish a fellow-man out of millions on millions of other men.

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The subject of inheritance is wonderful. When a new character arises, whatever its nature may be, it generally tends to be inherited, at least in a temporary and sometimes in a most persistent manner. What can be more wonderful than that some trifling peculiarity, not primordially attached to the species, should be transmitted through the male or female sexual cells, which are so minute as not to be visible to the naked eye, and afterward through the incessant changes of a long course of development, undergone either in the womb or in the egg, and ultimately appear in the offspring when mature, or even when quite old, as in the case of certain diseases? Or, again, what can be more wonderful than the well-ascertained fact that the minute ovule of a good milking-cow will produce a male, from whom a cell, in union with an ovule, will produce a female, and she, when mature, will have large mammary glands, yielding an abundant supply of milk, and even milk of a particular quality? Nevertheless, the real subject of surprise is, as Sir H. Holland has well remarked, not that a character should be inherited, but that any should ever fail to be inherited.

LAWS OF INHERITANCE THAT ARE FAIRLY WELL ESTABLISHED.

Though much remains obscure with respect to inheritance, we may look at the following laws as fairly well established: Firstly, a tendency in every character, new and old, to be transmitted by seminal and bud generation, though often counteracted by various known and unknown causes. Secondly, reversion or atavism, which depends on transmission and development being distinct powers: it acts in various degrees and manners through both seminal and bud generation. Thirdly, prepotency of transmission, which may be confined to one sex, or be common to both sexes. Fourthly, transmission, as limited by sex, generally to the same sex in which the inherited character first appeared; and this in many, probably most cases, depends on the new character having first appeared at a rather late period of life. Fifthly, inheritance at corresponding periods of life, with some tendency to the earlier development of the inherited character. In these laws of inheritance, as displayed under domestication, we see an ample provision for the production, through variability and natural selection, of new specific forms.

INHERITED PECULIARITIES IN MAN.

Gait, gestures, voice, and general bearing, are all inherited, as the illustrious Hunter and Sir A. Carlisle have insisted. My father communicated to me some striking instances, in one of which a man died during the early infancy of his son, and my father, who did not see this son until grown up and out of health, declared that it seemed to him as if his old friend had risen from the grave, with all his highly peculiar habits and manners. Peculiar manners pass into tricks, and several instances could be given of their inheritance; as in the case, often quoted, of the father who generally slept on his back, with his right leg crossed over the left, and whose daughter, while an infant in the cradle, followed exactly the same habit, though an attempt was made to cure her. I will give one instance which has fallen under my own observation, and which is curious from being a trick associated with a peculiar state of mind, namely, pleasurable emotion. A boy had the singular habit, when pleased, of rapidly moving his fingers parallel to each other, and, when much excited, of raising both hands, with the fingers still moving, to the sides of his face on a level with the eyes: when this boy was almost an old man, he could still hardly resist this trick when much pleased, but from its absurdity concealed it. He had eight children. Of these, a girl, when pleased, at the age of four and a half years, moved her fingers in exactly the same way, and, what is still odder, when much excited, she raised both her hands, with her fingers still moving, to the sides of her face, in exactly the same manner as her father had done, and sometimes even still continued to do so when alone. I never heard of any one, excepting this one man and his little daughter, who had this strange habit; and certainly imitation was in this instance out of the question.

INHERITED DISEASES.

Large classes of diseases usually appear at certain ages, such as St. Vitus’s dance in youth, consumption in early mid-life, gout later, and apoplexy still later; and these are naturally inherited at the same period. But, even in diseases of this class, instances have been recorded, as with St. Vitus’s dance, showing that an unusually early or late tendency to the disease is inheritable. In most cases the appearance of any inherited disease is largely determined by certain critical periods in each person’s life, as well as by unfavorable conditions. There are many other diseases, which are not attached to any particular period, but which certainly tend to appear in the child at about the same age at which the parent was first attacked. An array of high authorities, ancient and modern, could be given in support of this proposition. The illustrious Hunter believed in it; and Piorry cautions the physician to look closely to the child at the period when any grave inheritable disease attacked the parent. Dr. Prosper Lucas, after collecting facts from every source, asserts that affections of all kinds, though not related to any particular period of life, tend to reappear in the offspring at whatever period of life they first appeared in the progenitor.

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Esquirol gives several striking instances of insanity coming on at the same age as that of a grandfather, father, and son, who all committed suicide near their fiftieth year. Many other cases could be given, as of a whole family who became insane at the age of forty. Other cerebral affections sometimes follow the same rule—for instance, epilepsy and apoplexy. A woman died of the latter disease when sixty-three years old; one of her daughters at forty-three, and the other at sixty-seven: the latter had twelve children, who all died from tubercular meningitis. I mention this latter case because it illustrates a frequent occurrence, namely, a change in the precise nature of an inherited disease, though still affecting the same organ.

* * * * *

Two brothers, their father, their paternal uncles, seven cousins, and their paternal grandfather, were all similarly affected by a skin-disease, called pityriasis versicolor; “the disease, strictly limited to the males of the family (though transmitted through the females), usually appeared at puberty, and disappeared at about the age of forty or forty-five years.” The second case is that of four brothers, who, when about twelve years old, suffered almost every week from severe headaches, which were relieved only by a recumbent position in a dark room. Their father, paternal uncles, paternal grandfather, and grand-uncles all suffered in the same way from headaches, which ceased at the age of fifty-four or fifty-five in all those who lived so long. None of the females of the family were affected.

CAUSES OF NON-INHERITANCE.

A large number of cases of non-inheritance are intelligible on the principle that a strong tendency to inheritance does exist, but that it is overborne by hostile or unfavorable conditions of life. No one would expect that our improved pigs, if forced during several generations to travel about and root in the ground for their own subsistence, would transmit, as truly as they now do, their short muzzles and legs, and their tendency to fatten. Dray-horses assuredly would not long transmit their great size and massive limbs, if compelled to live in a cold, damp, mountainous region; we have, indeed, evidence of such deterioration in the horses which have run wild on the Falkland Islands. European dogs in India often fail to transmit their true character. Our sheep in tropical countries lose their wool in a few generations. There seems also to be a close relation between certain peculiar pastures and the inheritance of an enlarged tail in fat-tailed sheep, which form one of the most ancient breeds in the world. With plants, we have seen that tropical varieties of maize lose their proper character in the course of two or three generations, when cultivated in Europe; and conversely so it is with European varieties cultivated in Brazil. Our cabbages, which here come so true by seed, can not form heads in hot countries. According to Carrière, the purple-leafed beech and barberry transmit their character by seed far less truly in certain districts than in others. Under changed circumstances, periodical habits of life soon fail to be transmitted, as the period of maturity in summer and winter wheat, barley, and vetches. So it is with animals: for instance, a person, whose statement I can trust, procured eggs of Aylesbury ducks from that town, where they are kept in houses, and are reared as early as possible for the London market; the ducks bred from these eggs in a distant part of England, hatched their first brood on January 24th, while common ducks, kept in the same yard and treated in the same manner, did not hatch till the end of March; and this shows that the period of hatching was inherited. But the grandchildren of these Aylesbury ducks completely lost their habit of early incubation, and hatched their eggs at the same time with the common ducks of the same place.

Many cases of non-inheritance apparently result from the conditions of life continually inducing fresh variability. We have seen that when the seeds of pears, plums, apples, etc., are sown, the seedlings generally inherit some degree of family likeness. Mingled with these seedlings, a few, and sometimes many, worthless, wild-looking plants commonly appear, and their appearance may be attributed to the principle of reversion. But scarcely a single seedling will be found perfectly to resemble the parent-form; and this may be accounted for by constantly recurring variability induced by the conditions of life.

STEPS BY WHICH DOMESTIC RACES HAVE BEEN PRODUCED.

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 blood-hound, 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 rivaled 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-cock, 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 accumulative 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.

* * * * *

If selection consisted merely in separating some very distinct variety, and breeding from it, the principle would be so obvious as hardly to be worth notice; but its importance consists in the great effect produced by the accumulation in one direction, during successive generations, of differences absolutely inappreciable by an uneducated eye—differences which I for one have vainly attempted to appreciate. Not one man in a thousand has accuracy of eye and 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 assuredly fail. Few would readily believe in the natural capacity and years of practice requisite to become even a skillful pigeon-fancier.

UNCONSCIOUS SELECTION.

A man who intends keeping pointers naturally tries to get as good dogs as he can, and afterward 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 can never be recognized 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 civilized districts, where the breed has been less improved. There is reason to believe that King Charles’s 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 fox-hound; 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 favored 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.

* * * * *

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.

ADAPTATION OF ANIMALS TO THE FANCIES OF MAN.

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 œsophagus—a habit which is disregarded by all fanciers, as it is not one of the points of the breed.

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 can not 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 one hundred and eighty-two 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 two hundred and fifty-one species, whereas Mr. Bentham gives only one hundred and twelve—a difference of one hundred and thirty-nine doubtful forms!

SPECIES AN ARBITRARY TERM.

Certainly no clear line of demarkation 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 steps toward 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 coexist, 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.

THE TRUE PLAN OF CREATION.

When the views advanced by me in this volume, and by Mr. Wallace, or when analogous views on the origin of species are generally admitted, we can dimly foresee that there will be a considerable revolution in natural history. Systematists will be able to pursue their labors as at present; but they will not be incessantly haunted by the shadowy doubt whether this or that form be a true species.

* * * * *

Hereafter we shall be compelled to acknowledge that the only distinction between species and well-marked varieties is, that the latter are known, or believed, to be connected at the present day by intermediate gradations, whereas species were formerly thus connected. Hence, without rejecting the consideration of the present existence of intermediate gradations between any two forms, we shall be led to weigh more carefully and to value higher the actual amount of difference between them. It is quite possible that forms now generally acknowledged to be merely varieties may hereafter be thought worthy of specific names; and in this case scientific and common language will come into accordance. In short, we shall have to treat species in the same manner as those naturalists treat genera who admit that genera are merely artificial combinations made for convenience. This may not be a cheering prospect; but we shall at least be freed from the vain search for the undiscovered and undiscoverable essence of the term species.

The other and more general departments of natural history will rise greatly in interest. The terms used by naturalists, of affinity, relationship, community of type, paternity, morphology, adaptive characters, rudimentary80 and aborted organs, etc., will cease to be metaphorical, and will have a plain signification. When we no longer look at an organic being as a savage looks at a ship, as something wholly beyond his comprehension; when we regard every production of nature as one which has had a long history; when we contemplate every complex structure and instinct as the summing up of many contrivances, each useful to the possessor, in the same way as any great mechanical invention is the summing up of the labor, the experience, the reason, and even the blunders of numerous workmen; when we thus view each organic being, how far more interesting—I speak from experience—does the study of natural history become!

A grand and almost untrodden field of inquiry will be opened, on the causes and laws of variation, on correlation, on the effects of use and disuse, on the direct action of external conditions, and so forth. The study of domestic productions will rise immensely in value. A new variety raised by man will be a more important and interesting subject for study than one more species added to the infinitude of already recorded species. Our classifications will come to be, as far as they can be so made, genealogies, and will then truly give what may be called the plan of creation.

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