Fragment

INTRODUCTION.
When on board H.M.S. Beagle, as naturalist, I was much struck with certain facts in the distribution of the organic beings inhabiting South America, and in the geological relations of the present to the past inhabitants of that continent. These facts, as will be seen in the latter chapters of this volume, seemed to throw some light on the origin of species—that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, it occurred to me, in 1837, that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it. After five years’ work I allowed myself to speculate on the subject, and drew up some short notes; these I enlarged in 1844 into a sketch of the conclusions, which then seemed to me probable: from that period to the present day I have steadily pursued the same object. I hope that I may be excused for entering on these personal details, as I give them to show that I have not been hasty in coming to a decision.
My work is now (1859) nearly finished; but as it will take me many more years to complete it, and as my health is far from strong, I have been urged to publish this abstract. I have more especially been induced to do this, as Mr. Wallace, who is now studying the natural history of the Malay Archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. In 1858 he sent me a memoir on this subject, with a request that I would forward it to Sir Charles Lyell, who sent it to the Linnean Society, and it is published in the third volume of the Journal of that Society. Sir C. Lyell and Dr. Hooker, who both knew of my work—the latter having read my sketch of 1844—honoured me by thinking it advisable to publish, with Mr. Wallace’s excellent memoir, some brief extracts from my manuscripts.
This abstract, which I now publish, must necessarily be imperfect. I cannot here give references and authorities for my several statements; and I must trust to the reader reposing some confidence in my accuracy. No doubt errors may have crept in, though I hope I have always been cautious in trusting to good authorities alone. I can here give only the general conclusions at which I have arrived, with a few facts in illustration, but which, I hope, in most cases will suffice. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts, with references, on which my conclusions have been grounded; and I hope in a future work to do this. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced, often apparently leading to conclusions directly opposite to those at which I have arrived. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question; and this is here impossible.
I much regret that want of space prevents my having the satisfaction of acknowledging the generous assistance which I have received from very many naturalists, some of them personally unknown to me. I cannot, however, let this opportunity pass without expressing my deep obligations to Dr. Hooker, who, for the last fifteen years, has aided me in every possible way by his large stores of knowledge and his excellent judgment.
In considering the origin of species, it is quite conceivable that a naturalist, reflecting on the mutual affinities of organic beings, on their embryological relations, their geographical distribution, geological succession, and other such facts, might come to the conclusion that species had not been independently created, but had descended, like varieties, from other species. Nevertheless, such a conclusion, even if well founded, would be unsatisfactory, until it could be shown how the innumerable species, inhabiting this world have been modified, so as to acquire that perfection of structure and coadaptation which justly excites our admiration. Naturalists continually refer to external conditions, such as climate, food, &c., as the only possible cause of variation. In one limited sense, as we shall hereafter see, this may be true; but it is preposterous to attribute to mere external conditions, the structure, for instance, of the woodpecker, with its feet, tail, beak, and tongue, so admirably adapted to catch insects under the bark of trees. In the case of the mistletoe, which draws its nourishment from certain trees, which has seeds that must be transported by certain birds, and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its relations to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the plant itself.
It is, therefore, of the highest importance to gain a clear insight into the means of modification and coadaptation. At the commencement of my observations it seemed to me probable that a careful study of domesticated animals and of cultivated plants would offer the best chance of making out this obscure problem. Nor have I been disappointed; in this and in all other perplexing cases I have invariably found that our knowledge, imperfect though it be, of variation under domestication, afforded the best and safest clue. I may venture to express my conviction of the high value of such studies, although they have been very commonly neglected by naturalists.
From these considerations, I shall devote the first chapter of this abstract to variation under domestication. We shall thus see that a large amount of hereditary modification is at least possible; and, what is equally or more important, we shall see how great is the power of man in accumulating by his selection successive slight variations. I will then pass on to the variability of species in a state of nature; but I shall, unfortunately, be compelled to treat this subject far too briefly, as it can be treated properly only by giving long catalogues of facts. We shall, however, be enabled to discuss what circumstances are most favourable to variation. In the next chapter the struggle for existence among all organic beings throughout the world, which inevitably follows from the high geometrical ratio of their increase, will be considered. This is the doctrine of Malthus, applied to the whole animal and vegetable kingdoms. As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.
This fundamental subject of natural selection will be treated at some length in the fourth chapter; and we shall then see how natural selection almost inevitably causes much extinction of the less improved forms of life, and leads to what I have called divergence of character. In the next chapter I shall discuss the complex and little known laws of variation. In the five succeeding chapters, the most apparent and gravest difficulties in accepting the theory will be given: namely, first, the difficulties of transitions, or how a simple being or a simple organ can be changed and perfected into a highly developed being or into an elaborately constructed organ; secondly the subject of instinct, or the mental powers of animals; thirdly, hybridism, or the infertility of species and the fertility of varieties when intercrossed; and fourthly, the imperfection of the geological record. In the next chapter I shall consider the geological succession of organic beings throughout time; in the twelfth and thirteenth, their geographical distribution throughout space; in the fourteenth, their classification or mutual affinities, both when mature and in an embryonic condition. In the last chapter I shall give a brief recapitulation of the whole work, and a few concluding remarks.
No one ought to feel surprise at much remaining as yet unexplained in regard to the origin of species and varieties, if he make due allowance for our profound ignorance in regard to the mutual relations of the many beings which live around us. Who can explain why one species ranges widely and is very numerous, and why another allied species has a narrow range and is rare? Yet these relations are of the highest importance, for they determine the present welfare and, as I believe, the future success and modification of every inhabitant of this world. Still less do we know of the mutual relations of the innumerable inhabitants of the world during the many past geological epochs in its history. Although much remains obscure, and will long remain obscure, I can entertain no doubt, after the most deliberate study and dispassionate judgment of which I am capable, that the view which most naturalists until recently entertained, and which I formerly entertained—namely, that each species has been independently created—is erroneous. I am fully convinced that species are not immutable; but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species, in the same manner as the acknowledged varieties of any one species are the descendants of that species. Furthermore, I am convinced that natural selection has been the most important, but not the exclusive, means of modification.
 
CHAPTER I.
VARIATION UNDER DOMESTICATION.
Causes of Variability—Effects of Habit and the use and disuse of Parts—Correlated Variation—Inheritance—Character of Domestic Varieties—Difficulty of distinguishing between Varieties and Species—Origin of Domestic Varieties from one or more Species—Domestic Pigeons, their Differences and Origin—Principles of Selection, anciently followed, their Effects—Methodical and Unconscious Selection—Unknown Origin of our Domestic Productions—Circumstances favourable to Man’s power of Selection.
Causes of Variability.
When we compare the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us is, that they generally differ 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. There is, also, some probability in the view propounded by Andrew Knight, that this variability may be partly connected with excess of food. It seems clear that organic beings must be exposed during several generations to new conditions to cause any great amount of variation; and that, when the organisation has once begun to vary, it generally continues varying for many generations. No case is on record of a variable organism ceasing to vary under cultivation. Our oldest cultivated plants, such as wheat, still yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification.
As far as I am able to judge, after long attending to the subject, the conditions of life appear to act in two ways—directly on the whole organisation or on certain parts alone and in directly by affecting the reproductive system. With respect to the direct action, we must bear in mind that in every case, as Professor Weismann has lately insisted, and as I have incidently shown in my work on “Variation under Domestication,” there are two factors: namely, the nature of the organism and the nature of the conditions. The former seems to be much the more important; for nearly similar variations sometimes arise under, as far as we can judge, dissimilar conditions; and, on the other hand, dissimilar variations arise under conditions which appear to be nearly uniform. The effects on the offspring are either definite or in definite. They may be considered as definite when all or nearly all the offspring of individuals exposed to certain conditions during several generations are modified in the same manner. It is extremely difficult to come to any conclusion in regard to the extent of the changes which have been thus definitely induced. There can, however, be little doubt about many slight changes, such as size from the amount of food, colour from the nature of the food, thickness of the skin and hair from climate, &c. Each of the endless variations which we see in the plumage of our fowls must have had some efficient cause; and if the same cause were to act uniformly during a long series of generations on many individuals, all probably would be modified in the same manner. Such facts as the complex and extraordinary out growths which variably follow from the insertion of a minute drop of poison by a gall-producing insect, shows us what singular modifications might result in the case of plants from a chemical change in the nature of the sap.
In definite variability is a much more common result of changed conditions than definite variability, and has probably played a more important part in the formation of our domestic races. We see in definite variability in the endless slight peculiarities which distinguish the individuals of the same species, and which cannot be accounted for by inheritance from either parent or from some more remote ancestor. Even strongly-marked differences occasionally appear in the young of the same litter, and in seedlings from the same seed-capsule. At long intervals of time, out of millions of individuals reared in the same country and fed on nearly the same food, deviations of structure so strongly pronounced as to deserve to be called monstrosities arise; but monstrosities cannot be separated by any distinct line from slighter variations. All such changes of structure, whether extremely slight or strongly marked, which appear among many individuals living together, may be considered as the in definite effects of the conditions of life on each individual organism, in nearly the same manner as the chill effects different men in an in definite manner, according to their state of body or constitution, causing coughs or colds, rheumatism, or inflammation of various organs.
With respect to what I have called the in direct action of changed conditions, namely, through the reproductive system of being affected, we may infer that variability is thus induced, partly from the fact of this system being extremely sensitive to any change in the conditions, and partly from the similarity, as Kölreuter and others have remarked, between the variability which follows from the crossing of distinct species, and that which may be observed with plants and animals when reared under new or unnatural conditions. Many facts clearly show how eminently susceptible the reproductive system is to very slight changes in the surrounding conditions. Nothing is more easy than to tame an animal, and few things more difficult than to get it to breed freely under confinement, even when the male and female unite. How many animals there are which will not breed, though kept in an almost free state in their native country! This is generally, but erroneously attributed to vitiated instincts. Many cultivated plants display the utmost vigour, and yet rarely or never seed! In some few cases it has been discovered that a very trifling change, such as a little more or less water at some particular period of growth, will determine whether or not a plant will produce seeds. I cannot here give the details which I have collected and elsewhere published on this curious subject; but to show how singular the laws are which determine the reproduction of animals under confinement, I may mention that carnivorous animals, even from the tropics, breed in this country pretty freely under confinement, with the exception of the plantigrades or bear family, which seldom produce young; whereas, carnivorous birds, with the rarest exception, hardly ever lay fertile eggs. Many exotic plants have pollen utterly worthless, in the same condition as in the most sterile hybrids. When, on the one hand, we see domesticated animals and plants, though often weak and sickly, breeding freely under confinement; and when, on the other hand, we see individuals, though taken young from a state of nature perfectly tamed, long-lived, and healthy (of which I could give numerous instances), yet having their reproductive system so seriously affected by unperceived causes as to fail to act, we need not be surprised at this system, when it does act under confinement, acting irregularly, and producing offspring somewhat unlike their parents. I may add that as some organisms breed freely under the most unnatural conditions—for instance, rabbits and ferrets kept in hutches—showing that their reproductive organs are not easily affected; so will some animals and plants withstand domestication or cultivation, and vary very slightly—perhaps hardly more than in a state of nature.
Some naturalists have maintained that all variations are connected with the act of sexual reproduction; but this is certainly an error; for I have given in another work a long list of “sporting plants;” as they are called by gardeners; that is, of plants which have suddenly produced a single bud with a new and sometimes widely different character from that of the other buds on the same plant. These bud variations, as they may be named, can be propagated by grafts, offsets, &c., and sometimes by seed. They occur rarely under nature, but are far from rare under culture. As a single bud out of many thousands produced year after year on the same tree under uniform conditions, has been known suddenly to assume a new character; and as buds on distinct trees, growing under different conditions, have sometimes yielded nearly the same variety—for instance, buds on peach-trees producing nectarines, and buds on common roses producing moss-roses—we clearly see that the nature of the conditions is of subordinate importance in comparison with the nature of the organism in determining each particular form of variation; perhaps of not more importance than the nature of the spark, by which a mass of combustible matter is ignited, has in determining the nature of the flames.
Effects of Habit and of the Use or Disuse of Parts; Correlated Variation; Inheritance.
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 disuse of the muscles of the ear, from the animals being seldom much alarmed, seems probable.
Many laws regulate variation, some few of which can be dimly seen, and will hereafter be briefly discussed. I will here only allude to what may be called correlated variation. Important changes in the embryo or larva will probably entail changes in the mature animal. In monstrosities, the correlations between quite distinct parts are very curious; and many instances are given in Isidore Geoffroy St. Hilaire’s great work on this subject. Breeders believe that long limbs are almost always accompanied by an elongated head. Some instances of correlation are quite whimsical; thus cats which are entirely white and have blue eyes are generally deaf; but it has been lately stated by Mr. Tait that this is confined to the males. Colour and constitutional peculiarities go together, of which many remarkable cases could be given among animals and plants. From facts collected by Heusinger, it appears that white sheep and pigs are injured by certain plants, while dark-coloured 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 coloured 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.
The results of the various, unknown, or but dimly understood laws of variation are infinitely complex and diversified. It is well worth while carefully to study the several treatises on some of our old cultivated plants, as on the hyacinth, potato, even the dahlia, &c.; and it is really surprising to note the endless points of structure and constitution in which the varieties and sub-varieties differ slightly from each other. The whole organisation seems to have become plastic, and departs in a slight degree from that of the parental type.
Any variation which is not inherited is unimportant for us. But the number and diversity of inheritable deviations of structure, both those of slight and those of considerable physiological importance, are endless. Dr. Prosper Lucas’ treatise, in two large volumes, is the fullest and the best on this subject. No breeder doubts how strong is the tendency to inheritance; that like produces like is his fundamental belief: doubts have been thrown on this principle only by theoretical writers. When any deviation of structure often appears, and we see it in the father and child, we cannot tell whether it may not be due to the same cause having acted on both; but when among individuals, apparently exposed to the same conditions, any very rare deviation, due to some extraordinary combination of circumstances, appears in the parent—say, once among several million individuals—and it reappears in the child, the mere doctrine of chances almost compels us to attribute its reappearance to inheritance. Every one must have heard of cases of albinism, prickly skin, hairy bodies, &c., appearing in several members of the same family. If strange and rare deviations of structure are truly inherited, less strange and commoner deviations may be freely admitted to be inheritable. Perhaps the correct way of viewing the whole subject would be, to look at the inheritance of every character whatever as the rule, and non-inheritance as the anomaly.
The laws governing inheritance are 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 characteristics 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.
Having alluded to the subject of reversion, I may here refer to a statement often made by naturalists—namely, that our domestic varieties, when run wild, gradually but invariably revert in character to their aboriginal stocks. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature. I have in vain endeavoured to discover on what decisive facts the above statement has so often and so boldly been made. There would be great difficulty in proving its truth: we may safely conclude that very many of the most strongly marked domestic varieties could not possibly live in a wild state. In many cases we do not know what the aboriginal stock was, and so could not tell whether or not nearly perfect reversion had ensued. It would be necessary, in order to prevent the effects of intercrossing, that only a single variety should be turned loose in its new home. Nevertheless, as our varieties certainly do occasionally revert in some of their characters to ancestral forms, it seems to me not improbable that if we could succeed in naturalising, or were to cultivate, during many generations, the several races, for instance, of the cabbage, in very poor soil—in which case, however, some effect would have to be attributed to the definite action of the poor soil—that they would, to a large extent, or even wholly, revert to the wild aboriginal stock. Whether or not the experiment would succeed is not of great importance for our line of argument; for by the experiment itself the conditions of life are changed. If it could be shown that our domestic varieties manifested a strong tendency to reversion—that is, to lose their acquired characters, while kept under the same conditions and while kept in a considerable body, so that free intercrossing might check, by blending together, any slight deviations in their structure, in such case, I grant that we could deduce nothing from domestic varieties in regard to species. But there is not a shadow of evidence in favour of this view: to assert that we could not breed our cart and race-horses, long and short-horned cattle, and poultry of various breeds, and esculent vegetables, for an unlimited number of generations, would be opposed to all experience.
Character of Domestic Varieties; difficulty of distinguishing between Varieties and Species; origin of Domestic Varieties from one or more Species.
When we look to the hereditary varieties or races of our domestic animals and plants, and compare them with closely allied species, we generally perceive in each domestic race, as already remarked, less uniformity of character than in true species. Domestic races often have a somewhat monstrous character; by which I mean, that, although differing from each other and from other species of the same genus, in several trifling respects, they often differ in an extreme degree in some one part, both when compared one with another, and more especially when compared with the species under nature to which they are nearest allied. With these exceptions (and with that of the perfect fertility of varieties when crossed—a subject hereafter to be discussed), domestic races of the same species differ from each other in the same manner as do the closely allied species of the same genus in a state of nature, but the differences in most cases are less in degree. This must be admitted as true, for the domestic races of many animals and plants have been ranked by some competent judges as the descendants of aboriginally distinct species, and by other competent judges as mere varieties. If any well marked distinction existed between a domestic race and a species, this source of doubt would not so perpetually recur. It has often been stated that domestic races do not differ from each other in characters of generic value. It can be shown that this statement is not correct; but naturalists differ much in determining what characters are of generic value; all such valuations being at present empirical. When it is explained how genera originate under nature, it will be seen that we have no right to expect often to find a generic amount of difference in our domesticated races.
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 the different quarters of the world. I do not believe, as we shall presently see, that the whole amount of difference between the several breeds of the dog has been produced under domestication; I believe that a small part of the difference is due to their being descended from distinct species. In the case of strongly marked races of some other domesticated species, there is presumptive or even strong evidence that all are descended from a single wild stock.
It has often been assumed that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary, and likewise to withstand diverse climates. I do not dispute that these capacities have added largely to the value of most of our domesticated productions; but how could a savage possibly know, when he first tamed an animal, whether it would vary in succeeding generations, and whether it would endure other climates? Has the little variability of the ass and goose, or the small power of endurance of warmth by the reindeer, or of cold by the common camel, prevented their domestication? I cannot doubt that if other animals and plants, equal in number to our domesticated productions, and belonging to equally diverse classes and countries, were taken from a state of nature, and could be made to breed for an equal number of generations under domestication, they would on an average vary as largely as the parent species of our existing domesticated productions have varied.
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 civilisation, 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 civilisation; and this again implies a long continued previous period of less advanced civilisation, 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 men 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.
The origin of most of our domestic animals will probably forever remain vague. But I may here state that, looking to the domestic dogs of the whole world, I have, after a laborious collection of all known facts, come to the conclusion that several wild species of Canidæ have been tamed, and that their blood, in some cases mingled together, flows in the veins of our domestic breeds. In regard to sheep and goats I can form no decided opinion. From facts communicated to me by Mr. Blyth, on the habits, voice, constitution and structure of the humped Indian cattle, it is almost certain that they are descended from a different aboriginal stock from our European cattle; and some competent judges believe that these latter have had two or three wild progenitors, whether or not these deserve to be called species. This conclusion, as well as that of the specific distinction between the humped and common cattle, may, indeed, be looked upon as established by the admirable researches of Professor Rütimeyer. With respect to horses, from reasons which I cannot here give, I am doubtfully inclined to believe, in opposition to several authors, that all the races belong to the same species. Having kept nearly all the English breeds of the fowl alive, having bred and crossed them, and examined their skeletons, it appears to me almost certain that all are the descendants of the wild Indian fowl, Gallus bankiva; and this is the conclusion of Mr. Blyth, and of others who have studied this bird in India. In regard to ducks and rabbits, some breeds of which differ much from each other, the evidence is clear that they are all descended from the common duck and wild rabbit.
The doctrine of the origin of our several domestic races from several aboriginal stocks, has been carried to an absurd extreme by some authors. They believe that every race which breeds true, let the distinctive characters be ever so slight, has had its wild prototype. At this rate there must have existed at least a score of species of wild cattle, as many sheep, and several goats, in Europe alone, and several even within Great Britain. One author believes that there formerly existed eleven wild species of sheep peculiar to Great Britain! When we bear in mind that Britain has now not one peculiar mammal, and France but few distinct from those of Germany, and so with Hungary, Spain, &c., but that each of these kingdoms possesses several peculiar breeds of cattle, sheep, &c., we must admit that many domestic breeds must have originated in Europe; for whence otherwise could they have been derived? So it is in India. Even in the case of the breeds of the domestic dog throughout the world, which I admit are descended from several wild species, it cannot be doubted that there has been an immense amount of inherited variation; for who will believe that animals closely resembling the Italian greyhound, the bloodhound, the bull-dog, pug-dog, or Blenheim spaniel, &c.—so unlike all wild Canidæ—ever existed in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species; but by crossing we can only get forms in some degree intermediate between their parents; and if we account for our several domestic races by this process, we must admit the former existence of the most extreme forms, as the Italian greyhound, bloodhound, bull-dog, &c., in the wild state. Moreover, the possibility of making distinct races by crossing has been greatly exaggerated. Many cases are on record showing that a race may be modified by occasional crosses if aided by the careful selection of the individuals which present the desired character; but to obtain a race intermediate between two quite distinct races would be very difficult. Sir J. Sebright expressly experimented with this object and failed. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) quite uniform in character, and every thing seems simple enough; but when these mongrels are crossed one with another for several generations, hardly two of them are alike, and then the difficulty of the task becomes manifest.
Breeds of the Domestic Pigeon, their Differences and Origin.
Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the Hon. W. Elliot from India, and by the Hon. C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs. The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head, and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels. The runt is a bird of great size, with long, massive beak and large feet; some of the sub-breeds of runts have very long necks, others very long wings and tails, others singularly short tails. The barb is allied to the carrier, but, instead of a long beak, has a very short and broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter. The turbit has a short and conical beak, with a line of reversed feathers down the breast; and it has the habit of continually expanding, slightly, the upper part of the œsophagus. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood, and it has, proportionally to its size, elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds. The fantail has thirty or even forty tail-feathers, instead of twelve or fourteen, the normal number in all the members of the great pigeon family: these feathers are kept expanded and are carried so erect that in good birds the head and tail touch: the oil-gland is quite aborted. Several other less distinct breeds might be specified.
In the skeletons of the several breeds, the development of the bones of the face, in length and breadth and curvature, differs enormously. The shape, as well as the breadth and length of the ramus of the lower jaw, varies in a highly remarkable manner. The caudal and sacral vertebræ vary in number; as does the number of the ribs, together with their relative breadth and the presence of processes. The size and shape of the apertures in the sternum are highly variable; so is the degree of divergence and relative size of the two arms of the furcula. The proportional width of the gape of mouth, the proportional length of the eyelids, of the orifice of the nostrils, of the tongue (not always in strict correlation with the length of beak), the size of the crop and of the upper part of the œsophagus; the development and abortion of the oil-gland; the number of the primary wing and caudal feathers; the relative length of the wing and tail to each other and to the body; the relative length of the leg and foot; the number of scutellæ on the toes, the development of skin between the toes, are all points of structure which are variable. The period at which the perfect plumage is acquired varies, as does the state of the down with which the nestling birds are clothed when hatched. The shape and size of the eggs vary. The manner of flight, and in some breeds the voice and disposition, differ remarkably. Lastly, in certain breeds, the males and females have come to differ in a slight degree from each other.
Altogether at least a score of pigeons might be chosen, which, if shown to an ornithologist, and he were told that they were wild birds, would certainly be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would in this case place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail in the same genus; more especially as in each of these breeds several truly-inherited sub-breeds, or species, as he would call them, could be shown him.
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 flyers, 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, colouring, 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.
Some facts in regard to the colouring of pigeons well deserve consideration. The rock-pigeon is of a slaty-blue, with white loins; but the Indian sub-species, C. intermedia of Strickland, has this part bluish. The tail has a terminal dark bar, with the outer feathers externally edged at the base with white. The wings have two black bars. Some semi-domestic breeds, and some truly wild breeds, have, besides the two black bars, the wings chequered with black. These several marks do not occur together in any other species of the whole family. Now, in every one of the domestic breeds, taking thoroughly well-bred birds, all the above marks, even to the white edging of the outer tail-feathers, sometimes concur perfectly developed. Moreover, when birds belonging to two or more distinct breeds are crossed, none of which are blue or have any of the above-specified marks, the mongrel offspring are very apt suddenly to acquire these characters. To give one instance out of several which I have observed: I crossed some white fantails, which breed very true, with some black barbs—and it so happens that blue varieties of barbs are so rare that I never heard of an instance in England; and the mongrels were black, brown and mottled. I also crossed a barb with a spot, which is a white bird with a red tail and red spot on the forehead, and which notoriously breeds very true; the mongrels were dusky and mottled. I then crossed one of the mongrel barb-fantails with a mongrel barb-spot, and they produced a bird of as beautiful a blue colour, with the white loins, double black wing-bar, and barred and white-edged tail-feathers, as any wild rock-pigeon! We can understand these facts, on the well-known principle of reversion to ancestral characters, if all the domestic breeds are descended from the rock-pigeon. But if we deny this, we must make one of the two following highly improbable suppositions. Either, first, that all the several imagined aboriginal stocks were coloured and marked like the rock-pigeon, although no other existing species is thus coloured and marked, so that in each separate breed there might be a tendency to revert to the very same colours and markings. Or, secondly, that each breed, even the purest, has within a dozen, or at most within a score, of generations, been crossed by the rock-pigeon: I say within a dozen or twenty generations, for no instance is known of crossed descendants reverting to an ancestor of foreign blood, removed by a greater number of generations. In a breed which has been crossed only once the tendency to revert to any character derived from such a cross will naturally become less and less, as in each succeeding generation there will be less of the foreign blood; but when there has been no cross, and there is a tendency in the breed to revert to a character which was lost during some former generation, this tendency, for all that we can see to the contrary, may be transmitted undiminished for an indefinite number of generations. These two distinct cases of reversion are often confounded together by those who have written on inheritance.
Lastly, the hybrids or mongrels from between all the breeds of the pigeon are perfectly fertile, as I can state from my own observations, purposely made, on the most distinct breeds. Now, hardly any cases have been ascertained with certainty of hybrids from two quite distinct species of animals being perfectly fertile. Some authors believe that long-continued domestication eliminates this strong tendency to sterility in species. From the history of the dog, and of some other domestic animals, this conclusion is probably quite correct, if applied to species closely related to each other. But to extend it so far as to suppose that species, aboriginally as distinct as carriers, tumblers, pouters, and fantails now are, should yield offspring perfectly fertile, inter se, seems to me rash in the extreme.
From these several reasons, namely, the improbability of man having formerly made seven or eight supposed species of pigeons to breed freely under domestication—these supposed species being quite unknown in a wild state, and their not having become anywhere feral—these species presenting certain very abnormal characters, as compared with all other Columbidæ, though so like the rock-pigeon in most other respects—the occasional reappearance of the blue colour and various black marks in all the breeds, both when kept pure and when crossed—and lastly, the mongrel offspring being perfectly fertile—from these several reasons, taken together, we may safely conclude that all our domestic breeds are descended from the rock-pigeon or Columba livia with its geographical sub-species.
In favour of this view, I may add, firstly, that the wild C. livia has been found capable of domestication in Europe and in India; and that it agrees in habits and in a great number of points of structure with all the domestic breeds. Secondly, that although an English carrier or a short-faced tumbler differs immensely in certain characters from the rock-pigeon, yet that by comparing the several sub-breeds of these two races, more especially those brought from distant countries, we can make, between them and the rock-pigeon, an almost perfect series; so we can in some other cases, but not with all the breeds. Thirdly, those characters which are mainly distinctive of each breed are in each eminently variable, for instance, the wattle and length of beak of the carrier, the shortness of that of the tumbler, and the number of tail-feathers in the fantail; and the explanation of this fact will be obvious when we treat of selection. Fourthly, pigeons have been watched and tended with the utmost care, and loved by many people. They have been domesticated for thousands of years in several quarters of the world; the earliest known record of pigeons is in the fifth Ægyptian dynasty, about 3000 B.C., as was pointed out to me by Professor Lepsius; but Mr. Birch informs me that pigeons are given in a bill of fare in the previous dynasty. In the time of the Romans, as we hear from Pliny, immense prices were given for pigeons; “nay, they are come to this pass, that they can reckon up their pedigree and race.” Pigeons were much valued by Akber Khan in India, about the year 1600; never less than 20,000 pigeons were taken with the court. “The monarchs of Iran and Turan sent him some very rare birds;” and, continues the courtly historian, “His Majesty, by crossing the breeds, which method was never practised before, has improved them astonishingly.” About this same period the Dutch were as eager about pigeons as were the old Romans. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone, will likewise be obvious when we treat of Selection. We shall then, also, see how it is that the several breeds so often have a somewhat monstrous character. It is also a most favourable circumstance for the production of distinct breeds, that male and female pigeons can be easily mated for life; and thus different breeds can be kept together in the same aviary.
I have discussed the probable origin of domestic pigeons at some, yet quite insufficient, length; because when I first kept pigeons and watched the several kinds, well knowing how truly they breed, I felt fully as much difficulty in believing that since they had been domesticated they had all proceeded from a common parent, as any naturalist could in coming to a similar conclusion in regard to the many species of finches, or other groups of birds, in nature. One circumstance has struck me much; namely, that nearly all the breeders of the various domestic animals and the cultivators of plants, with whom I have conversed, or whose treatises I have read, are firmly convinced that the several breeds to which each has attended, are descended from so many aboriginally distinct species. Ask, as I have asked, a celebrated raiser of Hereford cattle, whether his cattle might not have descended from Long-horns, or both from a common parent-stock, and he will laugh you to scorn. I have never met a pigeon, or poultry, or duck, or rabbit fancier, who was not fully convinced that each main breed was descended from a distinct species. Van Mons, in his treatise on pears and apples, shows how utterly he disbelieves that the several sorts, for instance a Ribston-pippin or Codlin-apple, could ever have proceeded from the seeds of the same tree. Innumerable other examples could be given. The explanation, I think, is simple: from long-continued study they are strongly impressed with the differences between the several races; and though they well know that each race varies slightly, for they win their prizes by selecting such slight differences, yet they ignore all general arguments, and refuse to sum up in their minds slight differences accumulated during many successive generations. May not those naturalists who, knowing far less of the laws of inheritance than does the breeder, and knowing no more than he does of the intermediate links in the long lines of descent, yet admit that many of our domestic races are descended from the same parents—may they not learn a lesson of caution, when they deride the idea of species in a state of nature being lineal descendants of other species?
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-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.
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 space I could quote numerous passages to this effect from highly competent authorities. Youatt, who was probably better acquainted with the works of agriculturalists than almost any other individual, and who was himself a very good judge of 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 classed, so that the very best may ultimately be selected for breeding.
What English breeders have actually effected is proved by the enormous prices given for animals with a good pedigree; and these have 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 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 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 astonishing improvement in many florists’ flowers, when the flowers of the present day are compared with drawings made only twenty or thirty years ago. When a race of plants is once pretty well established, the seed-raisers do not pick out the best plants, but merely go over their seed-beds, and pull up the “rogues,” as they call the plants that deviate from the proper standard. With animals this kind of selection is, in fact, 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 accumulated 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 passed to prevent their exportation: the destruction of horses under a certain size was ordered, and this may be compared to the “roguing” of plants by nurserymen. The principle of selection I find distinctly given in an ancient Chinese encyclopædia. Explicit rules are laid down by some of the Roman classical writers. From passages in Genesis, it is clear that the colour of domestic animals was at that early period attended to. Savages now sometimes cross their dogs with wild canine animals, to improve the breed, and they formerly did so, as is attested by passages in Pliny. The savages in South Africa match their draught cattle by colour, as do some of the Esquimaux their teams of dogs. Livingstone states that good domestic breeds are highly valued by the negroes in the interior of Africa who have not associated with Europeans. Some of these facts do not show actual selection, but they show that the breeding of domestic animals was carefully attended to in ancient times, and is now attended to by the lowest savages. It would, indeed, have been a strange fact, had attention not been paid to breeding, for the inheritance of good and bad qualities is so obvious.
Unconscious Selection.
At the present time, eminent breeders try by methodical selection, with a distinct object in view, to make a new strain or sub-breed, superior to anything of the kind in the country. But, for our purpose, a form of selection, which may be called unconscious, and which results from every one trying to possess and breed from the best individual animals, is more important. Thus, a man who intends keeping pointers naturally tries to get as good dogs as he can, and afterwards breeds from his own best dogs, but he has no wish or expectation of permanently altering the breed. Neverth ×