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Title: The Variation of Animals and Plants Under Domestication, Vol. I.

Author: Charles Darwin

Release Date: March 27, 2008 [EBook #24923]

Language: English

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THE VARIATION

OF

ANIMALS AND PLANTS

UNDER DOMESTICATION.

By CHARLES DARWIN, M.A., F.R.S., &c.

IN TWO VOLUMES.—Vol. I.

WITH ILLUSTRATIONS.

LONDON:

JOHN MURRAY, ALBEMARLE STREET.

1868.

The right of Translation is reserved.

BY THE SAME AUTHOR.

ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or The Preservation of Favoured Races in the Struggle for Life. Fourth Edition (Eighth Thousand), with Additions and Corrections. 1866. ... Murray.

A NATURALIST'S VOYAGE ROUND THE WORLD; or, A Journal of Researches into the Natural History and Geology of the Countries visited during the Voyage of H.M.S. Beagle, under the Command of Capt. Fitz-Roy, R.N. Tenth Thousand. ... Murray.

ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. ... Smith, Elder, & Co.

GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. ... Smith, Elder, & Co.

GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. ... Smith, Elder, & Co.

A MONOGRAPH OF THE CIRRIPEDIA. With numerous Illustrations. 2 vols. 8vo. ... Hardwicke.

ON THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND FOREIGN ORCHIDS ARE FERTILISED BY INSECTS; and on the Good Effects of Crossing. With numerous Woodcuts. ... Murray.

ON THE MOVEMENTS and HABITS of CLIMBING PLANTS. With Woodcuts. ... Williams & Norgate.

LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, AND CHARING CROSS.

{iii}

CONTENTS OF VOLUME I.

INTRODUCTION ... Page 1

CHAPTER I.

DOMESTIC DOGS AND CATS.

ANCIENT VARIETIES OF THE DOG—RESEMBLANCE OF DOMESTIC DOGS IN VARIOUS COUNTRIES TO NATIVE CANINE SPECIES—ANIMALS NOT ACQUAINTED WITH MAN AT FIRST FEARLESS—DOGS RESEMBLING WOLVES AND JACKALS—HABIT OF BARKING ACQUIRED AND LOST—FERAL DOGS—TAN-COLOURED EYE-SPOTS—PERIOD OF GESTATION—OFFENSIVE ODOUR—FERTILITY OF THE RACES WHEN CROSSED—DIFFERENCES IN THE SEVERAL RACES IN PART DUE TO DESCENT FROM DISTINCT SPECIES—DIFFERENCES IN THE SKULL AND TEETH—DIFFERENCES IN THE BODY, IN CONSTITUTION—FEW IMPORTANT DIFFERENCES HAVE BEEN FIXED BY SELECTION—DIRECT ACTION OF CLIMATE—WATER-DOGS WITH PALMATED FEET—HISTORY OF THE CHANGES WHICH CERTAIN ENGLISH RACES OF THE DOG HAVE GRADUALLY UNDERGONE THROUGH SELECTION—EXTINCTION OF THE LESS IMPROVED SUB-BREEDS.

CATS, CROSSED WITH SEVERAL SPECIES—DIFFERENT BREEDS FOUND ONLY IN SEPARATED COUNTRIES—DIRECT EFFECTS OF THE CONDITIONS OF LIFE—FERAL CATS—INDIVIDUAL VARIABILITY ... Page 15

CHAPTER II.

HORSES AND ASSES.

HORSE.—DIFFERENCES IN THE BREEDS—INDIVIDUAL VARIABILITY OF—DIRECT EFFECTS OF THE CONDITIONS OF LIFE—CAN WITHSTAND MUCH COLD—BREEDS MUCH MODIFIED BY SELECTION—COLOURS OF THE HORSE—DAPPLING—DARK STRIPES ON THE SPINE, LEGS, SHOULDERS, AND FOREHEAD—DUN-COLOURED HORSES MOST FREQUENTLY STRIPED—STRIPES PROBABLY DUE TO REVERSION TO THE PRIMITIVE STATE OF THE HORSE.

ASSES.—BREEDS OF—COLOUR OF—LEG- AND SHOULDER-STRIPES—SHOULDER-STRIPES SOMETIMES ABSENT, SOMETIMES FORKED ... Page 49

CHAPTER III.

PIGS—CATTLE—SHEEP—GOATS.

PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICA—TORF-SCHWEIN—JAPAN PIG—FERTILITY OF CROSSED PIGS—CHANGES IN THE SKULL OF THE HIGHLY CULTIVATED RACES—CONVERGENCE OF CHARACTER—GESTATION—SOLID-HOOFED SWINE—CURIOUS APPENDAGES TO THE JAWS—DECREASE IN SIZE OF THE TUSKS—YOUNG PIGS LONGITUDINALLY STRIPED—FERAL PIGS—CROSSED BREEDS.

CATTLE.—ZEBU A DISTINCT SPECIES—EUROPEAN CATTLE PROBABLY DESCENDED FROM THREE WILD FORMS—ALL THE RACES NOW FERTILE TOGETHER—BRITISH PARK CATTLE—ON THE COLOUR OF THE ABORIGINAL SPECIES—CONSTITUTIONAL DIFFERENCES—SOUTH AFRICAN RACES—SOUTH AMERICAN RACES—NIATA CATTLE—ORIGIN OF THE VARIOUS RACES OF CATTLE. {iv}

SHEEP.—REMARKABLE RACES OF—VARIATIONS ATTACHED TO THE MALE SEX—ADAPTATIONS TO VARIOUS CONDITIONS—GESTATION OF—CHANGES IN THE WOOL—SEMI-MONSTROUS BREEDS.

GOATS.—REMARKABLE VARIATIONS OF ... Page 65

CHAPTER IV.

DOMESTIC RABBITS.

DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RABBIT—ANCIENT DOMESTICATION—ANCIENT SELECTION—LARGE LOP-EARED RABBITS—VARIOUS BREEDS—FLUCTUATING CHARACTERS—ORIGIN OF THE HIMALAYAN BREED—CURIOUS CASE OF INHERITANCE—FERAL RABBITS IN JAMAICA AND THE FALKLAND ISLANDS—PORTO SANTO FERAL RABBITS—OSTEOLOGICAL CHARACTERS—SKULL—SKULL OF HALF-LOP RABBITS—VARIATIONS IN THE SKULL ANALOGOUS TO DIFFERENCES IN DIFFERENT SPECIES OF HARES—VERTEBRÆ—STERNUM—SCAPULA—EFFECTS OF USE AND DISUSE ON THE PROPORTIONS OF THE LIMBS AND BODY—CAPACITY OF THE SKULL AND REDUCED SIZE OF THE BRAIN—SUMMARY ON THE MODIFICATIONS OF DOMESTICATED RABBITS ... Page 103

CHAPTER V.

DOMESTIC PIGEONS.

ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS—INDIVIDUAL VARIABILITY—VARIATIONS OF A REMARKABLE NATURE—OSTEOLOGICAL CHARACTERS: SKULL, LOWER JAW, NUMBER OF VERTEBRÆ—CORRELATION OF GROWTH: TONGUE WITH BEAK; EYELIDS AND NOSTRILS WITH WATTLED SKIN—NUMBER OF WING-FEATHERS, AND LENGTH OF WING—COLOUR AND DOWN—WEBBED AND FEATHERED FEET—ON THE EFFECTS OF DISUSE—LENGTH OF FEET IN CORRELATION WITH LENGTH OF BEAK—LENGTH OF STERNUM, SCAPULA, AND FURCULA—LENGTH OF WINGS—SUMMARY ON THE POINTS OF DIFFERENCE IN THE SEVERAL BREEDS ... Page 131

CHAPTER VI.

PIGEONS—continued.

ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES—HABITS OF LIFE—WILD RACES OF THE ROCK-PIGEON—DOVECOT-PIGEONS—PROOFS OF THE DESCENT OF THE SEVERAL RACES FROM COLUMBA LIVIA—FERTILITY OF THE RACES WHEN CROSSED—REVERSION TO THE PLUMAGE OF THE WILD ROCK-PIGEON—CIRCUMSTANCES FAVOURABLE TO THE FORMATION OF THE RACES—ANTIQUITY AND HISTORY OF THE PRINCIPAL RACES—MANNER OF THEIR FORMATION—SELECTION—UNCONSCIOUS SELECTION—CARE TAKEN BY FANCIERS IN SELECTING THEIR BIRDS—SLIGHTLY DIFFERENT STRAINS GRADUALLY CHANGE INTO WELL-MARKED BREEDS—EXTINCTION OF INTERMEDIATE FORMS—CERTAIN BREEDS REMAIN PERMANENT, WHILST OTHERS CHANGE—SUMMARY ... Page 180

{v}

CHAPTER VII.

FOWLS.

BRIEF DESCRIPTIONS OF THE CHIEF BREEDS—ARGUMENTS IN FAVOUR OF THEIR DESCENT FROM SEVERAL SPECIES—ARGUMENTS IN FAVOUR OF ALL THE BREEDS HAVING DESCENDED FROM GALLUS BANKIVA—-REVERSION TO THE PARENT-STOCK IN COLOUR—ANALOGOUS VARIATIONS—ANCIENT HISTORY OF THE FOWL—EXTERNAL DIFFERENCES BETWEEN THE SEVERAL BREEDS—EGGS—CHICKENS—SECONDARY SEXUAL CHARACTERS—WING- AND TAIL-FEATHERS, VOICE, DISPOSITION, ETC.—OSTEOLOGICAL DIFFERENCES IN THE SKULL, VERTEBRÆ, ETC.—EFFECTS OF USE AND DISUSE ON CERTAIN PARTS—CORRELATION OF GROWTH ... Page 225

CHAPTER VIII.

DUCKS—GOOSE—PEACOCK—TURKEY—GUINEA-FOWL—CANARY-BIRD—GOLD-FISH—HIVE-BEES—SILK-MOTHS.

DUCKS, SEVERAL BREEDS OF—PROGRESS OF DOMESTICATION—ORIGIN OF, FROM THE COMMON WILD-DUCK—DIFFERENCES IN THE DIFFERENT BREEDS—OSTEOLOGICAL DIFFERENCES—EFFECTS OF USE AND DISUSE ON THE LIMB-BONES.

GOOSE, ANCIENTLY DOMESTICATED—LITTLE VARIATION OF—SEBASTOPOL BREED.

PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.

TURKEY, BREEDS OF—CROSSED WITH THE UNITED STATES SPECIES—EFFECTS OF CLIMATE ON.

GUINEA-FOWL, CANARY-BIRD, GOLD-FISH, HIVE-BEES.

SILK-MOTHS, SPECIES AND BREEDS OF—ANCIENTLY DOMESTICATED—CARE IN THEIR SELECTION—DIFFERENCES IN THE DIFFERENT RACES—IN THE EGG, CATERPILLAR, AND COCOON STATES—INHERITANCE OF CHARACTERS—IMPERFECT WINGS—LOST INSTINCTS—CORRELATED CHARACTERS ... Page 276

CHAPTER IX.

CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.

PRELIMINARY REMARKS ON THE NUMBER AND PARENTAGE OF CULTIVATED PLANTS—FIRST STEPS IN CULTIVATION—GEOGRAPHICAL DISTRIBUTION OF CULTIVATED PLANTS.

CEREALIA.—DOUBTS ON THE NUMBER OF SPECIES.—WHEAT: VARIETIES OF—INDIVIDUAL VARIABILITY—CHANGED HABITS—SELECTION—ANCIENT HISTORY OF THE VARIETIES.—MAIZE: GREAT VARIATION OF—DIRECT ACTION OF CLIMATE ON.

CULINARY PLANTS.—CABBAGES: VARIETIES OF, IN FOLIAGE AND STEMS, BUT NOT IN OTHER PARTS—PARENTAGE OF—OTHER SPECIES OF BRASSICA.—PEAS: AMOUNT OF DIFFERENCE IN THE SEVERAL KINDS, CHIEFLY IN THE PODS AND SEED—SOME VARIETIES CONSTANT, SOME HIGHLY VARIABLE—DO NOT INTERCROSS.—BEANS.—POTATOES: NUMEROUS VARIETIES OF—DIFFERING LITTLE, EXCEPT IN THE TUBERS—CHARACTERS INHERITED ... Page 305

{vi}

CHAPTER X.

PLANTS continued—FRUITS—ORNAMENTAL TREES—FLOWERS.

FRUITS.—GRAPES—VARY IN ODD AND TRIFLING PARTICULARS.—MULBERRY.—THE ORANGE GROUP—SINGULAR RESULTS FROM CROSSING.—PEACH AND NECTARINE—BUD-VARIATION—ANALOGOUS VARIATION—RELATION TO THE ALMOND.—APRICOT.—PLUMS—VARIATION IN THEIR STONES.—CHERRIES—SINGULAR VARIETIES OF.—APPLE.—PEAR.—STRAWBERRY—INTERBLENDING OF THE ORIGINAL FORMS.—GOOSEBERRY—STEADY INCREASE IN SIZE OF THE FRUIT—VARIETIES OF.—WALNUT.—NUT.—CUCURBITACEOUS PLANTS—WONDERFUL VARIATION OF.

ORNAMENTAL TREES—THEIR VARIATION IN DEGREE AND KIND—ASH-TREE—SCOTCH-FIR—HAWTHORN.

FLOWERS—MULTIPLE ORIGIN OF MANY KINDS—VARIATION IN CONSTITUTIONAL PECULIARITIES—KIND OF VARIATION.—ROSES—SEVERAL SPECIES CULTIVATED.—PANSY.—DAHLIA.—HYACINTH, HISTORY AND VARIATION OF ... Page 332

CHAPTER XI.

ON BUD-VARIATION, AND ON CERTAIN ANOMALOUS MODES OF REPRODUCTION AND VARIATION.

BUD-VARIATIONS IN THE PEACH, PLUM, CHERRY, VINE, GOOSEBERRY, CURRANT, AND BANANA, AS SHOWN BY THE MODIFIED FRUIT—IN FLOWERS: CAMELLIAS, AZALEAS, CHRYSANTHEMUMS, ROSES, ETC.—ON THE RUNNING OF THE COLOUR IN CARNATIONS—BUD-VARIATIONS IN LEAVES—VARIATIONS BY SUCKERS, TUBERS, AND BULBS—ON THE BREAKING OF TULIPS—BUD-VARIATIONS GRADUATE INTO CHANGES CONSEQUENT ON CHANGED CONDITIONS OF LIFE—CYTISUS ADAMI, ITS ORIGIN AND TRANSFORMATION—ON THE UNION OF TWO DIFFERENT EMBRYOS IN ONE SEED—THE TRIFACIAL ORANGE—ON REVERSION BY BUDS IN HYBRIDS AND MONGRELS—ON THE PRODUCTION OF MODIFIED BUDS BY THE GRAFTING OF ONE VARIETY OR SPECIES ON ANOTHER—ON THE DIRECT OR IMMEDIATE ACTION OF FOREIGN POLLEN ON THE MOTHER-PLANT—ON THE EFFECTS IN FEMALE ANIMALS OF A FIRST IMPREGNATION ON THE SUBSEQUENT OFFSPRING—CONCLUSION AND SUMMARY ... Page 373

{vii}

LIST OF ILLUSTRATIONS.

{1}

THE

VARIATION OF ANIMALS AND PLANTS

UNDER DOMESTICATION.

INTRODUCTION.

The object of this work is not to describe all the many races of animals which have been domesticated by man, and of the plants which have been cultivated by him; even if I possessed the requisite knowledge, so gigantic an undertaking would be here superfluous. It is my intention to give under the head of each species only such facts as I have been able to collect or observe, showing the amount and nature of the changes which animals and plants have undergone whilst under man's dominion, or which bear on the general principles of variation. In one case alone, namely in that of the domestic pigeon, I will describe fully all the chief races, their history, the amount and nature of their differences, and the probable steps by which they have been formed. I have selected this case, because, as we shall hereafter see, the materials are better than in any other; and one case fully described will in fact illustrate all others. But I shall also describe domesticated rabbits, fowls, and ducks, with considerable fullness.

The subjects discussed in this volume are so connected that it is not a little difficult to decide how they can be best arranged. I have determined in the first part to give, under the heads of the various animals and plants, a large body of facts, some of which may at first appear but little related to our subject, and to devote the latter part to general discussions. Whenever I have found it necessary to give numerous details, in support of any proposition or conclusion, small type has been used. The reader {2}will, I think, find this plan a convenience, for, if he does not doubt the conclusion or care about the details, he can easily pass them over; yet I may be permitted to say that some of the discussions thus printed deserve attention, at least from the professed naturalist.

It may be useful to those who have read nothing about Natural Selection, if I here give a brief sketch of the whole subject and of its bearing on the origin of species.^[1] This is the more desirable, as it is impossible in the present work to avoid many allusions to questions which will be fully discussed in future volumes.

From a remote period, in all parts of the world, man has subjected many animals and plants to domestication or culture. Man has no power of altering the absolute conditions of life; he cannot change the climate of any country; he adds no new element to the soil; but he can remove an animal or plant from one climate or soil to another, and give it food on which it did not subsist in its natural state. It is an error to speak of man "tampering with nature" and causing variability. If organic beings had not possessed an inherent tendency to vary, man could have done nothing.^[2] He unintentionally exposes his animals and plants to various conditions of life, and variability supervenes, which he cannot even prevent or check. Consider the simple case of a plant which has been cultivated during a long time in its native country, and which consequently has not been subjected to any change of climate. It has been protected to a certain extent from the competing roots of plants of other kinds; it has generally been grown in manured soil, but probably not richer than that of many an alluvial flat; and lastly, it has been exposed to changes in its conditions, being grown sometimes in one district and sometimes in another, in different soils. Under such circumstances, {3}scarcely a plant can be named, though cultivated in the rudest manner, which has not given birth to several varieties. It can hardly be maintained that during the many changes which this earth has undergone, and during the natural migrations of plants from one land or island to another, tenanted by different species, that such plants will not often have been subjected to changes in their conditions analogous to those which almost inevitably cause cultivated plants to vary. No doubt man selects varying individuals, sows their seeds, and again selects their varying offspring. But the initial variation on which man works, and without which he can do nothing, is caused by slight changes in the conditions of life, which must often have occurred under nature. Man, therefore, may be said to have been trying an experiment on a gigantic scale; and it is an experiment which nature during the long lapse of time has incessantly tried. Hence it follows that the principles of domestication are important for us. The main result is that organic beings thus treated have varied largely, and the variations have been inherited. This has apparently been one chief cause of the belief long held by some few naturalists that species in a state of nature undergo change.

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. We shall learn something on the laws of inheritance, on the effects of crossing different breeds, and on that sterility which often supervenes when organic beings are removed from their natural conditions of life, and likewise when they are too closely interbred. During this investigation we shall see that the principle of Selection is all important. Although man does not cause variability and cannot even prevent it, he can select, preserve, and accumulate the variations given to him by the hand of nature 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 {4}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.

In a second work I shall discuss the variability of organic beings in a state of nature; namely, the individual differences presented by animals and plants, and those slightly greater and generally inherited differences which are ranked by naturalists as varieties or geographical races. We shall see how difficult, or rather how impossible it often is, to distinguish between races and sub-species, as the less well-marked forms have sometimes been denominated; and again between sub-species and true species. I shall further attempt to show that it is the common and widely ranging, or, as they may be called, the dominant species, which most frequently vary; and that it is the large and flourishing genera which include the greatest number of varying species. Varieties, as we shall see, may justly be called incipient species.

But it may be urged, granting that organic beings in a state of nature present some varieties,—that their organization is in some slight degree plastic; granting that many animals and plants have varied greatly under domestication, and that man by his power of selection has gone on accumulating such variations until he has made strongly marked and firmly inherited races; granting all this, how, it may be asked, have species arisen in a state of nature? The differences between natural varieties are slight; whereas the differences are {5}considerable between the species of the same genus, and great between the species of distinct genera. How do these lesser differences become augmented into the greater difference? How do varieties, or as I have called them incipient species, become converted into true and well-defined species? How has each new species been adapted to the surrounding physical conditions, and to the other forms of life on which it in any way depends? We see on every side of us innumerable adaptations and contrivances, which have justly excited in the mind of every observer the highest admiration. There is, for instance, a fly (Cecidomyia)^[3] which deposits its eggs within the stamens of a Scrophularia, and secretes a poison which produces a gall, on which the larva feeds; but there is another insect (Misocampus) which deposits its eggs within the body of the larva within the gall, and is thus nourished by its living prey; so that here a hymenopterous insect depends on a dipterous insect, and this depends on its power of producing a monstrous growth in a particular organ of a particular plant. So it is, in a more or less plainly marked manner, in thousands and tens of thousands of cases, with the lowest as well as with the highest productions of nature.

This problem of the conversion of varieties into species,—that is, the augmentation of the slight differences characteristic of varieties into the greater differences characteristic of species and genera, including the admirable adaptations of each being to its complex organic and inorganic conditions of life,—will form the main subject of my second work. We shall therein see that all organic beings, without exception, tend to increase at so high a ratio, that no district, no station, not even the whole surface of the land or the whole ocean, would hold the progeny of a single pair after a certain number of generations. The inevitable result is an ever-recurrent Struggle for Existence. It has truly been said that all nature is at war; the strongest ultimately prevail, the weakest fail; and we well know that myriads of forms have disappeared from the face of the earth. If then organic beings in a state of nature vary even in a slight degree, owing to changes in the surrounding {6}conditions, of which we have abundant geological evidence, or from any other cause; if, in the long course of ages, inheritable variations ever arise in any way advantageous to any being under its excessively complex and changing relations of life; and it would be a strange fact if beneficial variations did never arise, seeing how many have arisen which man has taken advantage of for his own profit or pleasure; if then these contingencies ever occur, and I do not see how the probability of their occurrence can be doubted, then the severe and often-recurrent struggle for existence will determine that those variations, however slight, which are favourable shall be preserved or selected, and those which are unfavourable shall be destroyed.

This preservation, during the battle for life, of varieties which possess any advantage in structure, constitution, or instinct, I have called Natural Selection; and Mr. Herbert Spencer has well expressed the same idea by the Survival of the Fittest. The term "natural selection" is in some respects a bad one, as it seems to imply conscious choice; but this will be disregarded after a little familiarity. No one objects to chemists speaking of "elective affinity;" and certainly an acid has no more choice in combining with a base, than the conditions of life have in determining whether or not a new form be selected or preserved. The term is so far a good one as it brings into connection the production of domestic races by man's power of selection, and the natural preservation of varieties and species in a state of nature. For brevity sake I sometimes speak of natural selection as an intelligent power;—in the same way as astronomers speak of the attraction of gravity as ruling the movements of the planets, or as agriculturists speak of man making domestic races by his power of selection. In the one case, as in the other, selection does nothing without variability, and this depends in some manner on the action of the surrounding circumstances on the organism. I have, also, often personified the word Nature; for I have found it difficult to avoid this ambiguity; but I mean by nature only the aggregate action and product of many natural laws,—and by laws only the ascertained sequence of events. {7}

In the chapter devoted to natural selection I shall show from experiment and from a multitude of facts, that the greatest amount of life can be supported on each spot by great diversification or divergence in the structure and constitution of its inhabitants. We shall, also, see that the continued production of new forms through natural selection, which implies that each new variety has some advantage over others, almost inevitably leads to the extermination of the older and less improved forms. These latter are almost necessarily intermediate in structure as well as in descent between the last-produced forms and their original parent-species. Now, if we suppose a species to produce two or more varieties, and these in the course of time to produce other varieties, the principle of good being derived from diversification of structure will generally lead to the preservation of the most divergent varieties; thus the lesser differences characteristic of varieties come to be augmented into the greater differences characteristic of species, and, by the extermination of the older intermediate forms, new species come to be distinctly defined objects. Thus, also, we shall see how it is that organic beings can be classed by what is called a natural method in distinct groups—species under genera, and genera under families.

As all the inhabitants of each country may be said, owing to their high rate of reproduction, to be striving to increase in numbers; as each form is related to many other forms in the struggle for life,—for destroy any one and its place will be seized by others; as every part of the organization occasionally varies in some slight degree, and as natural selection acts exclusively by the preservation of variations which are advantageous under the excessively complex conditions to which each being is exposed, no limit exists to the number, singularity, and perfection of the contrivances and co-adaptations which may thus be produced. An animal or a plant may thus slowly become related in its structure and habits in the most intricate manner to many other animals and plants, and to the physical conditions of its home. Variations in the organization will in some cases be aided by habit, or by the use and disuse of parts, and they will be governed by the direct action {8}of the surrounding physical conditions and by correlation of growth.

On the principles here briefly sketched out, there is no innate or necessary tendency in each being to its own advancement in the scale of organization. We are almost compelled to look at the specialization or differentiation of parts or organs for different functions as the best or even sole standard of advancement; for by such division of labour each function of body and mind is better performed. And, as natural selection acts exclusively through the preservation of profitable modifications of structure, and as the conditions of life in each area generally become more and more complex, from the increasing number of different forms which inhabit it and from most of these forms acquiring a more and more perfect structure, we may confidently believe, that, on the whole, organization advances. Nevertheless a very simple form fitted for very simple conditions of life might remain for indefinite ages unaltered or unimproved; for what would it profit an infusorial animalcule, for instance, or an intestinal worm, to become highly organized? Members of a high group might even become, and this apparently has occurred, fitted for simpler conditions of life; and in this case natural selection would tend to simplify or degrade the organization, for complicated mechanism for simple actions would be useless or even disadvantageous.

In a second work, after treating of the Variation of organisms in a state of nature, of the Struggle for Existence and the principle of Natural Selection, I shall discuss the difficulties which are opposed to the theory. These difficulties may be classed under the following heads:—the apparent impossibility in some cases of a very simple organ graduating by small steps into a highly perfect organ; the marvellous facts of Instinct; the whole question of Hybridity; and, lastly, the absence, at the present time and in our geological formations, of innumerable links connecting all allied species. Although some of these difficulties are of great weight, we shall see that many of them are explicable on the theory of natural selection, and are otherwise inexplicable.

In scientific investigations it is permitted to invent any hypothesis, and if it explains various large and independent classes of facts it rises to the rank of a well-grounded theory. The {9}undulations of the ether and even its existence are hypothetical, yet every one now admits the undulatory theory of light. The principle of natural selection may be looked at as a mere hypothesis, but rendered in some degree probable by what we positively know of the variability of organic beings in a state of nature,—by what we positively know of the struggle for existence, and the consequent almost inevitable preservation of favourable variations,—and from the analogical formation of domestic races. Now this hypothesis may be tested,—and this seems to me the only fair and legitimate manner of considering the whole question,—by trying whether it explains several large and independent classes of facts; such as the geological succession of organic beings, their distribution in past and present times, and their mutual affinities and homologies. If the principle of natural selection does explain these and other large bodies of facts, it ought to be received. On the ordinary view of each species having been independently created, we gain no scientific explanation of any one of these facts. We can only say that it has so pleased the Creator to command that the past and present inhabitants of the world should appear in a certain order and in certain areas; that He has impressed on them the most extraordinary resemblances, and has classed them in groups subordinate to groups. But by such statements we gain no new knowledge; we do not connect together facts and laws; we explain nothing.

In a third work I shall try the principle of natural selection by seeing how far it will give a fair explanation of the several classes of facts just alluded to. It was the consideration of these facts which first led me to take up the present subject. When I visited, during the voyage of H.M.S. Beagle, the Galapagos Archipelago, situated in the Pacific Ocean about 500 miles from the shore of South America, I found myself surrounded by peculiar species of birds, reptiles, and plants, existing nowhere else in the world. Yet they nearly all bore an American stamp. In the song of the mocking-thrush, in the harsh cry of the carrion-hawk, in the great candlestick-like opuntias, I clearly perceived the neighbourhood of America, though the islands were separated by so many miles of ocean from the mainland, and differed much from it in their geological {10}constitution and climate. Still more surprising was the fact that most of the inhabitants of each separate island in this small archipelago were specifically different, though most closely related to each other. The archipelago, with its innumerable craters and bare streams of lava, appeared to be of recent origin; and thus I fancied myself brought near to the very act of creation. I often asked myself how these many peculiar animals and plants had been produced: the simplest answer seemed to be that the inhabitants of the several islands had descended from each other, undergoing modification in the course of their descent; and that all the inhabitants of the archipelago had descended from those of the nearest land, namely America, whence colonists would naturally have been derived. But it long remained to me an inexplicable problem how the necessary degree of modification could have been effected, and it would have thus remained for ever, had I not studied domestic productions, and thus acquired a just idea of the power of Selection. As soon as I had fully realized this idea, I saw, on reading Malthus on Population, that Natural Selection was the inevitable result of the rapid increase of all organic beings; for I was prepared to appreciate the struggle for existence by having long studied the habits of animals.

Before visiting the Galapagos I had collected many animals whilst travelling from north to south on both sides of America, and everywhere, under conditions of life as different as it is possible to conceive, American forms were met with—species replacing species of the same peculiar genera. Thus it was when the Cordilleras were ascended, or the thick tropical forests penetrated, or the fresh waters of America searched. Subsequently I visited other countries, which in all the conditions of life were incomparably more like to parts of South America, than the different parts of that continent were to each other; yet in these countries, as in Australia or Southern Africa, the traveller cannot fail to be struck with the entire difference of their productions. Again the reflection was forced on me that community of descent from the early inhabitants or colonists of South America would alone explain the wide prevalence of American types of structure throughout that immense area.

To exhume with one's own hands the bones of extinct and {11}gigantic quadrupeds brings the whole question of the succession of species vividly before one's mind; and I had found in South America great pieces of tesselated armour exactly like, but on a magnificent scale, that covering the pigmy armadillo; I had found great teeth like those of the living sloth, and bones like those of the cavy. An analogous succession of allied forms had been previously observed in Australia. Here then we see the prevalence, as if by descent, in time as in space, of the same types in the same areas; and in neither case does the similarity of the conditions by any means seem sufficient to account for the similarity of the forms of life. It is notorious that the fossil remains of closely consecutive formations are closely allied in structure, and we can at once understand the fact if they are likewise closely allied by descent. The succession of the many distinct species of the same genus throughout the long series of geological formations seems to have been unbroken or continuous. New species come in gradually one by one. Ancient and extinct forms of life often show combined or intermediate characters, like the words of a dead language with respect to its several offshoots or living tongues. All these and other such facts seemed to me to point to descent with modification as the method of production of new groups of species.

The innumerable past and present inhabitants of the world are connected together by the most singular and complex affinities, and can be classed in groups under groups, in the same manner as varieties can be classed under species and sub-varieties under varieties, but with much higher grades of difference. It will be seen in my third work that these complex affinities and the rules for classification receive a rational explanation on the principle of descent, together with modifications acquired through natural selection, entailing divergence of character and the extinction of intermediate forms. How inexplicable is the similar pattern of the hand of a man, the foot of a dog, the wing of a bat, the flipper of a seal, on the doctrine of independent acts of creation! how simply explained on the principle of the natural selection of successive slight variations in the diverging descendants from {12}a single progenitor! So it is, if we look to the structure of an individual animal or plant, when we see the fore and hind limbs, the skull and vertebræ, the jaws and legs of a crab, the petals, stamens, and pistils of a flower, built on the same type or pattern. During the many changes to which in the course of time all organic beings have been subjected, certain organs or parts have occasionally become at first of little use and ultimately superfluous; and the retention of such parts in a rudimentary and utterly useless condition can, on the descent-theory, be simply understood. On the principle of modifications being inherited at the same age in the child, at which each successive variation first appeared in the parent, we shall see why rudimentary parts and organs are generally well developed in the individual at a very early age. On the same principle of inheritance at corresponding ages, and on the principle of variations not generally supervening at a very early period of embryonic growth (and both these principles can be shown to be probable from direct evidence), that most wonderful fact in the whole round of natural history, namely, the similarity of members of the same great class in their embryonic condition,—the embryo, for instance, of a mammal, bird, reptile, and fish being barely distinguishable,—becomes simply intelligible.

It is the consideration and explanation of such facts as these which has convinced me that the theory of descent with modification by means of natural selection is in the main true. These facts have as yet received no explanation on the theory of independent Creations; they cannot be grouped together under one point of view, but each has to be considered as an ultimate fact. As the first origin of life on this earth, as well as the continued life of each individual, is at present quite beyond the scope of science, I do not wish to lay much stress on the greater simplicity of the view of a few forms, or of only one form, having been originally created, instead of innumerable miraculous creations having been necessary at innumerable periods; though this more simple view accords well with Maupertuis's philosophical axiom "of least action."

In considering how far the theory of natural selection may be {13}extended,—that is, in determining from how many progenitors the inhabitants of the world have descended,—we may conclude that at least all the members of the same class have descended from a single ancestor. A number of organic beings are included in the same class, because they present, independently of their habits of life, the same fundamental type of structure, and because they graduate into each other. Moreover, members of the same class can in most cases be shown to be closely alike at an early embryonic age. These facts can be explained on the belief of their descent from a common form; therefore it may be safely admitted that all the members of the same class have descended from one progenitor. But as the members of quite distinct classes have something in common in structure and much in common in constitution, analogy and the simplicity of the view would lead us one step further, and to infer as probable that all living creatures have descended from a single prototype.

I hope that the reader will pause before coming to any final and hostile conclusion on the theory of natural selection. It is the facts and views to be hereafter given which have convinced me of the truth of the theory. The reader may consult my 'Origin of Species,' for a general sketch of the whole subject; but in that work he has to take many statements on trust. In considering the theory of natural selection, he will assuredly meet with weighty difficulties, but these difficulties relate chiefly to subjects—such as the degree of perfection of the geological record, the means of distribution, the possibility of transitions in organs, &c.—on which we are confessedly ignorant; nor do we know how ignorant we are. If we are much more ignorant than is generally supposed, most of these difficulties wholly disappear. Let the reader reflect on the difficulty of looking at whole classes of facts from a new point of view. Let him observe how slowly, but surely, the noble views of Lyell on the gradual changes now in progress on the earth's surface have been accepted as sufficient to account for all that we see in its past history. The present action of natural selection may seem more or less probable; but I believe in the truth of the theory, {14}because it collects under one point of view, and gives a rational explanation of, many apparently independent classes of facts.^[4]

{15}

CHAPTER I.

DOMESTIC DOGS AND CATS.

ANCIENT VARIETIES OF THE DOG—RESEMBLANCE OF DOMESTIC DOGS IN VARIOUS COUNTRIES TO NATIVE CANINE SPECIES—ANIMALS NOT ACQUAINTED WITH MAN AT FIRST FEARLESS—DOGS RESEMBLING WOLVES AND JACKALS—HABIT OF BARKING ACQUIRED AND LOST—FERAL DOGS—TAN-COLOURED EYE-SPOTS PERIOD OF GESTATION—OFFENSIVE ODOUR—FERTILITY OF THE RACES WHEN CROSSED—DIFFERENCES IN THE SEVERAL RACES IN PART DUE TO DESCENT FROM DISTINCT SPECIES—DIFFERENCES IN THE SKULL AND TEETH—DIFFERENCES IN THE BODY, IN CONSTITUTION—FEW IMPORTANT DIFFERENCES HAVE BEEN FIXED BY SELECTION—DIRECT ACTION OF CLIMATE—WATER-DOGS WITH PALMATED FEET—HISTORY OF THE CHANGES WHICH CERTAIN ENGLISH RACES OF THE DOG HAVE GRADUALLY UNDERGONE THROUGH SELECTION—EXTINCTION OF THE LESS IMPROVED SUB-BREEDS.

CATS, CROSSED WITH SEVERAL SPECIES—DIFFERENT BREEDS FOUND ONLY IN SEPARATED COUNTRIES—DIRECT EFFECTS OF THE CONDITIONS OF LIFE—FERAL CATS—INDIVIDUAL VARIABILITY.

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 favourite 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. Palæontology^[5] 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 {16}later tertiary deposits more like those of a large dog than of a wolf, which favours 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 the 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.

The reasons which have led various authors to infer that our dogs have descended from more than one wild species are as follows.^[6] Firstly, the great difference between the several breeds; but this will appear of comparatively little weight, after we shall have seen how great are the differences between the several races of various domesticated animals which certainly have descended from a single parent-form. Secondly, the more important fact that, at the most anciently known historical periods, several breeds of the dog existed, very unlike each other, and closely resembling or identical with breeds still alive.

We will briefly run back through the historical records. The materials are remarkably deficient between the fourteenth century and the Roman classical period.^[7] At this earlier period {17}various breeds, namely hounds, house-dogs, lapdogs, &c., existed; but as Dr. Walther has remarked it is impossible to recognise the greater number with any certainty. Youatt, however, gives a drawing of a beautiful sculpture of two greyhound puppies from the Villa of Antoninus. On an Assyrian monument, about 640 B.C., an enormous mastiff^[8] is figured; and according to Sir H. Rawlinson (as I was informed at the British Museum), similar dogs are still imported into this same country. I have looked through the magnificent works of Lepsius and Rosellini, and on the monuments from the fourth to the twelfth dynasties (i.e. from about 3400 B.C. to 2100 B.C.) several varieties of the dog are represented; most of them are allied to greyhounds; at the later of these periods a dog resembling a hound is figured, with drooping ears, but with a longer back and more pointed head than in our hounds. There is, also, a turnspit, with short and crooked legs, closely resembling the existing variety; but this kind of monstrosity is so common with various animals, as with the ancon sheep, and even, according to Rengger, with jaguars in Paraguay, that it would be rash to look at the monumental animal as the parent of all our turnspits: Colonel Sykes^[9] also has described an Indian Pariah dog as presenting the same monstrous character. The most ancient dog represented on the Egyptian monuments is one of the most singular; it resembles a greyhound, but has long pointed ears and a short curled tail: a closely allied variety still exists in Northern Africa; for Mr. E. Vernon Harcourt^[10] states that the Arab boar-hound is "an eccentric hieroglyphic animal, such as Cheops once hunted with, somewhat resembling the rough Scotch deer-hound; their tails are curled tight round on their backs, {18}and their ears stick out at right angles." With this most ancient variety a pariah-like dog coexisted.

We thus see that, at a period between four and five thousand years ago, various breeds, viz. pariah dogs, greyhounds, common hounds, mastiffs, house-dogs, lapdogs, 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.^[11] As long as man was believed to have existed on this earth only about 6000 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 embedded 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.

Long before the period of any historical record the dog was domesticated in Europe. In the Danish Middens of the Neolithic or Newer Stone period, bones of a canine animal are imbedded, and Steenstrup ingeniously argues that these belonged to a domestic dog; for a very large proportion of the bones of birds preserved in the refuse, consists of long bones, which it was found on trial dogs cannot devour.^[12] This ancient dog was succeeded in Denmark during the Bronze period by a larger kind, presenting certain differences, and this again during the Iron period, by a still larger kind. In Switzerland, we hear {19}from Prof. Rütimeyer,^[13] that during the Neolithic period a domesticated dog of middle size existed, which in its skull was about equally remote from the wolf and jackal, and partook of the characters of our hounds and setters or spaniels (Jagdhund und Wachtelhund). Rütimeyer insists strongly on the constancy of form during a very long period of time of this the most ancient known dog. During the Bronze period a larger dog appeared, and this closely resembled in its jaw a dog of the same age in Denmark. Remains of two notably distinct varieties of the dog were found by Schmerling in a cave;^[14] but their age cannot be positively determined.

The existence of a single race, remarkably constant in form during the whole Neolithic period, is an interesting fact in contrast with what we see of the changes which the races underwent during the period of the successive Egyptian monuments, and in contrast with our existing dogs. The character of this animal during the Neolithic period, as given by Rütimeyer, supports De Blainville's view that our varieties have descended from an unknown and extinct form. But we should not forget that we know nothing with respect to the antiquity of man in the warmer parts of the world. The succession of the different kinds of dogs in Switzerland and Denmark is thought to be due to the immigration of conquering tribes bringing with them their dogs; and this view accords with the belief that different wild canine animals were domesticated in different regions. Independently of the immigration of new races of man, we know from the wide-spread presence of bronze, composed of an alloy of tin, how much commerce there must have been throughout Europe at an extremely remote period, and dogs would then probably have been bartered. At the present time, amongst the savages of the interior of Guiana, the Taruma Indians are considered the best trainers of dogs, and possess a large breed, which they barter at a high price with other tribes.^[15]

The main argument in favour of the several breeds of the {20}dog being the descendants of distinct wild stocks, is their resemblance in various countries to distinct species still existing there. It must, however, be admitted that the comparison between the wild and domesticated animal has been made but in few cases with sufficient exactness. Before entering on details, it will be well to show that there is no a priori difficulty in the belief that several canine species have been domesticated; for there is much difficulty in this respect with some other domestic quadrupeds and birds. Members of the dog family inhabit nearly the whole world; and several species agree pretty closely in habits and structure with our several domesticated dogs. Mr. Galton has shown^[16] how fond savages are of keeping and taming animals of all kinds. Social animals are the most easily subjugated by man, and several species of Canidæ hunt in packs. It deserves notice, as bearing on other animals as well as on the dog, that at an extremely ancient period, when man first entered any country, the animals living there would have felt no instinctive or inherited fear of him, and would consequently have been tamed far more easily than at present. For instance, when the Falkland Islands were first visited by man, the large wolf-like dog (Canis antarcticus) fearlessly came to meet Byron's sailors, who, mistaking this ignorant curiosity for ferocity, ran into the water to avoid them: even recently a man, by holding a piece of meat in one hand and a knife in the other, could sometimes stick them at night. On an island in the Sea of Aral, when first discovered by Butakoff, the saigak antelopes, which are "generally very timid and watchful, did not fly from us, but on the contrary looked at us with a sort of curiosity." So, again, on the shores of the Mauritius, the manatee was not at first in the least afraid of man, and thus it has been in several quarters of the world with seals and the morse. I have elsewhere shown^[17] how slowly the native birds of several islands have acquired and inherited a salutary dread of man: at the Galapagos Archipelago I pushed with the muzzle of my gun hawks from a branch, and {21}held out a pitcher of water for other birds to alight on and drink. Quadrupeds and birds which have seldom been disturbed by man, dread him no more than do our English birds the cows or horses grazing in the fields.

It is a more important consideration that several canine species evince (as will be shown in a future chapter) no strong repugnance or inability to breed under confinement; and the incapacity to breed under confinement is one of the commonest bars to domestication. Lastly, savages set the highest value, as we shall see in the chapter on Selection, on dogs: even half-tamed animals are highly useful to them: the Indians of North America cross their half-wild dogs with wolves, and thus render them even wilder than before, but bolder: the savages of Guiana catch and partially tame and use the whelps of two wild species of Canis, as do the savages of Australia those of the wild Dingo. Mr. Philip King informs me that he once trained a wild Dingo puppy to drive cattle, and found it very useful. From these several considerations we see that there is no difficulty in believing that man might have domesticated various canine species in different countries. It would indeed have been a strange fact if one species alone had been domesticated throughout the world.

We will now enter into details. The accurate and sagacious Richardson says, "The resemblance between the Northern American wolves (Canis lupus, var. occidentalis) and the domestic dogs of the Indians is so great that the size and strength of the wolf seems to be the only difference. I have more than once mistaken a band of wolves for the dogs of a party of Indians; and the howl of the animals of both species is prolonged so exactly in the same key that even the practised ear of the Indian fails at times to discriminate them." He adds that the more northern Esquimaux dogs are not only extremely like the grey wolves of the Arctic circle in form and colour, but also nearly equal them in size. Dr. Kane has often seen in his teams of sledge-dogs the oblique eye (a character on which some naturalists lay great stress), the drooping tail, and scared look of the wolf. In disposition the Esquimaux dogs differ little from wolves, and, according to Dr. Hayes, they are capable of no attachment to man, and are so savage, that {22}when hungry they will attack even their masters. According to Kane they readily become feral. Their affinity is so close with wolves that they frequently cross with them, and the Indians take the whelps of wolves "to improve the breed of their dogs." The half-bred wolves sometimes (Lamare-Picquot) cannot be tamed, "though this case is rare;" but they do not become thoroughly well broken in till the second or third generation. These facts show that there can be but little, if any, sterility between the Esquimaux dog and the wolf, for otherwise they would not be used to improve the breed. As Dr. Hayes says of these dogs, "reclaimed wolves they doubtless are."^[18]

North America is inhabited by a second kind of wolf, the prairie-wolf (Canis latrans), which is now looked at by all naturalists as specifically distinct from the common wolf; and is, according to Mr. J. K. Lord, in some respects intermediate in habits between a wolf and a fox. Sir J. Richardson, after describing the Hare Indian dog, which differs in many respects from the Esquimaux dog, says, "It bears the same relation to the prairie wolf that the Esquimaux dog does to the great grey wolf." He could, in fact, detect no marked difference between them; and Messrs. Nott and Gliddon give additional details showing their close resemblance. The dogs derived from the above two aboriginal sources cross together and with the wild wolves, at least with the C. occidentalis, and with European dogs. In Florida, according to Bartram, the black wolf-dog of the Indians differs in nothing from the wolves of that country except in barking.^[19]

{23}

Turning to the southern parts of the New World, Columbus found two kinds of dogs in the West Indies; and Fernandez^[20] describes three in Mexico: some of these native dogs were dumb—that is, did not bark. In Guiana it has been known since the time of Buffon that the natives cross their dogs with an aboriginal species, apparently the Canis cancrivorus. Sir R. Schomburgk, who has so carefully explored these regions, writes to me, "I have been repeatedly told by the Arawaak Indians, who reside near the coast, that they cross their dogs with a wild species to improve the breed, and individual dogs have been shown to me which certainly resembled the C. cancrivorus much more than the common breed. It is but seldom that the Indians keep the C. cancrivorus for domestic purposes, nor is the Ai, another species of wild dog, and which I consider to be identical with the Dusicyon silvestris of H. Smith, now much used by the Arecunas for the purpose of hunting. The dogs of the Taruma Indians are quite distinct, and resemble Buffon's St. Domingo greyhound." It thus appears that the natives of Guiana have partially domesticated two aboriginal species, and still cross their dogs with them; these two species belong to a quite different type from the North American and European wolves. A careful observer, Rengger,^[21] gives reasons for believing that a hairless dog was domesticated when America was first visited by Europeans: some of these dogs in Paraguay are still dumb, and Tschudi^[22] states that they suffer from cold in the Cordillera. This naked dog is, however, quite distinct from that found preserved in the ancient Peruvian burial-places, and described by Tschudi, under the name of Canis Ingæ, as withstanding cold well and as barking. It is not known whether these two distinct kinds of dog are the descendants of native species, and it might be argued that when man first migrated into America he brought with him from the Asiatic continent dogs {24}which had not learned to bark; but this view does not seem probable, as the natives along the line of their march from the north reclaimed, as we have seen, at least two N. American species of Canidæ.

Turning to the Old World, some European dogs closely resemble the wolf; thus the shepherd dog of the plains of Hungary is white or reddish-brown, has a sharp nose, short, erect ears, shaggy coat, and bushy tail, and so much resembles a wolf that Mr. Paget, who gives this description, says he has known a Hungarian mistake a wolf for one of his own dogs. Jeitteles, also, remarks on the close similarity of the Hungarian dog and wolf. Shepherd dogs in Italy must anciently have closely resembled wolves, for Columella (vii. 12) advises that white dogs be kept, adding, "pastor album probat, ne pro lupo canem feriat." Several accounts have been given of dogs and wolves crossing naturally; and Pliny asserts that the Gauls tied their female dogs in the woods that they might cross with wolves.^[23] The European wolf differs slightly from that of North America, and has been ranked by many naturalists as a distinct species. The common wolf of India is also by some esteemed as a third species, and here again we find a marked resemblance between the pariah dogs of certain districts of India and the Indian wolf.^[24]

With respect to Jackals, Isidore Geoffroy Saint Hilaire^[25] says that not one constant difference can be pointed out between their structure and that of the smaller races of dogs. They agree closely in habits: jackals, when tamed and called by their {25}master, wag their tails, crouch, and throw themselves on their backs; they smell at the tails of dogs, and void their urine sideways.^[26] A number of excellent naturalists, from the time of Güldenstädt to that of Ehrenberg, Hemprich, and Cretzschmar, have expressed themselves in the strongest terms with respect to the resemblance of the half-domestic dogs of Asia and Egypt to jackals. M. Nordmann, for instance, says, "Les chiens d'Awhasie ressemblent étonnamment à des chacals." Ehrenberg^[27] asserts that the domestic dogs of Lower Egypt, and certain mummied dogs, have for their wild type a species of wolf (C. lupaster) of the country; whereas the domestic dogs of Nubia and certain other mummied dogs have the closest relation to a wild species of the same country, viz. C. sabbar, which is only a form of the common jackal. Pallas asserts that jackals and dogs sometimes naturally cross in the East; and a case is on record in Algeria.^[28] The greater number of naturalists divide the jackals of Asia and Africa into several species, but some few rank them all as one.

I may add that the domestic dogs on the coast of Guinea are fox-like animals, and are dumb.^[29] On the east coast of Africa, between lat. 4° and 6° south, and about ten days' journey in the interior, a semi-domestic dog, as the Rev. S. Erhardt informs me, is kept, which the natives assert is derived from a similar wild animal. Lichtenstein^[30] says that the dogs of the Bosjemans present a striking resemblance even in colour (excepting the black stripe down the back) with the C. mesomelas of South Africa. Mr. E. Layard informs me that he has seen a Caffre dog which closely resembled an Esquimaux dog. In Australia the Dingo is both domesticated and wild; though this animal may have been introduced aboriginally by man, yet it must be considered as almost an endemic form, for its remains have been found in a similar state of preservation and associated with {26}extinct mammals, so that its introduction must have been ancient.^[31]

From this resemblance in several countries of the half-domesticated dogs 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 favour their domestication, it is highly probable that the domestic dogs of the world have descended from two good species of wolf (viz. C. lupus and C. latrans), and from two or three other doubtful species of wolves (namely, the European, Indian, and North African forms); from at least one or two South American canine species; from several races or species of the jackal; and perhaps from one or more extinct species. Those authors who attribute great influence to the action of climate by itself may thus account for the resemblance of the domesticated dogs and native animals in the same countries; but I know of no facts supporting the belief in so powerful an action of climate.

It cannot be objected to the view of several canine species having been anciently domesticated, that these animals are tamed with difficulty: facts have been already given on this head, but I may add that the young of the Canis primævus of India were tamed by Mr. Hodgson,^[32] and became as sensible to caresses, and manifested as much intelligence, as any sporting dog of the same age. There is not much difference, as we have already shown and shall immediately further see, in habits between the domestic dogs of the North American Indians and the wolves of that country, or between the Eastern pariah dogs and jackals, or between the dogs which have run wild in various countries and the several natural species of the family. The habit of barking, however, which is almost universal with domesticated {27}dogs, and which does not characterise a single natural species of the family, seems an exception; but this habit is soon lost and soon reacquired. The case of the wild dogs on the island of Juan Fernandez having become dumb has often been quoted, and there is reason to believe^[33] that the dumbness ensued in the course of thirty-three years; on the other hand, dogs taken from this island by Ulloa slowly reacquired the habit of barking. The Mackenzie-river dogs, of the Canis latrans type, when brought to England, never learned to bark properly; but one born in the Zoological Gardens^[34] "made his voice sound as loudly as any other dog of the same age and size." According to Professor Nillson,^[35] a wolf-whelp reared by a bitch barks. I. Geoffroy Saint Hilaire exhibited a jackal which barked with the same tone as any common dog.^[36] An interesting account has been given by Mr. G. Clarke^[37] of some dogs run wild on Juan de Nova, in the Indian Ocean; "they had entirely lost the faculty of barking; they had no inclination for the company of other dogs, nor did they acquire their voice," during a captivity of several months. On the island they "congregate in vast packs, and catch sea-birds with as much address as foxes could display." The feral dogs of La Plata have not become dumb; they are of large size, hunt single or in packs, and burrow holes for their young.^[38] In these habits the feral dogs of La Plata resemble wolves and jackals; both of which hunt either singly or in packs, and burrow holes.^[39] These feral dogs have not become uniform in colour on Juan Fernandez, Juan de Nova, or La Plata.^[40] In Cuba the feral dogs are described by Poeppig as nearly all mouse-coloured, with short ears and light-blue eyes. {28}In St. Domingo, Col. Ham. Smith says^[41] that the feral dogs are very large, like greyhounds, of a uniform pale blue-ash, with small ears, and large light-brown eyes. Even the wild Dingo, though so anciently naturalised in Australia, "varies considerably in colour," as I am informed by Mr. P. P. King: a half-bred Dingo reared in England^[42] showed signs of wishing to burrow.

From the several foregoing facts we see that reversion in the feral state gives no indication of the colour or size of the aboriginal parent-species. One fact, however, with respect to the colouring of domestic dogs, I at one time hoped might have thrown some light on their origin; and it is worth giving, as showing how colouring follows laws, even in so anciently and thoroughly domesticated an animal as the dog. Black dogs with tan-coloured feet, whatever breed they may belong to, almost invariably have a tan-coloured spot on the upper and inner corners of each eye, and their lips are generally thus coloured. I have seen only two exceptions to this rule, namely, in a spaniel and terrier. Dogs of a light-brown colour often have a lighter, yellowish-brown spot over the eyes; sometimes the spot is white, and in a mongrel terrier the spot was black. Mr. Waring kindly examined for me a stud of fifteen greyhounds in Suffolk: eleven of them were black, or black and white, or brindled, and these had no eye-spots; but three were red and one slaty-blue, and these four had dark-coloured spots over their eyes. Although the spots thus sometimes differ in colour, they strongly tend to be tan-coloured; this is proved by my having seen four spaniels, a setter, two Yorkshire shepherd dogs, a large mongrel, and some fox-hounds, coloured black and white, with not a trace of tan-colour, excepting the spots over the eyes, and sometimes a little on the feet. These latter cases, and many others, show plainly that the colour of the feet and the eye-spots are in some way correlated. I have noticed, in various breeds, every gradation, from the whole face being tan-coloured, to a complete ring round the eyes, to a minute spot over the inner and upper corners. The spots occur in various sub-breeds of terriers and spaniels; in setters; in hounds of various kinds, including the turnspit-like German badger-hound; in shepherd dogs; in a mongrel, of which neither parent had the spots; in one pure bulldog, though the spots were in this case almost white; and in greyhounds,—but true black-and-tan greyhounds are excessively rare; nevertheless I have been assured by Mr. Warwick, that one ran at the Caledonian Champion meeting of April, 1860, and was "marked precisely like a black-and-tan terrier." Mr. Swinhoe at my request looked at the dogs in China, at Amoy, and he soon noticed a brown dog with yellow spots over the eyes. Colonel H. Smith^[43] figures the magnificent black mastiff of Thibet with a {29}tan-coloured stripe over the eyes, feet, and chaps; and what is more singular, he figures the Alco, or native domestic dog of Mexico, as black and white, with narrow tan-coloured rings round the eyes; at the Exhibition of dogs in London, May, 1863, a so-called forest-dog from North-West Mexico was shown, which had pale tan-coloured spots over the eyes. The occurrence of these tan-coloured spots in dogs of such extremely different breeds, living in various parts of the world, makes the fact highly remarkable.

We shall hereafter see, especially in the chapter on Pigeons, that coloured marks are strongly inherited, and that they often aid us in discovering the primitive forms of our domestic races. Hence, if any wild canine species had distinctly exhibited the tan-coloured spots over the eyes, it might have been argued that this was the parent-form of nearly all our domestic races. But after looking at many coloured plates, and through the whole collection of skins in the British Museum, I can find no species thus marked. It is no doubt possible that some extinct species was thus coloured. On the other hand, in looking at the various species, there seems to be a tolerably plain correlation between tan-coloured legs and face; and less frequently between black legs and a black face; and this general rule of colouring explains to a certain extent the above-given cases of correlation between the eye-spots and the colour of the feet. Moreover, some jackals and foxes have a trace of a white ring round their eyes, as in C. mesomelas, C. aureus, and (judging from Colonel Ham. Smith's drawing) in C. alopex and C. thaleb. Other species have a trace of a black line over the corners of the eyes, as in C. variegatus, cinereo-variegatus, and fulvus, and the wild Dingo. Hence I am inclined to conclude that a tendency for tan-coloured spots to appear over the eyes in the various breeds of dogs, is analogous to the case observed by Desmarest, namely, that when any white appears on a dog the tip of the tail is always white, "de manière a rappeler la tacho terminale de même couleur, qui caractérise la plupart des Canidées sauvages."^[44]

It has been objected that our domestic dogs cannot be descended from wolves or jackals, because their periods of gestation are different. The supposed difference rests on statements made by Buffon, Gilibert, Bechstein, and others; but these are now known to be erroneous; and the period is found to agree in the wolf, jackal, and dog, as closely as could be expected, for it is often in some degree variable.^[45] Tessier, who {30}has closely attended to this subject, allows a difference of four days in the gestation of the dog. The Rev. W. D. Fox has given me three carefully recorded cases of retrievers, in which the bitch was put only once to the dog; and not counting this day, but counting that of parturition, the periods were fifty-nine, sixty-two, and sixty-seven days. The average period is sixty-three days; but Bellingeri states that this holds good only with large dogs; and that for small races it is from sixty to sixty-three days; Mr. Eyton of Eyton, who has had much experience with dogs, also informs me that the time is apt to be longer with large than with small dogs.

F. Cuvier has objected that the jackal would not have been domesticated on account of its offensive smell; but savages are not sensitive in this respect. The degree of odour, also, differs in the different kinds of jackal;^[46] and Colonel H. Smith makes a sectional division of the group with one character dependent on not being offensive. On the other hand, dogs—for instance, rough and smooth terriers—differ much in this respect; and M. Godron states that the hairless so-called Turkish dog is more odoriferous than other dogs. Isidore Geoffroy^[47] gave to a dog the same odour as that from a jackal by feeding it on raw flesh.

The belief that our dogs are descended from wolves, jackals, South American Canidæ, and other species, suggests a far more important difficulty. These animals in their undomesticated state, judging from a widely-spread analogy, would have been in some degree sterile if intercrossed; and such sterility will be admitted as almost certain by all those who believe that the lessened fertility of crossed forms is an infallible criterion of specific distinctness. Anyhow these animals keep distinct in the countries which they inhabit in common. On the other hand, all domestic dogs, which are here supposed to be descended {31}from several distinct species, are, as far as is known, mutually fertile together. But, as Broca has well remarked,^[48] the fertility of successive generations of mongrel dogs has never been scrutinised with that care which is thought indispensable when species are crossed. The few facts leading to the conclusion that the sexual feelings and reproductive powers differ in the several races of the dog when crossed are (passing over mere size as rendering propagation difficult) as follows: the Mexican Alco^[49] apparently dislikes dogs of other kinds, but this perhaps is not strictly a sexual feeling; the hairless endemic dog of Paraguay, according to Rengger, mixes less with the European races than these do with each other; the Spitz-dog in Germany is said to receive the fox more readily than do other breeds; and Dr. Hodgkin states that a female Dingo in England attracted the male wild foxes. If these latter statements can be trusted, they prove some degree of sexual difference in the breeds of the dog. But the fact remains that our domestic dogs, differing so widely as they do in external structure, are far more fertile together than we have reason to believe their supposed wild parents would have been. Pallas assumes^[50] that a long course of domestication eliminates that sterility which the parent-species would have exhibited if only lately captured; no distinct facts are recorded in support of this hypothesis; but the evidence seems to me so strong (independently of the evidence derived from other domesticated animals) in favour of our domestic dogs having descended from several wild stocks, that I am led to admit the truth of this hypothesis.

There is another and closely allied difficulty consequent on the doctrine of the descent of our domestic dogs from several wild species, namely, that they do not seem to be perfectly fertile with their supposed parents. But the experiment has not been quite fairly tried; the Hungarian dog, for instance, {32}which in external appearance so closely resembles the European wolf, ought to be crossed with this wolf; and the pariah-dogs of India with Indian wolves and jackals; and so in other cases. That the sterility is very slight between certain dogs and wolves and other Canidæ is shown by savages taking the trouble to cross them. Buffon got four successive generations from the wolf and dog, and the mongrels were perfectly fertile together.^[51] But more lately M. Flourens states positively as the result of his numerous experiments that hybrids from the wolf and dog, crossed inter se, become sterile at the third generation, and those from the jackal and dog at the fourth generation.^[52] But these animals were closely confined; and many wild animals, as we shall see in a future chapter, are rendered by confinement in some degree or even utterly sterile. The Dingo, which breeds freely in Australia with our imported dogs, would not breed though repeatedly crossed in the Jardin des Plantes.^[53] Some hounds from Central Africa, brought home by Major Denham, never bred in the Tower of London;^[54] and a similar tendency to sterility might be transmitted to the hybrid offspring of a wild animal. Moreover, it appears that in M. Flourens' experiments the hybrids were closely bred in and in for three or four generations; but this circumstance, although it would almost certainly increase the tendency to sterility, would hardly account for the final result, even though aided by close confinement, unless there had been some original tendency to lessened fertility. Several years ago I saw confined in the Zoological Gardens of London a female hybrid from an English dog and jackal, which even in this the first generation was so sterile that, as I was assured by {33}her keeper, she did not fully exhibit her proper periods; but this case, from the numerous instances of fertile hybrids from these two animals, was certainly exceptional. In almost all experiments on the crossing of animals there are so many causes of doubt, that it is extremely difficult to come to any positive conclusion. It would, however, appear, that those who believe that our dogs are descended from several species will have not only to admit that their offspring after a long course of domestication generally lose all tendency to sterility when crossed together; but that between certain breeds of dogs and some of their supposed aboriginal parents a certain degree of sterility has been retained or possibly even acquired.

Notwithstanding the difficulties in regard to fertility given in the last two paragraphs, when we reflect on the inherent improbability of man having domesticated throughout the world one single species alone of so widely distributed, so easily tamed, and so useful a group as the Canidæ; when we reflect on the extreme antiquity of the different breeds; and especially when we reflect on the close similarity, both in external structure and habits, between the domestic dogs of various countries and the wild species still inhabiting these same countries, the balance of evidence is strongly in favour of the multiple origin of our dogs.

Differences between the several Breeds of the Dog.—If the several breeds have descended from several wild stocks, their difference can obviously in part be explained by that of their parent-species. For instance, the form of the greyhound may be partly accounted for by descent from some such animal as the slim Abyssinian Canis simensis,^[55] with its elongated muzzle; that of the larger dogs from the larger wolves, and the smaller and slighter dogs from jackals: and thus perhaps we may account for certain constitutional and climatal differences. But it would be a great error to suppose that there has not been in addition^[56] a large amount of variation. The intercrossing of the several aboriginal wild stocks, and of the subsequently formed {34}races, has probably increased the total number of breeds, and, as we shall presently see, has greatly modified some of them. But we cannot explain by crossing the origin of such extreme forms as thoroughbred greyhounds, bloodhounds, bulldogs, Blenheim spaniels, terriers, pugs, &c., unless we believe that forms equally or more strongly characterised in these different respects once existed in nature. But hardly any one has been bold enough to suppose that such unnatural forms ever did or could exist in a wild state. When compared with all known members of the family of Canidæ they betray a distinct and abnormal origin. No instance is on record of such dogs as bloodhounds, spaniels, true greyhounds having been kept by savages: they are the product of long-continued civilization.

The number of breeds and sub-breeds of the dog is great: Youatt, for instance, describes twelve kinds of greyhounds. I will not attempt to enumerate or describe the varieties, for we cannot discriminate how much of their difference is due to variation, and how much to descent from different aboriginal stocks. But it may be worth while briefly to mention some points. Commencing with the skull, Cuvier has admitted^[57] that in form the differences are "plus fortes que celles d'aucunes espèces sauvages d'un même genre naturel." The proportions of the different bones; the curvature of the lower jaw, the position of the condyles with respect to the plane of the teeth (on which F. Cuvier founded his classification), and in mastiffs the shape of its posterior branch; the shape of the zygomatic arch, and of the temporal fossæ; the position of the occiput—all vary considerably.^[58] The dog has properly six pairs of molar teeth in the upper jaw, and seven in the lower; but several naturalists have seen not rarely an additional pair in the upper jaw;^[59] and Professor Gervais says that there are dogs "qui ont sept paires de dents supérieures et huit inférieures.". De Blainville^[60] has given full particulars on the frequency of these deviations in the number of the teeth, and has shown that it is not always the same tooth which is supernumerary. In short-muzzled races, according to H. Müller,^[61] the molar teeth stand obliquely, whilst in long-muzzled races they are placed longitudinally, with open spaces between them. The naked, so-called Egyptian or Turkish dog is extremely deficient in its {35}teeth,^[62]—sometimes having none except one molar on each side; but this, though characteristic of the breed, must be considered as a monstrosity. M. Girard,^[63] who seems to have attended closely to the subject, says that the period of the appearance of the permanent teeth differs in different dogs, being earlier in large dogs; thus the mastiff assumes its adult teeth in four or five months, whilst in the spaniel the period is sometimes more than seven or eight months.

With respect to minor differences little need be said. Isidore Geoffroy has shown^[64] that in size some dogs are six times as long (the tail being excluded) as others; and that the height relatively to the length of the body varies from between one to two, and one to nearly four. In the Scotch deer-hound there is a striking and remarkable difference in the size of the male and female.^[65] Every one knows how the ears vary in size in different breeds, and with their great development their muscles become atrophied. Certain breeds of dogs are described as having a deep furrow between the nostrils and lips. The caudal vertebræ, according to F. Cuvier, on whose authority the two last statements rest, vary in number; and the tail in shepherd dogs is almost absent. The mammæ vary from seven to ten in number; Daubenton, having examined twenty-one dogs, found eight with five mammæ on each side; eight with four on each side; and the others with an unequal number on the two sides.^[66] Dogs have properly five toes in front and four behind, but a fifth toe is often added; and F. Cuvier states that, when a fifth toe is present, a fourth cuneiform bone is developed; and, in this case, sometimes the great cuneiform bone is raised, and gives on its inner side a large articular surface to the astragalus; so that even the relative connection of the bones, the most constant of all characters, varies. These modifications, however, in the feet of dogs are not important, because they ought to be ranked, as De Blainville has shown,^[67] as monstrosities. Nevertheless they are interesting from being correlated with the size of the body, for they occur much more frequently with mastiffs and other large breeds than with small dogs. Closely allied varieties, however, sometimes differ in this respect; thus Mr. Hodgson states that the black-and-tan Lassa variety of the Thibet mastiff has the fifth digit, whilst the Mustang sub-variety is not thus characterised. The extent to which the skin is developed between the toes varies much; but we shall return to this point. The degree to which the various breeds differ in the perfection of their senses, dispositions, and inherited habits is notorious to every one. The breeds present some constitutional differences: the pulse, says Youatt,^[68] "varies materially according to the breed, as well {36}as to the size of the animal." Different breeds of dogs are subject in different degrees to various diseases. They certainly become adapted to different climates under which they have long existed. It is notorious that most of our best European breeds deteriorate in India.^[69] The Rev. R. Everest^[70] believes that no one has succeeded in keeping the Newfoundland dog long alive in India; so it is, according to Lichtenstein,^[71] even at the Cape of Good Hope. The Thibet mastiff degenerates on the plains of India, and can live only on the mountains.^[72] Lloyd^[73] asserts that our bloodhounds and bulldogs have been tried, and cannot withstand the cold of the northern European forests.

Seeing in how many characters the races of the dog differ from each other, and remembering Cuvier's admission that their skulls differ more than do those of the species of any natural genus, and bearing in mind how closely the bones of wolves, jackals, foxes, and other Canidæ agree, it is remarkable that we meet with the statement, repeated over and over again, that the races of the dog differ in no important characters. A highly competent judge, Prof. Gervais,^[74] admits, "si l'on prenait sans contrôle les altérations dont chacun de ces organes est susceptible, on pourrait croire qu'il y a entre les chiens domestiques des différences plus grandes que celles qui séparent ailleurs les espèces, quelquefois même les genres." Some of the differences above enumerated are in one respect of comparatively little value, for they are not characteristic of distinct breeds: no one pretends that such is the case with the additional molar teeth or with the number of mammæ; the additional digit is generally present with mastiffs, and some of the more important differences in the skull and lower jaw are more or less characteristic of various breeds. But we must not forget that the predominant power of selection has not been applied in any of these cases; we have variability in important parts, but the differences have not been fixed by selection. Man {37}cares for the form and fleetness of his greyhounds, for the size of his mastiffs, for the strength of the jaw in his bulldogs, &c.; but he cares nothing about the number of their molar teeth or mammæ or digits; nor do we know that differences in these organs are correlated with, or owe their development to, differences in other parts of the body about which man does care. Those who have attended to the subject of selection will admit that, nature having given variability, man, if he so chose, could fix five toes to the hinder feet of certain breeds of dogs, as certainly as to the feet of his Dorking-fowls: he could probably fix, but with much more difficulty, an additional pair of molar teeth in either jaw, in the same way as he has given additional horns to certain breeds of sheep; if he wished to produce a toothless breed of dogs, having the so-called Turkish dog with its imperfect teeth to work on, he could probably do so, for he has succeeded in making hornless breeds of cattle and sheep.

With respect to the precise causes and steps by which the several races of dogs have come to differ so greatly from each other, we are, as in most other cases, profoundly ignorant. We may attribute part of the difference in external form and constitution to inheritance from distinct wild stocks, that is to changes effected under nature before domestication. We must attribute something to the crossing of the several domestic and natural races. I shall, however, soon recur to the crossing of races. We have already seen how often savages cross their dogs with wild native species; and Pennant gives a curious account^[75] of the manner in which Fochabers, in Scotland, was stocked "with a multitude of curs of a most wolfish aspect" from a single hybrid-wolf brought into that district.

It would appear that climate to a certain extent directly modifies the forms of dogs. We have lately seen that several of our English breeds cannot live in India, and it is positively asserted that when bred there for a few generations they degenerate not only in their mental faculties, but in form. Captain Williamson,^[76] who carefully attended to this subject, states that "hounds are the most rapid in their decline;" "greyhounds and {38}pointers, also, rapidly decline." But spaniels, after eight or nine generations, and without a cross from Europe, are as good as their ancestors. Dr. Falconer informs me that bulldogs, which have been known, when first brought into the country, to pin down even an elephant by its trunk, not only fall off after two of three generations in pluck and ferocity, but lose the under-hung character of their lower jaws; their muzzles become finer and their bodies lighter. English dogs imported into India are so valuable that probably due care has been taken to prevent their crossing with native dogs; so that the deterioration cannot be thus accounted for. The Rev. R. Everest informs me that he obtained a pair of setters, born in India, which perfectly resembled their Scotch parents: he raised several litters from them in Delhi, taking the most stringent precautions to prevent a cross, but he never succeeded, though this was only the second generation in India, in obtaining a single young dog like its parents in size or make; their nostrils were more contracted, their noses more pointed, their size inferior, and their limbs more slender. This remarkable tendency to rapid deterioration in European dogs subjected to the climate of India, may perhaps partly be accounted for by the tendency to reversion to a primordial condition which many animals exhibit, as we shall see in a future chapter, when exposed to new conditions of life.

Some of the peculiarities characteristic of the several breeds of the dog have probably arisen suddenly, and, though strictly inherited, may be called monstrosities; for instance, the shape of the legs and body in the turnspit of Europe and India; the shape of the head and the under-hanging jaw in the bull and pug-dog, so alike in this one respect and so unlike in all others. A peculiarity suddenly arising, and therefore in one sense deserving to be called a monstrosity, may, however, be increased and fixed by man's selection. We can hardly doubt that long-continued training, as with the greyhound in coursing hares, as with water-dogs in swimming—and the want of exercise, in the case of lapdogs—must have produced some direct effect on their structure and instincts. But we shall immediately see that the most potent cause of change has probably been the selection, both methodical and unconscious, of slight individual differences,—the {39}latter kind of selection resulting from the occasional preservation, during hundreds of generations, of those individual dogs which were the most useful to man for certain purposes and under certain conditions of life. In a future chapter on Selection I shall show that even barbarians attend closely to the qualities of their dogs. This unconscious selection by man would lie aided by a kind of natural selection; for the dogs of savages have partly to gain their own subsistence; for instance, in Australia, as we hear from Mr. Nind,^[77] the dogs are sometimes compelled by want to leave their masters and provide for themselves; but in a few days they generally return. And we may infer that dogs of different shapes, sizes, and habits, would have best chance of surviving under different circumstances,—on open, sterile plains, where they have to run down their own prey,—on rocky coasts, where they have to feed on crabs and fish left in the tidal pools, as in the case of New Guinea and Tierra del Fuego. In this latter country, as I am informed by Mr. Bridges, the Catechist to the Mission, the dogs turn over the stones on the shore to catch the crustaceans which lie beneath, and they "are clever enough to knock off the shell-fish at a first blow;" for if this be not done, shell-fish are well known to have an almost invincible power of adhesion.

It has already been remarked that dogs differ in the degree to which their feet are webbed. In dogs of the Newfoundland breed, which are eminently aquatic in their habits, the skin, according to Isidore Geoffroy,^[78] extends to the third phalanges, whilst in ordinary dogs it extends only to the second. In two Newfoundland dogs which I examined, when the toes were stretched apart and viewed on the under side, the skin extended in a nearly straight line between the outer margins of the balls of the toes; whereas, in two terriers of distinct sub-breeds, the skin viewed in the same manner was deeply scooped out. In Canada there is a dog which is peculiar to the country and common there, and this has "half-webbed feet and is fond of the water."^[79] English otter-hounds are said to have webbed feet: a friend examined for me the feet of two, in comparison {40}with the feet of some harriers and bloodhounds; he found the skin variable in extent in all, but more developed in the otter than in the other hounds.^[80] As aquatic animals which belong to quite different orders have webbed feet, there can be no doubt that this structure would be serviceable to dogs that frequent the water. We may confidently infer that no man ever selected his water-dogs by the extent to which the skin was developed between their toes; but what he does, is to preserve and breed from those individuals which hunt best in the water, or best retrieve wounded game, and thus he unconsciously selects dogs with feet slightly better webbed. Man thus closely imitates Natural Selection. We have an excellent illustration of this same process in North America, where, according to Sir J. Richardson,^[81] all the wolves, foxes, and aboriginal domestic dogs have their feet broader than in the corresponding species of the Old World, and "well calculated for running on the snow." Now, in these Arctic regions, the life or death of every animal will often depend on its success in hunting over the snow when softened; and this will in part depend on the feet being broad; yet they must not be so broad as to interfere with the activity of the animal when the ground is sticky, or with its power of burrowing holes, or with other habits of life.

As changes in domestic breeds which take place so slowly as not to be noticed at any one period, whether due to the selection of individual variations or of differences resulting from crosses, are most important in understanding the origin of our domestic productions, and likewise in throwing indirect light on the changes effected under nature, I will give in detail such cases as I have been able to collect. Lawrence,^[82] who paid particular attention to the history of the foxhound, writing in 1829, says that between eighty and ninety years before "an entirely new foxhound was raised through the breeder's art," the ears of the old southern hound being reduced, the bone and bulk lightened, the waist increased in length, and the stature {41}somewhat added to. It is believed that this was effected by a cross with the greyhound. With respect to this latter dog, Youatt,^[83] who is generally cautious in his statements, says that the greyhound within the last fifty years, that is before the commencement of the present century, "assumed a somewhat different character from that which he once possessed. He is now distinguished by a beautiful symmetry of form, of which he could not once boast, and he has even superior speed to that which he formerly exhibited. He is no longer used to struggle with deer, but contends with his fellows over a shorter and speedier course." An able writer^[84] believes that our English greyhounds are the descendants, progressively improved, of the large rough greyhounds which existed in Scotland so early as the third century. A cross at some former period with the Italian greyhound has been suspected; but this seems hardly probable, considering the feebleness of this latter breed. Lord Orford, as is well known, crossed his famous greyhounds, which failed in courage, with a bulldog—this breed being-chosen from being deficient in the power of scent; "after the sixth or seventh generation," says Youatt, "there was not a vestige left of the form of the bulldog, but his courage and indomitable perseverance remained."

Youatt infers, from a comparison of an old picture of King Charles's spaniels with the living dog, that "the breed of the present day is materially altered for the worse:" the muzzle has become shorter, the forehead more prominent, and the eyes larger: the changes in this case have probably been due to simple selection. The setter, as this author remarks in another place, "is evidently the large spaniel improved to his present peculiar size and beauty, and taught another way of marking his game. If the form of the dog were not sufficiently satisfactory on this point, we might have recourse to history:" he then refers to a document dated 1685 bearing on this subject, and adds that the pure Irish setter shows no signs of a cross with the pointer, which some authors suspect has been the case with the English setter. Another writer^[85] remarks {42}that, if the mastiff and English bulldog had formerly been as distinct as they are at the present time (i.e. 1828), so accurate an observer as the poet Gay (who was the author of 'Rural Sports' in 1711) would have spoken in his Fable of the Bull and the Bulldog, and not of the Bull and the Mastiff. There can be no doubt that the fancy bulldogs of the present day, now that they are not used for bull-baiting, have become greatly reduced in size, without any express intention on the part of the breeder. Our pointers are certainly descended from a Spanish breed, as even their names, Don, Ponto, Carlos, &c., would show: it is said that they were not known in England before the Revolution in 1688;^[86] but the breed since its introduction has been much modified, for Mr. Borrow, who is a sportsman and knows Spain intimately well, informs me that he has not seen in that country any breed "corresponding in figure with the English pointer; but there are genuine pointers near Xeres which have been imported by English gentlemen." A nearly parallel case is offered by the Newfoundland dog, which was certainly brought into England from that country, but which has since been so much modified that, as several writers have observed, it does not now closely resemble any existing native dog in Newfoundland.^[87]

These several cases of slow and gradual changes in our English dogs possess some interest; for though the changes have generally, but not invariably, been caused by one or two crosses with a distinct breed, yet we may feel sure, from the well-known extreme variability of crossed breeds, that rigorous and long-continued selection must have been practised, in order to improve them in a definite manner. As soon as any strain or family became slightly improved or better adapted to altered circumstances, it would tend to supplant the older and less improved strains. For instance, as soon as the old foxhound was improved by a cross with the greyhound, or by simple selection, and assumed its present character—and the change was probably required by {43}the increased fleetness of our hunters—it rapidly spread throughout the country, and is now everywhere nearly uniform. But the process of improvement is still going on, for every one tries to improve his strain by occasionally procuring dogs from the best kennels. Through this process of gradual substitution the old English hound has been lost; and so it has been with the old Irish greyhound and apparently with the old English bulldog. But the extinction of former breeds is apparently aided by another cause; for whenever a breed is kept in scanty numbers, as at present with the bloodhound, it is reared with difficulty, and this apparently is due to the evil effects of long-continued close interbreeding. As several breeds of the dog have been slightly but sensibly modified within so short a period as the last one or two centuries, by the selection of the best individual dogs, modified in many cases by crosses with other breeds; and as we shall hereafter see that the breeding of dogs was attended to in ancient times, as it still is by savages, we may conclude that we have in selection, even if only occasionally practised, a potent means of modification.

Domestic Cats.

Cats have been domesticated in the East from an ancient period; Mr. Blyth informs me that they are mentioned in a Sanskrit writing 2000 years old, and in Egypt their antiquity is known to be even greater, as shown by monumental drawings and their mummied bodies. These mummies, according to De Blainville^[88] who has particularly studied the subject, belong to no less than three species, namely, F. caligulata, bubastes, and chaus. The two former species are said to be still found, both wild and domesticated, in parts of Egypt. F. caligulata presents a difference in the first inferior milk molar tooth, as compared with the domestic cats of Europe, which makes De Blainville conclude that it is not one of the parent-forms of our cats. Several naturalists, as Pallas, Temminck, Blyth, believe that domestic cats are the descendants of several species {44}commingled: it is certain that cats cross readily with various wild species, and it would appear that the character of the domestic breeds has, at least in some cases, been thus affected. Sir W. Jardine has no doubt that, "in the north of Scotland, there has been occasional crossing with our native species (F. sylvestris), and that the result of these crosses has been kept in our houses. I have seen," he adds, "many cats very closely resembling the wild cat, and one or two that could scarcely be distinguished from it." Mr. Blyth^[89] remarks on this passage, "but such cats are never seen in the southern parts of England; still, as compared with any Indian tame cat, the affinity of the ordinary British cat to F. sylvestris is manifest; and due I suspect to frequent intermixture at a time when the tame cat was first introduced into Britain and continued rare, while the wild species was far more abundant than at present." In Hungary, Jeitteles^[90] was assured on trustworthy authority that a wild male cat crossed with a female domestic cat, and that the hybrids long lived in a domesticated state. In Algiers the domestic cat has crossed with the wild cat (F. Lybica) of that country.^[91] In South Africa, as Mr. E. Layard informs me, the domestic cat intermingles freely with the wild F. caffra; he has seen a pair of hybrids which were quite tame and particularly attached to the lady who brought them up; and Mr. Fry has found that these hybrids are fertile. In India the domestic cat, according to Mr. Blyth, has crossed with four Indian species. With respect to one of these species, F. chaus, an excellent observer, Sir W. Elliot, informs me that he once killed, near Madras, a wild brood, which were evidently hybrids from the domestic cat; these young animals had a thick lynx-like tail and the broad brown bar on the inside of the forearm characteristic of F. chaus. Sir W. Elliot adds that he has often observed this same mark on the forearms of domestic cats in India. Mr. Blyth states that domestic cats coloured nearly like F. chaus, but not resembling that species in shape, abound in {45}Bengal; he adds, "such a colouration is utterly unknown in European cats, and the proper tabby markings (pale streaks on a black ground, peculiarly and symmetrically disposed), so common in English cats, are never seen in those of India." Dr. D. Short has assured Mr. Blyth^[92] that at Hansi hybrids between the common cat and F. ornata (or torquata) occur, "and that many of the domestic cats of that part of India were undistinguishable from the wild F. ornata." Azara states, but only on the authority of the inhabitants, that in Paraguay the cat has crossed with two native species. From these several cases we see that in Europe, Asia, Africa, and America, the common cat, which lives a freer life than most other domesticated animals, has crossed with various wild species; and that in some instances the crossing has been sufficiently frequent to affect the character of the breed.

Whether domestic cats have descended from several distinct species, or have only been modified by occasional crosses, their fertility, as far as is known, is unimpaired. The large Angora or Persian cat is the most distinct in structure and habits of all the domestic breeds; and is believed by Pallas, but on no distinct evidence, to be descended from the F. manul of middle Asia; but I am assured by Mr. Blyth that this cat breeds freely with Indian cats, which, as we have already seen, have apparently been much crossed with F. chaus. In England half-bred Angora cats are perfectly fertile with the common cat; I do not know whether the half-breeds are fertile one with another; but as they are common in some parts of Europe, any marked degree of sterility could hardly fail to have been noticed.

Within the same country we do not meet with distinct races of the cat, as we do of dogs and of most other domestic animals; though the cats of the same country present a considerable amount of fluctuating variability. The explanation obviously is that, from their nocturnal and rambling habits, indiscriminate crossing cannot without much trouble be prevented. Selection cannot be brought into play to produce distinct breeds, or to keep those distinct which have been imported from foreign lands. On the other hand, in islands and {46}in countries completely separated from each other, we meet with breeds more or less distinct; and these cases are worth giving as showing that the scarcity of distinct races in the same country is not caused by a deficiency of variability in the animal. The tail-less cats of the Isle of Man are said to differ from common cats not only in the want of a tail, but in the greater length of their hind legs, in the size of their heads, and in habits. The Creole cat of Antigua, as I am informed by Mr. Nicholson, is smaller, and has a more elongated head, than the British cat. In Ceylon, as Mr. Thwaites writes to me, every one at first notices the different appearance of the native cat from the English animal; it is of small size, with closely lying hairs; its head is small, with a receding forehead; but the ears are large and sharp; altogether it has what is there called a "low-caste" appearance. Rengger^[93] says that the domestic cat, which has been bred for 300 years in Paraguay, presents a striking difference from the European cat; it is smaller by a fourth, has a more lanky body, its hair is short, shining, scanty, and lies close, especially on the tail: he adds that the change has been less at Ascension, the capital of Paraguay, owing to the continual crossing with newly imported cats; and this fact well illustrates the importance of separation. The conditions of life in Paraguay appear not to be highly favourable to the cat, for, though they have run half-wild, they do not become thoroughly feral, like so many other European animals. In another part of South America, according to Roulin,^[94] the introduced cat has lost the habit of uttering its hideous nocturnal howl. The Rev. W. D. Fox purchased a cat in Portsmouth, which he was told came from the coast of Guinea; its skin was black and wrinkled, fur bluish-grey and short, its ears rather bare, legs long, and whole aspect peculiar. This "negro" cat was fertile with common cats. On the opposite coast of Africa, at Mombas, Captain Owen, R.N.,^[95] states that all the cats are covered with short stiff hair instead of fur: he gives a curious account of a cat from Algoa Bay, which had been kept for some time on board and could be identified with certainty; this {47}animal was left for only eight weeks at Mombas, but during that short period it "underwent a complete metamorphosis, having parted with its sandy-coloured fur." A cat from the Cape of Good Hope has been described by Desmarest as remarkable from a red stripe extending along the whole length of its back. Throughout an immense area, namely, the Malayan archipelago, Siam, Pegu, and Burmah, all the cats have truncated tails about half the proper length,^[96] often with a sort of knot at the end. In the Caroline archipelago the cats have very long legs, and are of a reddish-yellow colour.^[97] In China a breed has drooping ears. At Tobolsk, according to Gmelin, there is a red-coloured breed. In Asia, also, we find the well-known Angora or Persian breed.

The domestic cat has run wild in several countries, and everywhere assumes, as far as can be judged by the short recorded descriptions, a uniform character. Near Maldonado, in La Plata, I shot one which seemed perfectly wild; it was carefully examined by Mr. Waterhouse,^[98] who found nothing remarkable in it, excepting its great size. In New Zealand, according to Dieffenbach, the feral cats assume a streaky grey colour like that of wild cats; and this is the case with the half-wild cats of the Scotch Highlands.

We have seen that distant countries possess distinct domestic races of the cat. The differences may be in part due to descent from several aboriginal species, or at least to crosses with them. In some cases, as in Paraguay, Mombas, and Antigua, the differences seem due to the direct action of different conditions of life. In other cases some slight effect may possibly be attributed to natural selection, as cats in many cases have largely to support themselves and to escape diverse dangers. But man, owing to the difficulty of pairing cats, has done nothing by methodical selection; and probably very little by unintentional selection; though in each litter he generally saves the prettiest, {48}and values most a good breed of mouse or rat-catchers. Those cats which have a strong tendency to prowl after game, generally get destroyed by traps. As cats are so much petted, a breed bearing the same relation to other cats, that lapdogs bear to larger dogs, would have been much valued; and if selection could have been applied, we should certainly have had many breeds in each long-civilized country, for there is plenty of variability to work upon.

We see in this country considerable diversity in size, some in the proportions of the body, and extreme variability in colouring. I have only lately attended to this subject, but have already heard of some singular cases of variation; one of a cat born in the West Indies toothless, and remaining so all its life. Mr. Tegetmeier has shown me the skull of a female cat with its canines so much developed that they protruded uncovered beyond the lips; the tooth with the fang being .95, and the part projecting from the gum .6 of an inch in length. I have heard of a family of six-toed cats. The tail varies greatly in length; I have seen a cat which always carried its tail flat on its back when pleased. The ears vary in shape, and certain strains, in England, inherit a pencil-like tuft of hairs, above a quarter of an inch in length, on the tips of their ears; and this same peculiarity, according to Mr. Blyth, characterises some cats in India. The great variability in the length of the tail and the lynx-like tufts of hairs on the ears are apparently analogous to differences in certain wild species of the genus. A much more important difference, according to Daubenton,^[99] is that the intestines of domestic cats are wider, and a third longer, than in wild cats of the same size; and this apparently has been caused by their less strictly carnivorous diet.

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CHAPTER II.

HORSES AND ASSES.

HORSE.—DIFFERENCES IN THE BREEDS—INDIVIDUAL VARIABILITY OF—DIRECT EFFECTS OF THE CONDITIONS OF LIFE—CAN WITHSTAND MUCH COLD—BREEDS MUCH MODIFIED BY SELECTION—COLOURS OF THE HORSE—DAPPLING—DARK STRIPES ON THE SPINE, LEGS, SHOULDERS, AND FOREHEAD—DUN-COLOURED HORSES MOST FREQUENTLY STRIPED—STRIPES PROBABLY DUE TO REVERSION TO THE PRIMITIVE STATE OF THE HORSE.

ASSES.—BREEDS OF—COLOUR OF—LEG- AND SHOULDER- STRIPES—SHOULDER-STRIPES SOMETIMES ABSENT, SOMETIMES FORKED.

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 latter part of the Stone period.^[100] At the present time the number of breeds is great, as may be seen by consulting any treatise on the Horse.^[101] 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 with each separate island in the great Malay archipelago.^[102] 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 six or seven other living species of the genus Equus.

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Of individual variations not known to characterise particular breeds, and not great or injurious enough to be called monstrosities, I have not collected many cases. Mr. G. Brown, of the Cirencester Agricultural College, who has particularly attended to the dentition of our domestic animals, writes to me that he has "several times noticed eight permanent incisors instead of six in the jaw." Male horses alone properly have canines, but they are occasionally found in the mare, though of small size.^[103] The number of ribs is properly eighteen, but Youatt^[104] asserts that not unfrequently there are nineteen on each side, the additional one being always the posterior rib. I have seen several notices of variations in the bones of the leg; thus Mr. Price^[105] speaks of an additional bone in the hock, and of certain abnormal appearances between the tibia and astragalus, as quite common in Irish horses, and not due to disease. Horses have often been observed, according to M. Gaudry,^[106] 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.^[107] Azara has described two cases in South America in which the projections were between three and four inches in length: other instances have occurred in Spain.

That there has been much inherited variation in the horse cannot be doubted, when we reflect on the number of the breeds existing throughout the world or even within the same country, and when we know that they have largely increased in number {51}since the earliest known records.^[108] Even in so fleeting a character as colour, Hofacker^[109] found that, out of two hundred and sixteen cases in which horses of the same colour were paired, only eleven pairs produced foals of a quite different colour. As Professor Low^[110] has remarked, the English race-horse offers the best possible evidence of inheritance. The pedigree of a race-horse is of more value in judging of its probable success than its appearance: "King Herod" gained in prizes 201,505l. sterling, and begot 497 winners; "Eclipse" begot 334 winners.

Whether the whole amount of difference between the various breeds be due to variation is doubtful. From the fertility of the most distinct breeds^[111] when crossed, naturalists have generally looked at all the breeds as having descended from a single species. Few will agree with Colonel H. Smith, who believes that they have descended from no less than five primitive and differently coloured stocks.^[112] But as several species and varieties of the horse existed^[113] during the later tertiary periods, and as Rütimeyer found differences in the size and form of the skull in the earliest known domesticated horses,^[114] we ought not to feel sure that all our breeds have descended from a single species. As we see that the savages of North and South America easily reclaim the feral horses, there is no improbability in savages in various quarters of the world having domesticated more than one native species or natural race. No aboriginal or truly wild horse is positively known now to exist; for it is thought by some authors that the wild horses of the East are escaped domestic animals.^[115] If our domestic breeds have descended from several {52}species or natural races, these apparently have all become extinct in the wild state. With our present knowledge, the common view that all have descended from a single species is, perhaps, the most probable.

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 Chile, which have lived under nearly the same conditions as their progenitors in Andalusia, remain unaltered, whilst 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,^[116] 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 unfavourable 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. Further south, in the Falkland Islands, the offspring of the horses imported in 1764 have already so much deteriorated in size^[117] 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. The horse can withstand intense cold, for wild troops live on the plains of Siberia under lat. 56°,^[118] and aboriginally the horse must {53}have inhabited countries annually covered with snow, for he long retains the instinct of scraping it away to get at the herbage beneath. The wild tarpans in the East have this instinct; and, as I am informed by Admiral Sulivan, this is likewise the case with the horses which have run wild on the Falkland Islands; now this is the more remarkable as the progenitors of these horses could not have followed this instinct during many generations in La Plata: the wild cattle of the Falklands never scrape away the snow, and perish when the ground is long covered. In the northern parts of America the horses, descended from those introduced by the Spanish conquerors of Mexico, have the same habit, as have the native bisons, but not so the cattle introduced from Europe.^[119]

The horse can flourish under intense heat as well as under intense cold, for he is known to come to the highest perfection, though not attaining a large size, in Arabia and northern Africa. Much humidity is apparently more injurious to the horse than heat or cold. In the Falkland Islands, horses suffer much from the dampness; and this same circumstance may perhaps partly account for the singular fact that to the eastward of the Bay of Bengal,^[120] over an enormous and humid area, in Ava, Pegu, Siam, the Malayan archipelago, the Loo Choo Islands, and a large part of China, no full-sized horse is found. When we advance as far eastward as Japan, the horse reacquires his full size.^[121]

With most of our domesticated animals, some breeds are kept on account of their curiosity or beauty; but the horse is valued almost solely for its utility. Hence semi-monstrous breeds are not preserved; and probably all the existing breeds have been slowly formed either by the direct action of the conditions of life, or through the selection of individual differences. No doubt semi-monstrous breeds might have been formed: thus Mr. Waterton records^[122] the case of a mare which produced {54}successively three foals without tails; so that a tailless race might have been formed like the tailless races of dogs and cats. A Russian breed of horses is said to have frizzled hair, and Azara^[123] relates that in Paraguay horses are occasionally born, but are generally destroyed, with hair like that on the head of a negro; and this peculiarity is transmitted even to half-breeds: it is a curious case of correlation that such horses have short manes and tails, and their hoofs are of a peculiar shape like those of a mule.

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 have proceeded from the commingled blood of Arabs, Turks, and Barbs; but selection and training have together made him a very different animal from his parent-stocks. As a writer in India, who evidently knows the pure Arab well, asks, who now, "looking at our present breed of race-horses, could have conceived that they were the result of the union of the Arab horse and African mare?" The improvement is so marked that in running for the Goodwood Cup "the first descendants of Arabian, Turkish, and Persian horses, are allowed a discount of 18 lbs. weight; and when both parents are of these countries a discount of 36 lbs."^[124] It is notorious that the Arabs have long been as careful about the pedigree of their horses as we are, and this implies great and continued care in breeding. Seeing what has been done in England by careful breeding, can we doubt that the Arabs must likewise have produced during the course of centuries a marked effect on the qualities of their horses? But we may go much farther back in time, for in the most ancient known book, the Bible, we hear of studs carefully kept for breeding, {55}and of horses imported at high prices from various countries.^[125] We may therefore conclude that, whether or not the various existing breeds of the horse have proceeded from one or more aboriginal stocks, yet that a great amount of change has resulted from the direct action of the conditions of life, and probably a still greater amount from the long-continued selection by man of slight individual differences.

With several domesticated quadrupeds and birds, certain coloured marks are either strongly inherited or tend to reappear after having long been lost. As this subject will hereafter be seen to be of importance, I will give a full account of the colouring of horses. All English breeds, however unlike in size and appearance, and several of those in India and the Malay archipelago, present a similar range and diversity of colour. The English race-horse, however, is said^[126] never to be dun-coloured; but as dun and cream-coloured horses are considered by the Arabs as worthless, "and fit only for Jews to ride,"^[127] these tints may have been removed by long-continued selection. Horses of every colour, and of such widely different kinds as dray-horses, cobs, and ponies, are all occasionally dappled,^[128] in the same manner as is so conspicuous with grey horses. This fact does not throw any clear light on the colouring of the aboriginal horse, but is a case of analogous variation, for even asses are sometimes dappled, and I have seen, in the British Museum, a hybrid from the ass and zebra dappled on its hinder quarters. By the expression analogous variation (and it is one that I shall often have occasion to use) I mean a variation occurring in a species or variety which resembles a normal character in another and distinct species or variety. Analogous variations may arise, as will be explained in a future chapter, {56}from two or more forms with a similar constitution having been exposed to similar conditions,—or from one of two forms having reacquired through reversion a character inherited by the other form from their common progenitor,—or from both forms having reverted to the same ancestral character. We shall immediately see that horses occasionally exhibit a tendency to become striped over a large part of their bodies; and as we know that stripes readily pass into spots and cloudy marks in the varieties of the domestic cat and in several feline species—even the cubs of the uniformly-coloured lion being spotted with dark marks on a lighter ground—we may suspect that the dappling of the horse, which has been noticed by some authors with surprise, is a modification or vestige of a tendency to become striped.

Fig. 1.—Dun Devonshire Pony, with shoulder, spinal, and leg stripes.

This tendency in the horse to become striped is in several respects an interesting feet. Horses of all colours, of the most diverse breeds, in various parts of the world, often have a dark stripe extending along the spine, from the mane to the tail; but this is so common that I need enter into no particulars.^[129] Occasionally horses are transversely barred on the legs, chiefly on the under side; and more rarely they have a distinct stripe on the shoulder, like that on the shoulder of the ass, or a broad dark patch representing a stripe. Before entering on any details I must premise that {57}the term dun-coloured is vague, and includes three groups of colour, viz. that between cream-colour and reddish-brown, which graduates into light-bay or light-chesnut—this, I believe, is often called fallow-dun; secondly, leaden or slate-colour or mouse-dun, which graduates into an ash-colour; and, lastly, dark-dun, between brown and black. In England I have examined a rather large, lightly-built, fallow-dun Devonshire pony (fig. 1), with a conspicuous stripe along the back, with light transverse stripes on the under sides of its front legs, and with four parallel stripes on each shoulder. Of these four stripes the posterior one was very minute and faint; the anterior one, on the other hand, was long and broad, but interrupted in the middle, and truncated at its lower extremity, with the anterior angle produced into a long tapering point. I mention this latter fact because the shoulder-stripe of the ass occasionally presents exactly the same appearance. I have had an outline and description sent to me of a small, purely-bred, light fallow-dun Welch pony, with a spinal stripe, a single transverse stripe on each leg, and three shoulder-stripes; the posterior stripe corresponding with that on the shoulder of the ass was the longest, whilst the two anterior parallel stripes, arising from the mane, decreased in length, in a reversed manner as compared with the shoulder-stripes on the above-described Devonshire pony. I have seen a bright fallow-dun, strong cob, with its front legs transversely barred on the under sides in the most conspicuous manner; also a dark-leaden mouse-coloured pony with similar leg stripes, but much less conspicuous; also a bright fallow-dun colt, fully three-parts thoroughbred, with very plain transverse stripes on the legs; also a chesnut-dun cart-horse with a conspicuous spinal stripe, with distinct traces of shoulder-stripes, but none on the legs; I could add other cases. My son made a sketch for me of a large, heavy, Belgian cart-horse, of a fallow-dun, with a conspicuous spinal stripe, traces of leg-stripes, and with two parallel (three inches apart) stripes about seven or eight inches in length on both shoulders. I have seen another rather light cart-horse, of a dirty dark cream-colour, with striped legs, and on one shoulder a large ill-defined dark cloudy patch, and on the opposite shoulder two parallel faint stripes. All the cases yet mentioned are duns of various tints; but Mr. W. W. Edwards has seen a nearly thoroughbred chesnut horse which had the spinal stripe, and distinct bars on the legs; and I have seen two bay carriage-horses with black spinal stripes; one of these horses had on each shoulder a light shoulder-stripe, and the other had a broad black ill-defined stripe, running obliquely half-way down each shoulder; neither had leg-stripes.

The most interesting case which I have met with occurred in a colt of my own breeding. A bay mare (descended from a dark-brown Flemish mare by a light grey Turcoman horse) was put to Hercules, a thoroughbred dark bay, whose sire (Kingston) and dam were both bays. The colt ultimately turned out brown; but when only a fortnight old it was a dirty bay, shaded with mouse-grey, and in parts with a yellowish tint: it had only a trace of the spinal stripe, with a few obscure transverse bars on the legs; but almost the whole body was marked with very narrow dark stripes, in most parts so obscure as to be visible only in certain lights, like the {58}stripes which may be seen on black kittens. These stripes were distinct on the hind-quarters, where they diverged from the spine, and pointed a little forwards; many of them as they diverged from the spine became a little branched, exactly in the same manner as in some zebrine species. The stripes were plainest on the forehead between the ears, where they formed a set of pointed arches, one under the other, decreasing in size downwards towards the muzzle; exactly similar marks may be seen on the forehead of the quagga and Burchell's zebra. When this foal was two or three months old all the stripes entirely disappeared. I have seen similar marks on the forehead of a fully grown, fallow-dun, cob-like horse, having a conspicuous spinal stripe, and with its front legs well barred.

In Norway the colour of the native horse or pony is dun, varying from almost cream-colour to dark mouse-dun; and an animal is not considered purely bred unless it has the spinal and leg stripes.^[130] In one part of the country my son estimated that about a third of the ponies had striped legs; he counted seven stripes on the fore-legs and two on the hind-legs of one pony; only a few of them exhibited traces of shoulder-stripes; but I have heard of a cob imported from Norway which had the shoulder as well as the other stripes well developed. Colonel Ham. Smith^[131] alludes to dun-horses with the spinal stripe in the Sierras of Spain; and the horses originally derived from Spain, in some parts of South America, are now duns. Sir W. Elliot informs me that he inspected a herd of 300 South American horses imported into Madras, and many of these had transverse stripes on the legs and short shoulder-stripes; the most strongly marked individual, of which a coloured drawing was sent me, was a mouse-dun, with the shoulder-stripes slightly forked.

In the North-Western parts of India striped horses of more than one breed are apparently commoner than in any other part of the world; and I have received information respecting them from several officers, especially from Colonel Poole, Colonel Curtis, Major Campbell, Brigadier St. John, and others. The Kattywar horses are often fifteen or sixteen hands in height, and are well but lightly built. They are of all colours, but the several kinds of duns prevail; and these are so generally striped, that a horse without stripes is not considered pure. Colonel Poole believes that all the duns have the spinal stripe, the leg-stripes are generally present, and he thinks that about half the horses have the shoulder-stripe; this stripe is sometimes double or treble on both shoulders. Colonel Poole has often seen stripes on the cheeks and sides of the nose. He has seen stripes on the grey and bay Kattywars when first foaled, but they soon faded away. I have received other accounts of cream-coloured, bay, brown, and grey Kattywar horses being striped. Eastward of India, the Shan (north of Burmah) ponies, as I am informed by Mr. Blyth, have spinal, leg, and shoulder stripes. Sir W. Elliot informs me that he saw two bay Pegu ponies with {59}leg-stripes. Burmese and Javanese ponies are frequently dun-coloured, and have the three kinds of stripes, "in the same degree as in England."^[132] Mr. Swinhoe informs me that he examined two light-dun ponies of two Chinese breeds, viz. those of Shangai and Amoy; both had the spinal stripe, and the latter an indistinct shoulder-stripe.

We thus see that in all parts of the world breeds of the horse as different as possible, when of a dun-colour (including under this term a wide range of tint from cream to dusky black), and rarely when of bay, grey, and chesnut shades, have the several above-specified stripes. Horses which are of a yellow colour with white mane and tail, and which are sometimes called duns, I have never seen with stripes.^[133]

From reasons which will be apparent in the chapter on Reversion, I have endeavoured, but with poor success, to discover whether duns, which are so much oftener striped than other coloured horses, are ever produced from the crossing of two horses, neither of which are duns. Most persons to whom I have applied believe that one parent must be a dun; and it is generally asserted, that, when this is the case, the dun-colour and the stripes are strongly inherited.^[134] One case has fallen under my own observation of a foal from a black mare by a bay horse, which when fully grown was a dark fallow-dun and had a narrow but plain spinal stripe. Hofacker^[135] gives two instances of mouse-duns (Mausrapp) being produced from two parents of different colours and neither duns.

I have also endeavoured with little success to find out whether the stripes are generally plainer or less plain in the foal than in the adult horse. Colonel Poole informs me that, as he believes, "the stripes are plainest when the colt is first foaled; they then become less and less distinct till after the first coat is shed, when they come out as strongly as before; but certainly often fade away as the age of the horse increases." Two other accounts confirm this fading of the stripes in old horses in India. One writer, on the other hand, states that colts are often born without stripes, but that they appear as the colt grows older. Three authorities affirm that in Norway the stripes are less plain in the foal than in the adult. Perhaps there is no fixed rule. In the case described by me of the young foal which was narrowly striped over nearly all its body, there was no doubt about the the early and complete disappearance of the stripes. Mr. W. W. Edwards examined for me twenty-two foals of race-horses, and twelve had the spinal stripe more or less plain; this fact, and some other accounts which I have received, lead me to believe that the spinal stripe often disappears in the English race-horse when old. On the whole I infer that the stripes are generally plainest in the foal, and tend to disappear in old age.

The stripes are variable in colour, but are always darker than the rest of the body. They do not by any means always {60}coexist on the different parts of the body: the legs may be striped without any shoulder-stripe, or the converse case, which is rarer, may occur; but I have never heard of either shoulder or leg-stripes without the spinal stripe. The latter is by far the commonest of all the stripes, as might have been expected, as it characterises the other seven or eight species of the genus. It is remarkable that so trifling a character as the shoulder-stripe being double or triple should occur in such different breeds as Welch and Devonshire ponies, the Shan pony, heavy cart-horses, light South American horses, and the lanky Kattywar breed. Colonel Hamilton Smith believes that one of his five supposed primitive stocks was dun-coloured and striped; and that the stripes in all the other breeds result from ancient crosses with this one primitive dun; but it is extremely improbable that different breeds living in such distant quarters of the world should all have been crossed with any one aboriginally distinct stock. Nor have we any reason to believe that the effects of a cross at a very remote period could be propagated for so many generations as is implied on this view.

With respect to the primitive colour of the horse having been dun, Colonel Hamilton Smith^[136] has collected a large body of evidence showing that this tint was common in the East as far back as the time of Alexander, and that the wild horses of Western Asia and Eastern Europe now are, or recently were, of various shades of dun. It seems that not very long ago a wild breed of dun-coloured horses with a spinal stripe was preserved in the royal parks in Prussia. I hear from Hungary that the inhabitants of that country look at the duns with a spinal stripe as the aboriginal stock, and so it is in Norway. Dun-coloured ponies are not rare in the mountainous parts of Devonshire, Wales, and Scotland, where the aboriginal breed would have had the best chance of being preserved. In South America in the time of Azara, when the horse had been feral for about 250 years, 90 out of 100 horses were "bai-châtains," and the remaining ten were "zains," and not more than one in 2000 {61}black. Zain is generally translated as dark without any white; but as Azara speaks of mules being "zain-clair," I suspect that zain must have meant dun-coloured. In some parts of the world feral horses show a strong tendency to become roans.^[137]

In the following chapters on the Pigeon we shall see that in pure breeds of various colours, when a blue bird is occasionally produced, certain black marks invariably appear on the wings and tail; so again, when variously coloured breeds are crossed, blue birds with the same black marks are frequently produced. We shall further see that these facts are explained by, and afford strong evidence in favour of, the view that all the breeds are descended from the rock-pigeon, or Columba livia, which is thus coloured and marked. But the appearance of the stripes on the various breeds of the horse, when of a dun-colour, does not afford nearly such good evidence of their descent from a single primitive stock as in the case of the pigeon; because no certainly wild horse is known as a standard of comparison; because the stripes when they do appear are variable in character; because there is far from sufficient evidence of the appearance of the stripes from the crossing of distinct breeds; and lastly, because all the species of the genus Equus have the spinal stripe, and several have shoulder and leg stripes. Nevertheless the similarity in the most distinct breeds in their general range of colour, in their dappling, and in the occasional appearance, especially in duns, of leg-stripes and of double or triple shoulder-stripes, taken together, indicate the probability of the descent of all the existing races from a single, dun-coloured, more or less striped, primitive stock, to which our horses still occasionally revert.

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The Ass.

Four species of Asses, besides three of zebras, have been described by naturalists; but there can now be little doubt that our domesticated animal is descended from one alone, namely, the Asinus tæniopus of Abyssinia.^[138] The ass is sometimes advanced as an instance of an animal domesticated, as we know by the Old Testament, from an ancient period, which has varied only in a very slight degree. But this is by no means strictly true; for in Syria alone there are four breeds;^[139] first, a light and graceful animal, with an agreeable gait, used by ladies; secondly, an Arab breed reserved exclusively for the saddle; thirdly, a stouter animal used for ploughing and various purposes; and lastly, the large Damascus breed, with a peculiarly long body and ears. In this country, and generally in Central Europe, though the ass is by no means uniform in appearance, it has not given rise to distinct breeds like those of the horse. This may probably be accounted for by the animal being kept chiefly by poor persons, who do not rear large numbers, nor carefully match and select the young. For, as we shall see in a future chapter, the ass can with ease be greatly improved in size and strength by careful selection, combined no doubt with good food; and we may infer that all its other characters would be equally amenable to selection. The small size of the ass in England and Northern Europe is apparently due far more to want of care in breeding than to cold; for in Western India, where the ass is used as a beast of burden by some of the lower castes, it is not much larger than a Newfoundland dog, "being generally not more than from twenty to thirty inches high."^[140]

The ass varies greatly in colour; and its legs, especially the fore-legs, both in England and other countries—for instance, in China—are occasionally barred transversely more plainly than those of dun-coloured horses. With the horse the occasional appearance of leg-stripes was accounted for, through the principle of reversion, by the supposition that the primitive horse was {63}thus striped; with the ass we may confidently advance this explanation, for the parent-form, the A. tæniopus, is known to be barred, though only in a slight degree, across the legs. The stripes are believed to occur most frequently and to be plainest on the legs of the domestic ass during early youth,^[141] as is apparently likewise the case with the horse. The shoulder-stripe, which is so eminently characteristic of the species, is nevertheless variable in breadth, length, and manner of termination. I have measured a shoulder-stripe four times as broad as another; and some more than twice as long as others. In one light-grey ass the shoulder-stripe was only six inches in length, and as thin as a piece of string; and in another animal of the same colour there was only a dusky shade representing a stripe. I have heard of three white asses, not albinoes, with no trace of shoulder or spinal stripes;^[142] and I have seen nine other asses with no shoulder-stripe, and some of them had no spinal stripe. Three of the nine were light-greys, one a dark-grey, another grey passing into reddish-roan, and the others were brown, two being tinted on parts of their bodies with a reddish or bay shade. Hence we may conclude that, if grey and reddish-brown asses had been steadily selected and bred from, the shoulder-stripe would have been almost as generally and as completely lost as in the case of the horse.

The shoulder-stripe on the ass is sometimes double, and Mr. Blyth has seen even three or four parallel stripes.^[143] I have observed in ten cases shoulder-stripes abruptly truncated at the lower end, with the anterior angle produced into a tapering point, precisely as has been figured in the dun Devonshire pony. I have seen three cases of the terminal portion abruptly and angularly bent; and two cases of a distinct though slight forking. In Syria, Dr. Hooker and his party observed for me no less than five instances of the shoulder-stripe being plainly forked over the fore leg. In the common mule it is likewise sometimes forked. When I first noticed the forking and angular bending of the shoulder-stripe, I had seen enough of the stripes {64}in the various equine species to feel convinced that even a character so unimportant as this had a distinct meaning, and was thus led to attend to the subject. I now find that in the Asinus Burchellii and quagga, the stripe which corresponds with the shoulder-stripe of the ass, as well as some of the stripes on the neck, bifurcate, and that some of those near the shoulder have their extremities angularly bent backwards. The forking and angular bending of the stripes on the shoulders apparently stand in relation with the changed direction of the nearly upright stripes on the sides of the body and neck to the transverse bars on the legs. Finally we see that the presence of shoulder, leg, and spinal stripes in the horse,—their occasional absence in the ass,—the occurrence of double and triple shoulder-stripes in both animals, and the similar manner in which these stripes terminate at their lower extremities,—are all cases of analogous variation in the horse and ass. These cases are probably not due to similar conditions acting on similar constitutions, but to a partial reversion in colour to the common progenitor of these two species, as well as of the other species of the genus. We shall hereafter have to return to this subject, and discuss it more fully.

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CHAPTER III.

PIGS—CATTLE—SHEEP—GOATS.

PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICA—TORF-SCHWEIN—JAPAN PIG—FERTILITY OF CROSSED PIGS—CHANGES IN THE SKULL OF THE HIGHLY CULTIVATED RACES—CONVERGENCE OF CHARACTER—GESTATION—SOLID-HOOFED SWINE—CURIOUS APPENDAGES TO THE JAWS—DECREASE IN SIZE OF THE TUSKS—YOUNG PIGS LONGITUDINALLY STRIPED—FERAL PIGS—CROSSED BREEDS.

CATTLE.—ZEBU A DISTINCT SPECIES—EUROPEAN CATTLE PROBABLY DESCENDED FROM THREE WILD FORMS—ALL THE RACES NOW FERTILE TOGETHER—BRITISH PARK CATTLE—ON THE COLOUR OF THE ABORIGINAL SPECIES—CONSTITUTIONAL DIFFERENCES—SOUTH AFRICAN RACES—SOUTH AMERICAN RACES—NIATA CATTLE—ORIGIN OF THE VARIOUS RACES OF CATTLE.

SHEEP.—REMARKABLE RACES OF—VARIATIONS ATTACHED TO THE MALE SEX—ADAPTATIONS TO VARIOUS CONDITIONS—GESTATION OF—CHANGES IN THE WOOL—SEMI-MONSTROUS BREEDS.

GOATS.—REMARKABLE VARIATIONS OF.

The breeds of the pig have recently been more closely studied, though much still remains to be done, than those of almost any other domesticated animal. This has been effected by Hermann von Nathusius in two admirable works, especially in the later one on the Skulls of the several races, and by Rütimeyer in his celebrated Fauna of the ancient Swiss lake-dwellings.^[144] Nathusius has shown that all the known breeds may be divided in two great groups: one resembling in all important respects and no doubt descended from the common wild boar; so that this may be called the Sus scrofa group. The other group differs in several important and constant osteological characters; its wild parent-form is unknown; the name given to it by Nathusius, according to the law of priority, is Sus Indica of Pallas. This name must now be followed, though an unfortunate one, as the wild aboriginal does not inhabit India, and the best-known domesticated breeds have been imported from Siam and China.

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Firstly, the Sus scrofa breeds, or those resembling the common wild boar. These still exist, according to Nathusius (Schweineschädel, s. 75), in various parts of central and northern Europe; formerly every kingdom,^[145] and almost every province in Britain, possessed its own native breed; but these are now everywhere rapidly disappearing, being replaced by improved breeds crossed with the S. Indica form. The skull in the breeds of the S. scrofa type resembles, in all important respects, that of the European wild boar; but it has become (Schweineschädel, s. 63-68) higher and broader relatively to its length; and the hinder part is more upright. The differences, however, are all variable in degree. The breeds which thus resemble S. scrofa in their essential skull-characters differ conspicuously from each other in other respects, as in the length of the ears and legs, curvature of the ribs, colour, hairiness, size and proportions of the body.

The wild Sus scrofa has a wide range, namely, Europe, North Africa, as identified by osteological characters by Rütimeyer, and Hindostan, as similarly identified by Nathusius. But the wild boars inhabiting these several countries differ so much from each other in external characters, that they have been ranked by some naturalists as specifically distinct. Even within Hindostan these animals, according to Mr. Blyth, form very distinct races in the different districts; in the N. Western provinces, as I am informed by the Rev. R. Everest, the boar never exceeds 36 inches in height, whilst in Bengal one has been measured 44 inches in height. In Europe, Northern Africa, and Hindostan, domestic pigs have been known to cross with the wild native species;^[146] and in Hindostan an accurate observer,^[147] Sir Walter Elliot, after describing the differences between wild Indian and wild German boars, remarks that "the same differences are perceptible in the domesticated {67}individuals of the two countries." We may therefore conclude that the breeds of the Sus scrofa type have either descended from, or been modified by crossing with, forms which may be ranked as geographical races, but which are, according to some naturalists, distinct species.

Pigs of the Sus Indica type are best known to Englishmen under the form of the Chinese breed. The skull of S. Indica, as described by Nathusius, differs from that of S. scrofa in several minor respects, as in its greater breadth and in some details in the teeth; but chiefly in the shortness of the lachrymal bones, in the greater width of the fore part of the palate-bones, and in the divergence of the premolar teeth. It deserves especial notice that these latter characters are not gained, even in the least degree, by the domesticated forms of S. scrofa. After reading the remarks and descriptions given by Nathusius, it seems to me to be merely playing with words to doubt whether S. Indica ought to be ranked as a species; for the above-specified differences are more strongly marked than any that can be pointed out between, for instance, the fox and the wolf, or the ass and the horse. As already stated, S. Indica is not known in a wild state; but its domesticated forms, according to Nathusius, come near to S. vittatus of Java and some allied species. A pig found wild in the Aru islands (Schweineschädel, s. 169) is apparently identical with S. Indica; but it is doubtful whether this is a truly native animal. The domesticated breeds of China, Cochin-China, and Siam belong to this type. The Roman or Neapolitan breed, the Andalusian, the Hungarian, and the "Krause" swine of Nathusius, inhabiting south-eastern Europe and Turkey, and having fine curly hair, and the small Swiss "Bündtnerschwein" of Rütimeyer, all agree in their more important skull characters with S. Indica, and, as is supposed, have all been largely crossed with this form. Pigs of this type have existed during a long period on the shores of the Mediterranean, for a figure (Schweineschädel, s. 142) closely resembling the existing Neapolitan pig has been found in the buried city of Herculaneum.

Rütimeyer has made the remarkable discovery that there lived contemporaneously in Switzerland, during the later Stone or Neolithic period, two domesticated forms, the S. scrofa, and {68}the S. scrofa palustris or Torfschwein. Rütimeyer perceived that the latter approached the Eastern breeds, and, according to Nathusius, it certainly belongs to the S. Indica group; but Rütimeyer has subsequently shown that it differs in some well-marked characters. This author was formerly convinced that his Torfschwein existed as a wild animal during the first part of the Stone period, and was domesticated during a later part of the same period.^[148] Nathusius, whilst he fully admits the curious fact first observed by Rütimeyer, that the bones of domesticated and wild animals can be distinguished by their different aspect, yet, from special difficulties in the case of the bones of the pig (Schweineschädel, s. 147), is not convinced of the truth of this conclusion; and Rütimeyer himself seems now to feel some doubt. As the Torfschwein was domesticated at so early a period, and as its remains have been found in several parts of Europe, belonging to various historic and prehistoric ages,^[149] and as closely allied forms still exist in Hungary and on the shores of the Mediterranean, one is led to suspect that the wild S. Indica formerly ranged from Europe to China, in the same manner as S. scrofa now ranges from Europe to Hindostan. Or, as Rütimeyer apparently suspects, a third allied species may formerly have lived in Europe and Eastern Asia.

Several breeds, differing in the proportions of the body, in the length of the ears, in the nature of the hair, in colour, &c., come under the S. Indica type. Nor is this surprising, considering how ancient the domestication of this form has been both in Europe and in China. In this latter country the date is believed by an eminent Chinese scholar^[150] to go back at least 4900 years from the present time. This same scholar alludes to the existence of many local varieties of the pig in China; and at the present time the Chinese take extraordinary pains in feeding and tending their pigs, not even allowing them to walk from place to place.^[151] Hence the Chinese breed, as Nathusius has remarked,^[152] displays in an eminent degree the characters of a highly-cultivated race, and hence, no doubt, its {69}high value in the improvement of our European breeds. Nathusius makes a remarkable statement (Schweineschädel, s. 138), that the infusion of the 1/32nd, or even of the 1/64th, part of the blood of S. Indica into a breed of S. scrofa, is sufficient plainly to modify the skull of the latter species. This singular fact may perhaps be accounted for by several of the chief distinctive characters of S. Indica, such as the shortness of the lachrymal bones, &c., being common to several of the species of the genus; for in crosses the characters which are common to many species apparently tend to be prepotent over those appertaining to only a few species.

The Japan pig (S. pliciceps of Gray), which has been recently exhibited in the Zoological Gardens, has an extraordinary appearance from its short head, broad forehead and nose, great fleshy ears, and deeply furrowed skin. The following woodcut is copied from that given by Mr. Bartlett.^[153] Not only {70}is the face furrowed, but thick folds of skin, which are harder than the other parts, almost like the plates on the Indian rhinoceros, hang about the shoulders and rump. It is coloured black, with white feet, and breeds true. That it has long been domesticated there can be little doubt; and this might have been inferred even from the fact that its young are not longitudinally striped; for this is a character common to all the species included within the genus Sus and the allied genera whilst in their natural state.^[154] Dr. Gray^[155] has described the skull of this animal, which he ranks not only as a distinct species, but places it in a distinct section of the genus. Nathusius, however, after his careful study of the whole group, states positively (Schweineschädel, s. 153-158) that the skull in all essential characters closely resembles that of the short-eared Chinese breed of the S. Indica type. Hence Nathusius considers the Japan pig as only a domesticated variety of S. Indica: if this really be the case, it is a wonderful instance of the amount of modification which can be effected under domestication.

Fig. 2.—Head of Japan or Masked Pig. (Copied from Mr. Bartlett's paper in Proc. Zoolog. Soc. 1861, p. 263.)

Formerly there existed in the central islands of the Pacific Ocean a singular breed of pigs. These are described by the Rev. D. Tyerman and G. Bennett^[156] as of small size, hump-backed, with a disproportionately long head, with short ears turned backwards, with a bushy tail not more than two inches in length, placed as if it grew from the back. Within half a century after the introduction into these islands of European and Chinese pigs, the native breed, according to the above authors, became almost completely lost by being repeatedly crossed with them. Secluded islands, as might have been expected, seem favourable for the production or retention of peculiar breeds; thus, in the Orkney Islands, the hogs have been described as very small, with erect and sharp ears, and "with an appearance altogether different from the hogs brought from the south."^[157]

Seeing how different the Chinese pigs, belonging to the Sus Indica type, are in their osteological characters and in external {71}appearance from the pigs of the S. scrofa type, so that they must be considered specifically distinct, it is a fact well deserving attention, that Chinese and common pigs have been repeatedly crossed in various manners, with unimpaired fertility. One great breeder who had used pure Chinese pigs assured me that the fertility of the half-breeds inter se and of their recrossed progeny was actually increased; and this is the general belief of agriculturists. Again, the Japan pig or S. pliciceps of Gray is so distinct in appearance from all common pigs, that it stretches one's belief to the utmost to admit that it is simply a domestic variety; yet this breed has been found perfectly fertile with the Berkshire breed; and Mr. Eyton informs me that he paired a half-bred brother and sister and found them quite fertile together.

The modifications of the skull in the most highly cultivated races are wonderful. To appreciate the amount of change, Nathusius' work, with its excellent figures, should be studied. The whole of the exterior of the skull in all its parts has been altered; the hinder surface, instead of sloping backwards, is directed forwards, entailing many changes in other parts; the front of the head is deeply concave; the orbits have a different shape; the auditory meatus has a different direction and shape; the incisors of the upper and lower jaws do not touch each other, and they stand in both jaws above the plane of the molars; the canines of the upper jaw stand in front of those of the lower jaw, and this is a remarkable anomaly: the articular surfaces of the occipital condyles are so greatly changed in shape, that, as Nathusius remarks (s. 133), no naturalist, seeing this important part of the skull by itself, would suppose that it belonged to the genus Sus. These and various other modifications, as Nathusius observes, can hardly be considered as monstrosities, for they are not injurious, and are strictly inherited. The whole head is much shortened; thus, whilst in common breeds its length to that of the body is as 1 to 6, in the "cultur-races" the proportion is as 1 to 9, and even recently as 1 to 11.^[158] The following woodcut^[159] {72}of the head of a wild boar and of a sow from a photograph of the Yorkshire Large Breed, may aid in showing how greatly the head in a highly cultivated race has been modified and shortened.

Fig. 3.—Head of Wild Boar, and of "Golden Days," a pig of the Yorkshire Large Breed; the latter from a photograph. (Copied from Sidney's edit. of 'The Pig,' by Youatt.)

Nathusius has well discussed the causes of the remarkable changes in the skull and shape of the body which the highly cultivated races have undergone. These modifications occur chiefly in the pure and crossed races of the S. Indica type; but their commencement may be clearly detected in the slightly improved breeds of the S. scrofa type.^[160] Nathusius states positively (s. 99, 103), as the result of common experience and of his experiments, that rich and abundant food, given during youth, tends by some direct action to make the head broader and shorter; and that poor food works a contrary result. He lays much stress on the fact that all wild and semi-domesticated pigs, in ploughing up the ground with their muzzles, have; whilst young, to exert the powerful muscles fixed to the hinder part of the head. In highly cultivated races this habit is no longer followed, and consequently the back of the skull becomes modified in shape, entailing other changes in other parts. There can hardly be a doubt that so great a change in habits would {73}affect the skull; but it seems rather doubtful how far this will account for the greatly reduced length of the skull and for its concave front. It is well known (Nathusius himself advancing many cases, s. 104) that there is a strong tendency in many domestic animals—in bull- and pug-dogs, in the niata cattle, in sheep, in Polish fowls, short-faced tumbler pigeons, and in one variety of the carp—for the bones of the face to become greatly shortened. In the case of the dog, as H. Müller has shown, this seems caused by an abnormal state of the primordial cartilage. We may, however, readily admit that abundant and rich food supplied during many generations would give an inherited tendency to increased size of body, and that, from disuse, the limbs would become finer and shorter.^[161] We shall in a future chapter also see that the skull and limbs are apparently in some manner correlated, so that any change in the one tends to affect the other.

Nathusius has remarked, and the observation is an interesting one, that the peculiar form of the skull and body in the most highly cultivated races is not characteristic of any one race, but is common to all when improved up to the same standard. Thus the large-bodied, long-eared, English breeds with a convex back, and the small-bodied, short-eared, Chinese breeds with a concave back, when bred to the same state of perfection, nearly resemble each other in the form of the head and body. This result, it appears, is partly due to similar causes of change acting on the several races, and partly to man breeding the pig for one sole purpose, namely, for the greatest amount of flesh and fat; so that selection has always tended towards one and the same end. With most domestic animals the result of selection has been divergence of character, here it has been convergence.^[162]

The nature of the food supplied during many generations has apparently affected the length of the intestines; for, according to Cuvier,^[163] their length to that of the body in the wild boar is as 9 to 1,—in the common domestic boar as 13.5 to 1,—and in the Siam breed as 16 to 1. In this latter breed the greater {74}length may be due either to descent from a distinct species or to more ancient domestication. The number of mammæ vary, as does the period of gestation. The latest authority says^[164] that "the period averages from 17 to 20 weeks," but I think there must be some error in this statement: in M. Tessier's observations on 25 sows it varied from 109 to 123 days. The Rev. W. D. Fox has given me ten carefully recorded cases with well-bred pigs, in which the period varied from 101 to 116 days. According to Nathusius the period is shortest in the races which come early to maturity; but in these latter the course of development does not appear to be actually shortened, for the young animal is born, judging from the state of the skull, less fully developed, or in a more embryonic condition,^[165] than in the case of common swine, which arrive at maturity at a later age. In the highly cultivated and early matured races, the teeth, also, are developed earlier.

The difference in the number of the vertebræ and ribs in different kinds of pigs, as observed by Mr. Eyton,^[166] and as given in the following table, has often been quoted. The African sow probably belongs to the S. scrofa type; and Mr. Eyton informs me that, since the publication of his paper, cross-bred animals from the African and English races were found by Lord Hill to be perfectly fertile.

{75}

Some semi-monstrous breeds deserve notice. From the time of Aristotle to the present time solid-hoofed swine have occasionally been observed in various parts of the world. Although this peculiarity is strongly inherited, it is hardly probable that all the animals with solid hoofs have descended from the same parents; it is more probable that the same peculiarity has reappeared at various times and places. Dr. Struthers has lately described and figured^[167] the structure of the feet; in both front and hind feet the distal phalanges of the two greater toes are represented by a single, great, hoof-bearing phalanx; and in the front feet, the middle phalanges are represented by a bone which is single towards the lower end, but bears two separate articulations towards the upper end. From other accounts it appears that an intermediate toe is likewise sometimes superadded.

Fig. 4.—Old Irish Pig, with jaw-appendages. (Copied from H. D. Richardson on Pigs.)

Another curious anomaly is offered by the appendages, described by M. Eudes-Deslongchamps as often characterizing the Normandy pigs. These appendages are always attached to the same spot, to the corners of the jaw; they are cylindrical, about three inches in length, covered with bristles, and with a pencil of bristles rising out of a sinus on one side: they have a cartilaginous centre, with two small longitudinal muscles; they occur either symmetrically on both sides of the face or on one {76}side alone. Richardson figures them on the gaunt old "Irish Greyhound pig;" and Nathusius states that they occasionally appear in all the long-eared races, but are not strictly inherited, for they occur or fail in animals of the same litter.^[168] As no wild pigs are known to have analogous appendages, we have at present no reason to suppose that their appearance is due to reversion; and if this be so, we are forced to admit that somewhat complex, though apparently useless, structures may be suddenly developed without the aid of selection. This case perhaps throws some little light on the manner of appearance of the hideous fleshy protuberances, though of an essentially different nature from the above-described appendages, on the cheeks of the wart-hog or Phacochœrus Africanus.

It is a remarkable fact that the boars of all domesticated breeds have much shorter tusks than wild boars. Many facts show that with all animals the state of the hair is much affected by exposure to, or protection from, climate; and as we see that the state of the hair and teeth are correlated in Turkish dogs (other analogous facts will be hereafter given), may we not venture to surmise that the reduction of the tusks in the domestic boar is related to his coat of bristles being diminished from living under shelter? On the other hand, as we shall immediately see, the tusks and bristles reappear with feral boars, which are no longer protected from the weather. It is not surprising that the tusks should be more affected than the other teeth; as parts developed to serve as secondary sexual characters are always liable to much variation.

It is a well-known fact that the young of wild European and Indian pigs,^[169] for the first six months, are longitudinally banded with light-coloured stripes. This character generally disappears under domestication. The Turkish domestic pigs, however, have striped young, as have those of Westphalia, "whatever may be their hue;"^[170] whether these latter pigs belong to the {77}same curly-haired race with the Turkish swine, I do not know. The pigs which have run wild in Jamaica and the semi-feral pigs of New Granada, both those which are black and those which are black with a white band across the stomach, often extending over the back, have resumed this aboriginal character and produce longitudinally-striped young. This is likewise the case, at least occasionally, with the neglected pigs in the Zambesi settlement on the coast of Africa.^[171]

The common belief that all domesticated animals, when they run wild, revert completely to the character of their parent-stock, is chiefly founded, as far as I can discover, on feral pigs. But even in this case the belief is not grounded on sufficient evidence; for the two main types of S. scrofa and Indica have never been distinguished in a feral state. The young, as we have just seen, reacquire their longitudinal stripes, and the boars invariably reassume their tusks. They revert also in the general shape of their bodies, and in the length of their legs and muzzles, to the state of the wild animal, as might have been expected from the amount of exercise which they are compelled to take in search of food. In Jamaica the feral pigs do not acquire the full size of the European wild boar, "never attaining a greater height than 20 inches at the shoulder." In various countries they reassume their original bristly covering, but in different {78}degrees, dependent on the climate; thus, according to Roulin, the semi-feral pigs in the hot valleys of New Granada are very scantily clothed; whereas, on the Paramos, at the height of 7000 to 8000 feet, they acquire a thick covering of wool lying under the bristles, like that on the truly wild pigs of France. These pigs on the Paramos are small and stunted. The wild boar of India is said to have the bristles at the end of its tail arranged like the plumes of an arrow, whilst the European boar has a simple tuft; and it is a curious fact that many, but not all, of the feral pigs in Jamaica, derived from a Spanish stock, have a plumed tail.^[172] With respect to colour, feral pigs generally revert to that of the wild boar; but in certain parts of S. America, as we have seen, some of the semi-feral pigs have a curious white band across their stomachs; and in certain other hot places the pigs are red, and this colour has likewise occasionally been observed in the feral pigs of Jamaica. From these several facts we see that with pigs when feral there is a strong tendency to revert to the wild type; but that this tendency is largely governed by the nature of the climate, amount of exercise, and other causes of change to which they have been subjected.

The last point worth notice is that we have unusually good evidence of breeds of pigs now keeping perfectly true, which have been formed by the crossing of several distinct breeds. The Improved Essex pigs, for instance, breed very true; but there is no doubt that they largely owe their present excellent qualities to crosses originally made by Lord Western with the Neapolitan race, and to subsequent crosses with the Berkshire breed (this also having been improved by Neapolitan crosses), and likewise, probably, with the Sussex breed.^[173] In breeds thus formed by complex crosses, the most careful and unremitting selection during many generations has been found to be indispensable. Chiefly in consequence of so much crossing, some well-known breeds have undergone rapid changes; thus, according to Nathusius,^[174] the Berkshire breed of 1780 is quite {79}different from that of 1810; and, since this latter period, at least two distinct forms have borne the same name.

Cattle.

Domestic cattle are almost certainly the descendants of more than one wild form, in the same manner as has been shown to be the case with our dogs and pigs. Naturalists have generally made two main divisions of cattle: the humped kinds inhabiting tropical countries, called in India Zebus, to which the specific name of Bos Indicus has been given; and the common non-humped cattle, generally included under the name of Bos taurus. The humped cattle were domesticated, as may be seen on the Egyptian monuments, at least as early as the twelfth dynasty, that is 2100 B.C. They differ from common cattle in various osteological characters, even in a greater degree, according to Rütimeyer,^[175] than do the fossil species of Europe, namely Bos primigenius, longifrons, and frontosus, from each other. They differ, also, as Mr. Blyth,^[176] who has particularly attended to this subject, remarks, in general configuration, in the shape of their ears, in the point where the dewlap commences, in the typical curvature of their horns, in their manner of carrying their heads when at rest, in their ordinary variations of colour, especially in the frequent presence of "nilgau-like markings on their feet," and "in the one being born with teeth protruding through the jaws, and the other not so." They have different habits, and their voice is entirely different. The humped cattle in India "seldom seek shade, and never go into the water and there stand knee-deep, like the cattle of Europe." They have run wild in parts of Oude and Rohilcund, and can maintain themselves in a region infested by tigers. They have given rise to many races differing greatly in size, in the presence {80}of one or two humps, in length of horns, and other respects. Mr. Blyth sums up emphatically that the humped and humpless cattle must be considered as distinct species. When we consider the number of points in external structure and habits, independently of their important osteological differences, in which they differ from each other; and that many of these points are not likely to have been affected by domestication, there can hardly be a doubt, notwithstanding the adverse opinion of some naturalists, that the humped and non-humped cattle must be ranked as specifically distinct.

The European breeds of humpless cattle are numerous. Professor Low enumerates 19 British breeds, only a few of which are identical with those on the Continent. Even the small Channel islands of Guernsey, Jersey, and Alderney, possess their own sub-breeds;^[177] and these again differ from the cattle of the other British islands, such as Anglesea, and the western isles of Scotland. Desmarest, who paid attention to the subject, describes 15 French races, excluding sub-varieties and those imported from other countries. In other parts of Europe there are several distinct races, such as the pale-coloured Hungarian cattle, with their light and free step, and their enormous horns sometimes measuring above five feet from tip to tip:^[178] the Podolian cattle are remarkable from the height of their fore-quarters. In the most recent work on Cattle,^[179] engravings are given of fifty-five European breeds; it is, however, probable that several of these differ very little from each other, or are merely synonyms. It must not be supposed that numerous breeds of cattle exist only in long-civilized countries, for we shall presently see that several kinds are kept by the savages of Southern Africa.

With respect to the parentage of the several European breeds, we already know much from Nilsson's Memoir,^[180] and more especially from Rütimeyer's 'Pfahlbauten' and succeeding works. Two or three species or forms of {81}Bos, closely allied to still living domestic races, have been found fossil in the more recent tertiary deposits of Europe. Following Rütimeyer, we have:—

Bos primigenius.—This magnificent, well-known species was domesticated in Switzerland during the Neolithic period; even at this early period it varied a little, having apparently been crossed with other races. Some of the larger races on the Continent, as the Friesland, &c., and the Pembroke race in England, closely resemble in essential structure B. primigenius, and no doubt are its descendants. This is likewise the opinion of Nilsson. Bos primigenius existed as a wild animal in Cæsar's time, and is now semi-wild, though much degenerated in size, in the park of Chillingham; for I am informed by Professor Rütimeyer, to whom Lord Tankerville sent a skull, that the Chillingham cattle are less altered from the true primigenius type than any other known breed.^[181]

Bos trochoceros.—This form is not included in the three species above mentioned, for it is now considered by Rütimeyer to be the female of an early domesticated form of B. primigenius, and as the progenitor of his frontosus race. I may add that specific names have been given to four other fossil oxen, now believed to be identical with B. primigenius.^[182]

Bos longifrons (or brachyceros) of Owen.—This very distinct species was of small size, and had a short body with fine legs. It has been found in England associated with the remains of the elephant and rhinoceros.^[183] It was the commonest form in a domesticated condition in Switzerland during the earliest part of the Neolithic period. It was domesticated in England during the Roman period, and supplied food to the Roman legionaries.^[184] Some remains have been found in Ireland in certain crannoges, of which the dates are believed to be from 843-933 A.D.^[185] Professor Owen^[186] thinks it probable that the Welsh and Highland cattle are descended from this form; as likewise is the case, according to Rütimeyer, with some of the existing Swiss breeds. These latter are of different shades of colour from light-grey to blackish-brown, with a lighter stripe along the spine, but they have no pure white marks. The cattle of North Wales and the Highlands, on the other hand, are generally black or dark-coloured.

Bos frontosus of Nilsson.—This species is allied to B. longifrons, but in the opinion of some good judges is distinct from it. Both co-existed in Scania during the same late geological period,^[187] and both have been found in the Irish crannoges.^[188] Nilsson believes that his B. frontosus may be the {82}parent of the mountain cattle of Norway, which have a high protuberance on the skull between the base of the horns. As Professor Owen believes that the Scotch Highland cattle are descended from his B. longifrons, it is worth notice that a capable judge^[189] has remarked that he saw no cattle in Norway like the Highland breed, but that they more nearly resembled the Devonshire breed.

Hence we see that three forms or species of Bos, originally inhabitants of Europe, have been domesticated; but there is no improbability in this fact, for the genus Bos readily yields to domestication. Besides these three species and the zebu, the yak, the gayal, and the arni^[190] (not to mention the buffalo or genus Bubalus) have been domesticated; making altogether seven species of Bos. The zebu and the three European species are now extinct in a wild state, for the cattle of the B. primigenius type in the British parks can hardly be considered as truly wild. Although certain races of cattle, domesticated at a very ancient period in Europe, are the descendants of the three above-named fossil species, yet it does not follow that they were here first domesticated. Those who place much reliance on philology argue that our cattle were imported from the East.^[191] But as races of men invading any country would probably give their own names to the breeds of cattle which they might there find domesticated, the argument seems inconclusive. There is indirect evidence that our cattle are the descendants of species which originally inhabited a temperate or cold climate, but not a land long covered with snow; for our cattle, as we have seen in the chapter on Horses, apparently have not the instinct of scraping away the snow to get at the herbage beneath. No one could behold the magnificent wild bulls on the bleak Falkland Islands in the southern hemisphere, and doubt about the climate being admirably suited to them. Azara has remarked that in the temperate regions of La Plata the cows conceive when two years old, whilst in the much hotter country of Paraguay they do not conceive till three years old; "from which fact," as he adds, "one may conclude that cattle do not succeed so well in warm countries."^[192]

The above-named three fossil forms of Bos have been ranked {83}by nearly all palæontologists as distinct species; and it would not be reasonable to change their denomination simply because they are now found to be the parents of several domesticated races. But what is of most importance for us, as showing that they deserve to be ranked as species, is that they co-existed in different parts of Europe during the same period, and yet kept distinct. Their domesticated descendants, on the other hand, if not separated, cross with the utmost freedom and become commingled. The several European breeds have so often been crossed, both intentionally and unintentionally, that, if any sterility ensued from such unions, it would certainly have been detected. As zebus inhabit a distant and much hotter region, and as they differ in so many characters from our European cattle, I have taken pains to ascertain whether the two forms are fertile when crossed. The late Lord Powis imported some zebus and crossed them with common cattle in Shropshire; and I was assured by his steward that the cross-bred animals were perfectly fertile with both parent-stocks. Mr. Blyth informs me that in India hybrids, with various proportions of either blood, are quite fertile; and this can hardly fail to be known, for in some districts^[193] the two species are allowed to breed freely together. Most of the cattle which were first introduced into Tasmania were humped, so that at one time thousands of crossed animals existed there; and Mr. B. O'Neile Wilson, M.A., writes to me from Tasmania that he has never heard of any sterility having been observed. He himself formerly possessed a herd of such crossed cattle, and all were perfectly fertile; so much so, that he cannot remember even a single cow failing to calve. These several facts afford an important confirmation of the Pallasian doctrine that the descendants of species which when first domesticated would if crossed probably have been in some degree sterile, become perfectly fertile after a long course of domestication. In a future chapter we shall see that this doctrine throws much light on the difficult subject of Hybridism.

I have alluded to the cattle in Chillingham Park, which, according to Rütimeyer, have been very little changed from the Bos primigenius type. This park is so ancient that it is {84}referred to in a record of the year 1220. The cattle in their instincts and habits are truly wild. They are white, with the inside of the ears reddish-brown, eyes rimmed with black, muzzles brown, hoofs black, and horns white tipped with black. Within a period of thirty-three years about a dozen calves were born with "brown and blue spots upon the cheeks or necks; but these, together with any defective animals, were always destroyed." According to Bewick, about the year 1770 some calves appeared with black ears; but these were also destroyed by the keeper, and black ears have not since reappeared. The wild white cattle in the Duke of Hamilton's park, where I have heard of the birth of a black calf, are said by Lord Tankerville to be inferior to those at Chillingham. The cattle kept until the year 1780 by the Duke of Queensberry, but now extinct, had their ears, muzzle, and orbits of the eyes black. Those which have existed from time immemorial at Chartley; closely resemble the cattle at Chillingham, but are larger, "with some small difference in the colour of the ears." "They frequently tend to become entirely black; and a singular superstition prevails in the vicinity that, when a black calf is born, some calamity impends over the noble house of Ferrers. All the black calves are destroyed." The cattle at Burton Constable in Yorkshire, now extinct, had ears, muzzle, and the tip of the tail black. Those at Gisburne, also in Yorkshire, are said by Bewick to have been sometimes without dark muzzles, with the inside alone of the ears brown; and they are elsewhere said to have been low in stature and hornless.^[194]

The several above-specified differences in the park-cattle, slight though they be, are worth recording, as they show that animals living nearly in a state of nature, and exposed to nearly uniform conditions, if not allowed to roam freely and to cross with other herds, do not keep as uniform as truly {85}wild animals. For the preservation of a uniform character, even within the same park, a certain degree of selection—that is, the destruction of the dark-coloured calves—is apparently necessary.

The cattle in all the parks are white; but, from the occasional appearance of dark-coloured calves, it is extremely doubtful whether the aboriginal Bos primigenius was white. The following facts, however, show that there is a strong, though not invariable, tendency in wild or escaped cattle, under widely different conditions of life, to become white with coloured ears. If the old writers Boethius and Leslie^[195] can be trusted, the wild cattle of Scotland were white and furnished with a great mane; but the colour of their ears is not mentioned. The primæval forest formerly extended across the whole country from Chillingham to Hamilton, and Sir Walter Scott used to maintain that the cattle still preserved in these two parks, at the two extremities of the forest, were remnants of its original inhabitants; and this view certainly seems probable. In Wales,^[196] during the tenth century, some of the cattle are described as being white with red ears. Four hundred cattle thus coloured were sent to King John; and an early record speaks of a hundred cattle with red ears having been demanded as a compensation for some offence, but, if the cattle were of a dark or black colour, one hundred and fifty were to be presented. The black cattle of North Wales apparently belong, as we have seen, to the small longifrons type: and as the alternative was offered of either 150 dark cattle, or 100 white cattle with red ears, we may presume that the latter were the larger beasts, and probably belonged to the primigenius type. Youatt has remarked that at the present day, whenever cattle of the short-horn breed are white, the extremities of their ears are more or less tinged with red.

The cattle which have run wild on the Pampas, in Texas, and in two parts of Africa, have become of a nearly uniform dark {86}brownish-red.^[197] On the Ladrone Islands, in the Pacific Ocean, immense herds of cattle, which were wild in the year 1741, are described as "milk-white, except their ears, which are generally black."^[198] The Falkland Islands, situated far south, with all the conditions of life as different as it is possible to conceive from those of the Ladrones, offer a more interesting case. Cattle have run wild there during eighty or ninety years; and in the southern districts the animals are mostly white, with their feet, or whole heads, or only their ears black; but my informant, Admiral Sulivan,^[199] who long resided on these islands, does not believe that they are ever purely white. So that in these two archipelagos we see that the cattle tend to become white with coloured ears. In other parts of the Falkland Islands, other colours prevail: near Port Pleasant brown is the common tint; round Mount Usborne, about half the animals in some of the herds were lead or mouse-coloured, which elsewhere is an unusual tint. These latter cattle, though generally inhabiting high land, breed about a month earlier than the other cattle; and this circumstance would aid in keeping them distinct and in perpetuating this peculiar colour. It is worth recalling to mind that blue or lead-coloured marks have occasionally appeared on the white cattle of Chillingham. So plainly different were the colours of the wild herds in different parts of the Falkland Islands, that in hunting them, as Admiral Sulivan informs me, white spots in one district, and dark spots in another district, were always looked out for on the distant hills. In the intermediate districts intermediate colours prevailed. Whatever the cause may be, this tendency in the wild cattle of the Falkland Islands, which are all descended from a few brought from La Plata, to break up into herds of three different colours, is an interesting fact.

Returning to the several British breeds, the conspicuous difference in general appearance between Short-horns, Long-horns (now rarely seen), Herefords, Highland cattle, Alderneys, &c., must be familiar to every one. A large part of the {87}difference, no doubt, may be due to descent from primordially distinct species; but we may feel sure that there has been in addition a considerable amount of variation. Even during the Neolithic period, the domestic cattle were not actually identical with the aboriginal species. Within recent times most of the breeds have been modified by careful and methodical selection. How strongly the characters thus acquired are inherited, may be inferred from the prices realised by the improved breeds; even at the first sale of Colling's Short-horns, eleven bulls reached an average of 214l., and lately Short-horn bulls have been sold for a thousand guineas, and have been exported to all quarters of the world.

Some constitutional differences may be here noticed. The Short-horns arrive at maturity far earlier than the wilder breeds, such as those of Wales or the Highlands. This fact has been shown in an interesting manner by Mr. Simonds,^[200] who has given a table of the average period of their dentition, which proves that there is a difference of no less than six months in the appearance of the permanent incisors. The period of gestation, from observations made by Tessier on 1131 cows, varies to the extent of eighty-one days; and what is more interesting, M. Lefour affirms "that the period of gestation is longer in the large German cattle than in the smaller breeds."^[201] With respect to the period of conception, it seems certain that Alderney and Zetland cows often become pregnant earlier than other breeds.^[202] Lastly, as four fully-developed mammæ is a generic character in the genus Bos,^[203] it is worth notice that with our domestic cows the two rudimentary mammæ often become fairly well developed and yield milk.

As numerous breeds are generally found only in long-civilized countries, it may be well to show that in some countries inhabited by barbarous races, who are frequently at war with each other and therefore have little free {88}communication, several distinct breeds of cattle now exist or formerly existed. At the Cape of Good Hope Leguat observed, in the year 1720, three kinds.^[204] At the present day various travellers have noticed the differences in the breeds in Southern Africa. Sir Andrew Smith several years ago remarked to me that the cattle possessed by the different tribes of Caffres, though living near each other under the same latitude and in the same kind of country, yet differed, and he expressed much surprise at the fact. Mr. Andersson has described^[205] the Damara, Bechuana, and Namaqua cattle; and he informs me in a letter that the cattle north of Lake Ngami are likewise different, as Mr. Galton has heard is the case with the cattle of Benguela. The Namaqua cattle in size and shape nearly resemble European cattle, and have short stout horns and large hoofs. The Damara cattle are very peculiar, being big-boned, with slender legs and small hard feet; their tails are adorned with a tuft of long bushy hair nearly touching the ground, and their horns are extraordinarily large. The Bechuana cattle have even larger horns, and there is now a skull in London with the two horns 8 ft. 8¼ in. long, as measured in a straight line from tip to tip, and no less than 13ft. 5in. as measured along their curvature! Mr. Andersson in his letter to me says that, though he will not venture to describe the differences between the breeds belonging to the many different sub-tribes, yet such certainly exist, as shown by the wonderful facility with which the natives discriminate them.

That many breeds of cattle have originated through variation, independently of descent from distinct species, we may infer from what we see in South America, where the genus Bos was not endemic, and where the cattle which now exist in such vast numbers are the descendants of a few imported from Spain and Portugal. In Columbia, Roulin^[206] describes two peculiar breeds, namely, pelones, with extremely thin and fine hair, and calongos, absolutely naked. According to Castelnau there are two races in Brazil, one like European cattle, the other different, with {89}remarkable horns. In Paraguay, Azara describes a breed which certainly originated in S. America, called chivos, "because they have straight vertical horns, conical, and very large at the base." He likewise describes a dwarf race in Corrientes, with short legs and a body larger than usual. Cattle without horns, and others with reversed hair, have also originated in Paraguay.

Another monstrous breed, called niatas or natas, of which I saw two small herds on the northern bank of the Plata, is so remarkable as to deserve a fuller description. This breed bears the same relation to other breeds, as bull or pug dogs do to other dogs, or as improved pigs, according to H. von Nathusius, do to common pigs.^[207] Rütimeyer believes that these cattle belong to the primigenius type.^[208] The forehead is very short and broad, with the nasal end of the skull, together with the whole plane of the upper molar-teeth, curved upwards. The lower jaw projects beyond the upper, and has a corresponding upward curvature. It is an interesting fact that an almost similar conformation characterizes, as I have been informed by Dr. Falconer, the extinct and gigantic Sivatherium of India, and is not known in any other ruminant. The upper lip is much drawn back, the nostrils are seated high up and are widely open, the eyes project outwards, and the horns are large. In walking the head is carried low, and the neck is short. The hind legs appear to be longer, compared with the front legs, than is usual. The exposed incisor teeth, the short head and upturned nostrils, give these cattle the most ludicrous, self-confident air of defiance. The skull which I presented to the College of Surgeons has been thus described by Professor Owen:^[209] "It is remarkable from the stunted development of the nasals, premaxillaries, and fore-part of the lower jaw, which is unusually {90}curved upwards to come into contact with the premaxillaries. The nasal bones are about one-third the ordinary length, but retain almost their normal breadth. The triangular vacuity is left between them, the frontal and lachrymal, which latter bone articulates with the premaxillary, and thus excludes the maxillary from any junction with the nasal." So that even the connexion of some of the bones is changed. Other differences might be added: thus the plane of the condyles is somewhat modified, and the terminal edge of the premaxillaries forms an arch. In fact, on comparison with the skull of a common ox, scarcely a single bone presents the same exact shape, and the whole skull has a wonderfully different appearance.

The first brief published notice of this race was by Azara, between the years 1783-96; but Don F. Muniz, of Luxan, who has kindly collected information for me, states that about 1760 these cattle were kept as curiosities near Buenos Ayres. Their origin is not positively known, but they must have originated subsequently to the year 1552, when cattle were first introduced. Signor Muniz informs me that the breed is believed to have originated with the Indians southward of the Plata. Even to this day those reared near the Plata show their less civilized nature in being fiercer than common cattle, and in the cow, if visited too often, easily deserting her first calf. The breed is very true, and a niata bull and cow invariably produce niata calves. The breed has already lasted at least a century. A niata bull crossed with a common cow, and the reverse cross, yield offspring having an intermediate character, but with the niata character strongly displayed. According to Signor Muniz, there is the clearest evidence, contrary to the common belief of agriculturists in analogous cases, that the niata cow when crossed with a common bull transmits her peculiarities more strongly than does the niata bull when crossed with a common cow. When the pasture is tolerably long, these cattle feed as well as common cattle with their tongue and palate; but during the great droughts, when so many animals perish on the Pampas, the niata breed lies under a great disadvantage, and would, if not attended to, become extinct; for the common cattle, like horses, are able just to keep alive by browsing on the twigs of trees and on reeds with their lips: this the niatas cannot so {91}well do, as their lips do not join, and hence they are found to perish before the common cattle. This strikes me as a good illustration of how little we are able to judge from the ordinary habits of an animal, on what circumstances, occurring only at long intervals of time, its rarity or extinction may depend. It shows us, also, how natural selection would have determined the rejection of the niata modification had it arisen in a state of nature.

Having described the semi-monstrous niata breed, I may allude to a white bull, said to have been brought from Africa, which was exhibited in London in 1829, and which has been well figured by Mr. Harvey.^[210] It had a hump, and was furnished with a mane. The dewlap was peculiar, being divided between its fore-legs into parallel divisions. Its lateral hoofs were annually shed, and grew to the length of five or six inches. The eye was very peculiar, being remarkably prominent, and "resembled a cup and ball, thus enabling the animal to see on all sides with equal ease; the pupil was small and oval, or rather a parallelogram with the ends cut off, and lying transversely across the ball," A new and strange breed might probably have been formed by careful breeding and selection from this animal.

I have often speculated on the probable causes through which each separate district in Great Britain came to possess in former times its own peculiar breed of cattle; and the question is, perhaps, even more perplexing in the case of Southern Africa. We now know that the differences may be in part attributed to descent from distinct species; but this will not suffice. Have the slight differences in climate and in the nature of the pasture, in the different districts of Britain, directly induced corresponding differences in the cattle? We have seen that the semi-wild cattle in the several British parks are not identical in colouring or size, and that some degree of selection has been requisite to keep them true. It is almost certain that abundant food given during many generations directly affects the size of a breed.^[211] That climate directly affects the thickness of the {92}skin and the hair is likewise certain: thus Roulin asserts^[212] that the hides of the feral cattle on the hot Llanos "are always much less heavy than those of the cattle raised on the high platform of Bogota; and that these hides yield in weight and in thickness of hair to those of the cattle which have run wild on the lofty Paramos." The same difference has been observed in the hides of the cattle reared on the bleak Falkland Islands and on the temperate Pampas. Low has remarked^[213] that the cattle which inhabit the more humid parts of Britain have longer hair and thicker skins than other British cattle; and the hair and horns are so closely related to each other, that, as we shall see in a future chapter, they are apt to vary together; thus climate might indirectly affect, through the skin, the form and size of the horns. When we compare highly improved stall-fed cattle with the wilder breeds, or compare mountain and lowland breeds, we cannot doubt that an active life, leading to the free use of the limbs and lungs, affects the shape and proportions of the whole body. It is probable that some breeds, such as the semi-monstrous niata cattle, and some peculiarities, such as being hornless, &c., have appeared suddenly from what we may call a spontaneous variation; but even in this case a rude kind of selection is necessary, and the animals thus characterized must be at least partially separated from others. This degree of care, however, has sometimes been taken even in little-civilized districts, where we should least have expected it, as in the case of the niata, chivo, and hornless cattle in S. America.

That methodical selection has done wonders within a recent period in modifying our cattle, no one doubts. During the process of methodical selection it has occasionally happened that deviations of structure, more strongly pronounced than mere individual differences, yet by no means deserving to be called monstrosities, have been taken advantage of: thus the famous Long-horn Bull, Shakespeare, though of the pure Canley stock, "scarcely inherited a single point of the long-horned breed, his horns excepted;^[214] yet in the hands of Mr. Fowler, {93}this bull greatly improved his race. We have also reason to believe that selection, carried on so far unconsciously that there was at no one time any distinct intention to improve or change the breed, has in the course of time modified most of our cattle; for by this process, aided by more abundant food, all the lowland British breeds have increased greatly in size and in early maturity since the reign of Henry VII.^[215] It should never be forgotten that many animals have to be annually slaughtered; so that each owner must determine which shall be killed and which preserved for breeding. In every district, as Youatt has remarked, there is a prejudice in favour of the native breed; so that animals possessing qualities, whatever they may be, which are most valued in each district, will be oftenest preserved; and this unmethodical selection assuredly will in the long run affect the character of the whole breed. But it may be asked, can this rude kind of selection have been practised by barbarians such as those of southern Africa? In a future chapter on Selection we shall see that this has certainly occurred to some extent. Therefore, looking to the origin of the many breeds of cattle which formerly inhabited the several districts of Britain, I conclude that, although slight differences in the nature of the climate, food, &c., as well as changed habits of life, aided by correlation of growth, and the occasional appearance from unknown causes of considerable deviations of structure, have all probably played their parts; yet that the occasional preservation in each district of those individual animals which were most valued by each owner has perhaps been even more effective in the production of the several British breeds. As soon as two or more breeds had once been formed in any district, or when new breeds descended from distinct species were introduced, their crossing, especially if aided by some selection, will have multiplied the number and modified the characters of the older breeds.

Sheep.

I shall treat this subject briefly. Most authors look at our domestic sheep as descended from several distinct species; but how many still exist is doubtful. Mr. Blyth believes that there {94}are in the whole world fourteen species, one of which, the Corsican moufflon, he concludes (as I am informed by him) to be the parent of the smaller, short-tailed breeds, with crescent-shaped horns, such as the old Highland sheep. The larger, long-tailed breeds, having horns with a double flexure, such as the Dorsets, merinos, &c., he believes to be descended from an unknown and extinct species. M. Gervais makes six species of Ovis;^[216] but concludes that our domestic sheep form a distinct genus, now completely extinct. A German naturalist^[217] believes that our sheep descend from ten aboriginally distinct species, of which only one is still living in a wild state! Another ingenious observer,^[218] though not a naturalist, with a bold defiance of everything known on geographical distribution, infers that the sheep of Great Britain alone are the descendants of eleven endemic British forms! Under such a hopeless state of doubt it would be useless for my purpose to give a detailed account of the several breeds; but a few remarks may be added.

Sheep have been domesticated from a very ancient period. Rütimeyer^[219] found in the Swiss lake-dwellings the remains of a small breed, with thin and tall legs, and with horns like those of a goat: this race differs somewhat from any one now known. Almost every country has its own peculiar breed; and many countries have many breeds differing greatly from each other. One of the most strongly marked races is an Eastern one with a long tail, including, according to Pallas, twenty vertebræ, and so loaded with fat, that, from being esteemed a delicacy, it is sometimes placed on a truck which is dragged about by the living animal. These sheep, though ranked by Fitzinger as a distinct aboriginal form, seem to bear in their drooping ears the stamp of long domestication. This is likewise the case with those sheep which have two great masses of fat on the rump, with the tail in a rudimentary condition. The Angola variety of {95}the long-tailed race has curious masses of fat on the back of the head and beneath the jaws.^[220] Mr. Hodgson in an admirable paper^[221] on the sheep of the Himalaya infers from the distribution of the several races, "that this caudal augmentation in most of its phases is an instance of degeneracy in these pre-eminently Alpine animals." The horns present an endless diversity in character; being, especially in the female sex, not rarely absent, or, on the other hand, amounting to four or even eight in number. The horns, when numerous, arise from a crest on the frontal bone, which is elevated in a peculiar manner. It is remarkable that multiplicity of horns "is generally accompanied by great length and coarseness of the fleece."^[222] This correlation, however, is not invariable; for I am informed by Mr. D. Forbes, that the Spanish sheep in Chile resemble, in fleece and in all other characters, their parent merino-race, except that instead of a pair they generally bear four horns. The existence of a pair of mammæ is a generic character in the genus Ovis as well as in several allied forms; nevertheless, as Mr. Hodgson has remarked, "this character is not absolutely constant even among the true and proper sheep: for I have more than once met with Cágias (a sub-Himalayan domestic race) possessed of four teats."^[223] This case is the more remarkable as, when any part or organ is present in reduced number in comparison with the same part in allied groups, it usually is subject to little variation. The presence of interdigital pits has likewise been considered as a generic distinction in sheep; but Isidore Geoffroy^[224] has shown that these pits or pouches are absent in some breeds.

In sheep there is a strong tendency for characters, which have apparently been acquired under domestication, to become attached either exclusively to the male sex, or to be more highly developed in this than in the other sex. Thus in many breeds the horns are deficient in the ewe, though this likewise occurs occasionally with the female of the wild musmon. In the rams of the Wallachian breed "the horns spring almost perpendicularly {96}from the frontal bone, and then take a beautiful spiral form; in the ewes they protrude nearly at right angles from the head, and then become twisted in a singular manner."^[225] Mr. Hodgson states that the extraordinarily arched nose or chaffron, which is so highly developed in several foreign breeds, is characteristic of the ram alone, and apparently is the result of domestication.^[226] I hear from Mr. Blyth that the accumulation of fat in the fat-tailed sheep of the plains of India is greater in the male than in the female; and Fitzinger^[227] remarks that the mane in the African maned race is far more developed in the ram than in the ewe.

Different races of sheep, like cattle, present constitutional differences. Thus the improved breeds arrive at maturity at an early age, as has been well shown by Mr. Simonds through their early average period of dentition. The several races have become adapted to different kinds of pasture and climate: for instance, no one can rear Leicester sheep on mountainous regions, where Cheviots flourish. As Youatt has remarked, "in all the different districts of Great Britain we find various breeds of sheep beautifully adapted to the locality which they occupy. No one knows their origin; they are indigenous to the soil, climate, pasturage, and the locality on which they graze; they seem to have been formed for it and by it."^[228] Marshall relates^[229] that a flock of heavy Lincolnshire and light Norfolk sheep which had been bred together in a large sheep-walk, part of which was low, rich, and moist, and another part high and dry, with benty grass, when turned out, regularly separated from each other; the heavy sheep drawing off to the rich soil, and the lighter sheep to their own soil; so that "whilst there was plenty of grass the two breeds kept themselves as distinct as rooks and pigeons." Numerous sheep from various parts of the world have been brought during a long course of years to the Zoological Gardens of London; but as Youatt, who attended the animals as a {97}veterinary surgeon, remarks, "few or none die of the rot, but they are phthisical; not one of them from a torrid climate lasts out the second year, and when they die their lungs are tuberculated."^[230] Even in certain parts of England it has been found impossible to keep certain breeds of sheep; thus on a farm on the banks of the Ouse, the Leicester sheep were so rapidly destroyed by pleuritis^[231] that the owner could not keep them; the coarser-skinned sheep never being affected.

The period of gestation was formerly thought to be so unalterable a character, that a supposed difference between the wolf and the dog in this respect was esteemed a sure sign of specific distinction; but we have seen that the period is shorter in the improved breeds of the pig, and in the larger breeds of the ox, than in other breeds of these two animals. And now we know, on the excellent authority of Hermann von Nathusius,^[232] that Merino and Southdown sheep, when both have long been kept under exactly the same conditions, differ in their average period of gestation, as is seen in the following Table:—

In this graduated difference, in these cross-bred animals having different proportions of Southdown blood, we see how strictly the two periods of gestation have been transmitted. Nathusius remarks that, as Southdowns grow with remarkable rapidity after birth, it is not surprising that their fœtal development should have been shortened. It is of course possible that the difference in these two breeds may be due to their descent from distinct parent-species; but as the early maturity of the Southdowns has long been carefully attended to by breeders, the difference is more probably the result of such attention. Lastly, the fecundity of the several breeds differs much; some generally producing twins or even triplets at a birth, of which fact the curious Shangai sheep (with their truncated and rudimentary {98}ears, and great Roman noses), lately exhibited in the Zoological Gardens, offer a remarkable instance.

Sheep are perhaps more readily affected by the direct action of the conditions of life to which they have been exposed than almost any other domestic animal. According to Pallas, and more recently according to Erman, the fat-tailed Kirghisian sheep, when bred for a few generations in Russia, degenerate, and the mass of fat dwindles away, "the scanty and bitter herbage of the steppes seems so essential to their development." Pallas makes an analogous statement with respect to one of the Crimean breeds. Burnes states that the Karakool breed, which produces a fine, curled, black, and valuable fleece, when removed from its own canton near Bokhara to Persia or to other quarters, loses its peculiar fleece.^[233] In all such cases, however, it may be that a change of any kind in the conditions of life causes variability and consequent loss of character, and not that certain conditions are necessary for the development of certain characters.

Great heat, however, seems to act directly on the fleece: several accounts have been published of the change which sheep imported from Europe undergo in the West Indies. Dr. Nicholson of Antigua informs me that, after the third generation, the wool disappears from the whole body, except over the loins; and the animal then appears like a goat with a dirty door-mat on its back. A similar change is said to take place on the west coast of Africa.^[234] On the other hand, many wool-bearing sheep live on the hot plains of India. Roulin asserts that in the lower and heated valleys of the Cordillera, if the lambs are sheared as soon as the wool has grown to a certain thickness, all goes on afterwards as usual; but if not sheared, the wool detaches itself in flakes, and short shining hair like that {99}on a goat is produced ever afterwards. This curious result seems merely to be an exaggerated tendency natural to the Merino breed, for as a great authority, namely, Lord Somerville, remarks, "the wool of our Merino sheep after shear-time is hard and coarse to such a degree as to render it almost impossible to suppose that the same animal could bear wool so opposite in quality, compared to that which has been clipped from it: as the cold weather advances, the fleeces recover their soft quality." As in sheep of all breeds the fleece naturally consists of longer and coarser hair covering shorter and softer wool, the change which it often undergoes in hot climates is probably merely a case of unequal development; for even with those sheep which like goats are covered with hair, a small quantity of underlying wool may always be found.^[235] In the wild mountain-sheep (Ovis montana) of North America there is an annual analogous change of coat; "the wool begins to drop out in early spring, leaving in its place a coat of hair resembling that of the elk, a change of pelage quite different in character from the ordinary thickening of the coat or hair, common to all furred animals in winter,—for instance, in the horse, the cow, &c., which shed their winter coat in the spring."^[236]

A slight difference in climate or pasture sometimes slightly affects the fleece, as has been observed even in different districts in England, and as is well shown by the great softness of the wool brought from Southern Australia. But it should be observed, as Youatt repeatedly insists, that the tendency to change may generally be counteracted by careful selection. M. Lasterye, after discussing this subject, sums up as follows: "The preservation of the Merino race in its utmost purity at the Cape of Good Hope, in the marshes of Holland, and under the rigorous climate of Sweden, furnishes an additional support of this my unalterable principle, that fine-woolled sheep may be kept wherever industrious men and intelligent breeders exist."

That methodical selection has effected great changes in several {100}breeds of sheep no one, who knows anything on the subject, entertains a doubt. The case of the Southdowns, as improved by Ellman, offers perhaps the most striking instance. Unconscious or occasional selection has likewise slowly produced a great effect, as we shall see in the chapters on Selection. That crossing has largely modified some breeds, no one who will study what has been written on this subject—for instance, Mr. Spooner's paper—will dispute; but to produce uniformity, in a crossed breed, careful selection and "rigorous weeding," as this author expresses it, are indispensable.^[237]

In some few instances new breeds have suddenly originated; thus, in 1791, a ram-lamb was born in Massachusetts, having short crooked legs and a long back, like a turnspit-dog. From this one lamb the otter or ancon semi-monstrous breed was raised; as these sheep could not leap over the fences, it was thought that they would be valuable; but they have been supplanted by merinos, and thus exterminated. These sheep are remarkable from transmitting their character so truly that Colonel Humphreys^[238] never heard of "but one questionable case" of an ancon ram and ewe not producing ancon offspring. When they are crossed with other breeds the offspring, with rare exceptions, instead of being intermediate in character, perfectly resemble either parent; and this has occurred even in the case of twins. Lastly, "the ancons have been observed to keep together, separating themselves from the rest of the flock when put into enclosures with other sheep."

A more interesting case has been recorded in the Report of the Juries for the Great Exhibition (1851), namely, the production of a merino ram-lamb on the Mauchamp farm, in 1828, which was remarkable for its long, smooth, straight, and silky wool. By the year 1833 M. Graux had raised rams enough to serve his whole flock, and after a few more years he was able to sell stock of his new breed. So peculiar and valuable is the wool, that it sells at 25 per cent. above the best merino wool: even the fleeces of half-bred animals are valuable, and are known in France as the "Mauchamp-merino." It is interesting, as {101}showing how generally any marked deviation of structure is accompanied by other deviations, that the first ram and his immediate offspring were of small size, with large heads, long necks, narrow chests, and long flanks; but these blemishes were removed by judicious crosses and selection. The long smooth wool was also correlated with smooth horns; and as horns and hair are homologous structures, we can understand the meaning of this correlation. If the Mauchamp and ancon breeds had originated a century or two ago, we should have had no record of their birth; and many a naturalist would no doubt have insisted, especially in the case of the Mauchamp race, that they had each descended from, or been crossed with, some unknown aboriginal form.

Goats.

From the recent researches of M. Brandt, most naturalists now believe that all our goats are descended from the Capra ægagrus of the mountains of Asia, possibly mingled with the allied Indian species C. Falconeri of India.^[239] In Switzerland, during the early Stone period, the domestic goat was commoner than the sheep; and this very ancient race differed in no respect from that now common in Switzerland.^[240] At the present time, the many races found in several parts of the world differ greatly from each other; nevertheless, as far as they have been tried,^[241] they are all quite fertile when crossed. So numerous are the breeds, that Mr. G. Clark^[242] has described eight distinct kinds imported into the one island of Mauritius. The ears of one kind were enormously developed, being, as measured by Mr. Clark, no less than 19 inches in length and 4¾ inches in breadth. As with cattle, the mammæ of those breeds which are regularly milked become greatly developed; and, as Mr. Clark remarks, "it is not rare to see their teats touching the ground." The following cases are worth notice as presenting unusual {102}points of variation. According to Godron,^[243] the mammæ differ greatly in shape in different breeds, being elongated in the common goat, hemispherical in the Angora race, and bilobed and divergent in the goats of Syria and Nubia. According to this same author, the males of certain breeds have lost their usual offensive odour. In one of the Indian breeds the males and females have horns of widely-different shapes;^[244] and in some breeds the females are destitute of horns.^[245] The presence of interdigital pits or glands on all four feet has been thought to characterise the genus Ovis, and their absence to be characteristic of the genus Capra; but Mr. Hodgson has found that they exist in the front feet of the majority of Himalayan goats.^[246] Mr. Hodgson measured the intestines in two goats of the Dúgú race, and he found that the proportional length of the great and small intestines differed considerably. In one of these goats the cæcum was thirteen inches, and in the other no less than thirty-six inches in length!

{103}

CHAPTER IV.

DOMESTIC RABBITS.

DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RABBIT—ANCIENT DOMESTICATION—ANCIENT SELECTION—LARGE LOP-EARED RABBITS—VARIOUS BREEDS—FLUCTUATING CHARACTERS—ORIGIN OF THE HIMALAYAN BREED—CURIOUS CASE OF INHERITANCE—FERAL RABBITS IN JAMAICA AND THE FALKLAND ISLANDS—PORTO SANTO FERAL RABBITS—OSTEOLOGICAL CHARACTERS—SKULL—SKULL OF HALF-LOP RABBITS—VARIATIONS IN THE SKULL ANALOGOUS TO DIFFERENCES IN DIFFERENT SPECIES OF HARES—VERTEBRÆ—STERNUM—SCAPULA—EFFECTS OF USE AND DISUSE ON THE PROPORTIONS OF THE LIMBS AND BODY—CAPACITY OF THE SKULL AND REDUCED SIZE OF THE BRAIN—SUMMARY ON THE MODIFICATIONS OF DOMESTICATED RABBITS.

All naturalists, with, as far as I know, a single exception, believe that the several domestic breeds of the rabbit are descended from the common wild species; I shall therefore describe them more carefully than in the previous cases. Professor Gervais^[247] states "that the true wild rabbit is smaller than the domestic; its proportions are not absolutely the same; its tail is smaller; its ears are shorter and more thickly clothed with hair; and these characters, without speaking of colour, are so many indications opposed to the opinion which unites these animals under the same specific denomination." Few naturalists will agree with this author that such slight differences are sufficient to separate as distinct species the wild and domestic rabbit. How extraordinary it would be, if close confinement, perfect tameness, unnatural food, and careful breeding, all prolonged during many generations, had not produced at least some effect! The tame rabbit has been domesticated from an ancient period. Confucius ranges rabbits among animals worthy to be sacrificed to the gods, and, as he prescribes their multiplication, they were probably at this early period domesticated in China. They are mentioned by several of the classical writers. {104}In 1631 Gervaise Markham writes, "You shall not, as in other cattell, looke to their shape, but to their richnesse, onely elect your buckes, the largest and goodliest conies you can get; and for the richnesse of the skin, that is accounted the richest which hath the equallest mixture of blacke and white haire together, yet the blacke rather shadowing the white; the furre should be thicke, deepe, smooth, and shining; ... they are of body much fatter and larger, and, when another skin is worth two or three pence, they are worth two shillings." From this full description we see that silver-grey rabbits existed in England at this period; and, what is far more important, we see that the breeding or selection of rabbits was then carefully attended to. Aldrovandi, in 1637, describes, on the authority of several old writers (as Scaliger, in 1557), rabbits of various colours, some "like a hare," and he adds that P. Valerianus (who died a very old man in 1558) saw at Verona rabbits four times bigger than ours.^[248]

From the fact of the rabbit having been domesticated at an ancient period, we must look to the northern hemisphere of the Old World, and to the warmer temperate regions alone, for the aboriginal parent-form; for the rabbit cannot live without protection in countries as cold as Sweden, and, though it has run wild in the tropical island of Jamaica, it has never greatly multiplied there. It now exists, and has long existed, in the warmer temperate parts of Europe, for fossil remains have been found in several countries.^[249] The domestic rabbit readily becomes feral in these same countries, and when variously coloured kinds are turned out they generally revert to the ordinary grey colour.^[250] The wild rabbits, if taken young, can be domesticated, though the process is generally very troublesome.^[251] The various {105}domestic races are often crossed, and are believed to be perfectly fertile together, and a perfect gradation can be shown to exist from the largest domestic kinds, having enormously developed ears, to the common wild kind. The parent-form must have been a burrowing animal, a habit not common, as far as I can discover, to any other species in the large genus Lepus. Only one wild species is known with certainty to exist in Europe; but the rabbit (if it be a true rabbit) from Mount Sinai, and likewise that from Algeria, present slight differences; and these forms have been considered by some authors as specifically distinct.^[252] But such slight differences would aid us little in explaining the more considerable differences characteristic of the several domestic races. If the latter are the descendants of two or more closely allied species, all, excepting the common rabbit, have been exterminated in a wild state; and this is very improbable, seeing with what pertinacity this animal holds its ground. From these several reasons we may infer with safety that all the domestic breeds are the descendants of the common wild species. But from what we hear of the late marvellous success in rearing hybrids between the hare and rabbit,^[253] it is possible, though not probable, from the great difficulty in making the first cross, that some of the larger races, which are coloured like the hare, may have been modified by crosses with this animal. Nevertheless, the chief differences in the skeletons of the several domestic breeds cannot, as we shall presently see, have been derived from a cross with the hare.

There are many breeds which transmit their characters more or less truly. Every one has seen the enormous lop-eared rabbits exhibited at our shows; various allied sub-breeds are reared on the Continent, such as the so-called Andalusian, which is said to have a large head with a round forehead, and to attain a greater size than any other kind; another large Paris breed is named the Rouennais, and has a square head; the so-called Patagonian rabbit has remarkably short ears and a large round head. Although I have not seen all these breeds, I feel some doubt about there being any marked difference in the {106}shape of their skulls.^[254] English lop-eared rabbits often weigh 8 lbs. or 10 lbs., and one has been exhibited weighing 18 lbs.; whereas a full-sized wild rabbit weighs only about 3¼ lbs. The head or skull in all the large lop-eared rabbits examined by me is much longer relatively to its breadth than in the wild rabbit. Many of them have loose transverse folds of skin or dewlaps beneath the throat, which can be pulled out so as to reach nearly to the ends of the jaws. Their ears are prodigiously developed, and hang down on each side of their faces. A rabbit has been exhibited with its two ears, measured from the tip of one to the tip of the other, 22 inches in length, and each ear was 5⅜ inches in breadth. In a common wild rabbit I found that the length of the two ears, from tip to tip, was 7⅝ inches, and the breadth only 1⅞ inch. The great weight of the body in the larger rabbits, and the immense development of their ears, are the qualities which win prizes, and have been carefully selected.

The hare-coloured, or, as it is sometimes called, the Belgian rabbit, differs in nothing except colour from the other large breeds; but Mr. J. Young, of Southampton, a great breeder of this kind, informs me that the females, in all the specimens examined by him, had only six mammæ; and this certainly was the case with two females which came into my possession. Mr. B. P. Brent, however, assures me that the number is variable with other domestic rabbits. The common wild rabbit always has ten mammæ. The Angora rabbit is remarkable from the length and fineness of its fur, which even on the soles of the feet is of considerable length. This breed is the only one which differs in its mental qualities, for it is said to be much more sociable than other rabbits, and the male shows no wish to destroy its young.^[255] Two live rabbits were brought to me from Moscow, of about the size of the wild species, but with long soft fur, different from that of the Angora. These Moscow rabbits had pink eyes and were snow-white, excepting the ears, two spots near the nose, the upper and under surface of the tail, and the hinder tarsi, which were blackish-brown. In short, they were {107}coloured nearly like the so-called Himalayan rabbits, presently to be described, and differed from them only in the character of their fur. There are two other breeds which come true to colour, but differ in no other respect, namely silver-greys and chinchillas. Lastly, the Nicard or Dutch rabbit may be mentioned, which varies in colour, and is remarkable from its small size, some specimens weighing only 1¼ lb.; rabbits of this breed make excellent nurses for other and more delicate kinds.^[256]

Certain characters are remarkably fluctuating, or are very feebly transmitted by domestic rabbits: thus, one breeder tells me that with the smaller kinds he has hardly ever raised a whole litter of the same colour: with the large lop-eared breeds "it is impossible," says a great judge,^[257] "to breed true to colour, but by judicious crossing a great deal may be done towards it. The fancier should know how his does are bred, that is, the colour of their parents." Nevertheless, certain colours, as we shall presently see, are transmitted truly. The dewlap is not strictly inherited. Lop-eared rabbits, with their ears hanging flat down on each side of the face, do not transmit this character at all truly. Mr. Delamer remarks that, "with fancy rabbits, when both the parents are perfectly formed, have model ears, and are handsomely marked, their progeny do not invariably turn out the same." When one parent, or even both, are oar-laps, that is, have their ears sticking out at right angles, or when one parent or both are half-lops, that is, have only one ear dependent, there is nearly as good a chance of the progeny having both ears full-lop, as if both parents had been thus characterized. But I am informed, if both parents have upright ears, there is hardly a chance of a full-lop. In some half-lops the ear that hangs down is broader and longer than the upright ear;^[258] so that we have the unusual case of a want of symmetry on the two sides. This difference in the position and size of the two ears probably indicates that the lopping of the ear results {108}from its great length and weight, favoured no doubt by the weakness of the muscles consequent on disuse. Anderson^[259] mentions a breed having only a single ear; and Professor Gervais another breed which is destitute of ears.

Fig. 5.—Half-lop Rabbit. (Copied from E. S. Delamer's work.)

The origin of the Himalayan breed (sometimes called Chinese, or Polish, or Russian) is so curious, both in itself, and as throwing some light on the complex laws of inheritance, that it is worth giving in detail. These pretty rabbits are white, except their ears, nose, all four feet, and the upper side of tail, which are all brownish-black; but as they have red eyes, they may be considered as albinoes. I have received several accounts of their breeding perfectly true. From their symmetrical marks, they were at first ranked as specifically distinct, and were provisionally named L. nigripes^[260] Some good observers thought that they could detect a difference in their habits, and stoutly maintained that they formed a new species. Their origin is now well known. A writer, in 1857,^[261] stated that he had produced Himalayan rabbits in the following manner. But it is first necessary briefly to describe two other breeds: silver-greys or silver-sprigs generally have black heads and legs, and their fine grey fur is interspersed with numerous black and white long hairs. {109}They breed perfectly true, and have long been kept in warrens. When they escape and cross with common rabbits, the product, as I hear from Mr. Wyrley Birch, of Wretham Hall, is not a mixture of the two colours, but about half take after the one parent, and the other half after the other parent. Secondly, chinchillas or tame silver-greys (I will use the former name) have short, paler, mouse or slate-coloured fur, interspersed with long, blackish, slate-coloured, and white hairs.^[262] These rabbits breed perfectly true. Now, the writer above referred to had a breed of chinchillas which had been crossed with the common black rabbit, and their offspring were either blacks or chinchillas. These latter were again crossed with other chinchillas (which had also been crossed with silver-greys), and from this complicated cross Himalayan rabbits were raised. From these and other similar statements, Mr. Bartlett^[263] was led to make a careful trial in the Zoological Gardens, and he found that by simply crossing silver-greys with chinchillas he could always produce some few Himalayans; and the latter, notwithstanding their sudden origin, if kept separate, bred perfectly true.

The Himalayans, when first born, are quite white, and are then true albinoes; but in the course of a few months they gradually assume their dark ears, nose, feet, and tail. Occasionally, however, as I am informed by Mr. W. A. Wooler and the Rev. W. D. Fox, the young are born of a very pale grey colour, and specimens of such fur were sent me by the former gentleman. The grey tint, however, disappears as the animal comes to maturity. So that with these Himalayans there is a tendency, strictly confined to early youth, to revert to the colour of the adult silver-grey parent-stock. Silver-greys and chinchillas, on the other hand, present a remarkable contrast in their colour whilst quite young, for they are born perfectly black, but soon assume their characteristic grey or silver tints. The same thing occurs with grey horses, which, as long as they are foals, are generally of a nearly black colour, but soon become grey, and get whiter and whiter as they grow older. Hence the usual rule is that Himalayans are born white and afterwards become in certain parts of their bodies dark-coloured; whilst {110}silver-greys are born black and afterwards become sprinkled with white. Exceptions, however, and of a directly opposite nature, occasionally occur in both cases. For young silver-greys are sometimes born in warrens, as I hear from Mr. W. Birch, of a cream-colour, but these young animals ultimately become black, The Himalayans, on the other hand, sometimes produce, as is stated by an experienced amateur,^[264] a single black young one in a litter; but such, before two months elapse, become perfectly white.

To sum up the whole curious case: wild silver-greys may be considered as black rabbits which become grey at an early period of life. When they are crossed with common rabbits, the offspring are said not to have blended colours, but to take after either parent; and in this respect they resemble black and albino varieties of most quadrupeds, which often transmit their colours in this same manner. When they are crossed with chinchillas, that is, with a paler sub-variety, the young are at first pure albinoes, but soon become dark-coloured in certain parts of their bodies, and are then called Himalayans. The young Himalayans, however, are sometimes at first either pale grey or completely black, in either case changing after a time to white. In a future chapter I shall advance a large body of facts showing that, when two varieties are crossed both of which differ in colour from their parent-stock, there is a strong tendency in the young to revert to the aboriginal colour; and what is very remarkable, this reversion occasionally supervenes, not before birth, but during the growth of the animal. Hence, if it could be shown that silver-greys and chinchillas were the offspring of a cross between a black and albino variety with the colours intimately blended—a supposition in itself not improbable, and supported by the circumstance of silver-greys in warrens sometimes producing creamy-white young, which ultimately become black—then all the above-given paradoxical facts on the changes of colour in silver-greys and in their descendants the Himalayans would come under the law of reversion, supervening at different periods of growth and in different degrees, either to the original black or to the original albino parent-variety.

{111}

It is, also, remarkable that Himalayans, though produced so suddenly, breed true. But as, whilst young, they are albinoes, the case falls under a very general rule; for albinism is well known to be strongly inherited, as with white mice and many other quadrupeds, and even with white flowers. But why, it may be asked, do the ears, tail, nose, and feet, and no other part of the body, revert to a black colour? This apparently depends on a law, which generally holds good, namely, that characters common to many species of a genus—and this, in fact, implies long inheritance in common from the ancient progenitor of the genus—are found to resist variation, or to reappear if lost, more persistently than the characters which are confined to the separate species. Now, in the genus Lepus, a large majority of the species have their ears and the upper surface of the tail tinted black; but the persistence of these marks is best seen in those species which in winter become white: thus, in Scotland the L. variabilis^[265] in its winter dress has a shade of colour on its nose, and the tips of its ears are black: in the L. tibetanus the ears are black, the upper surface of the tail greyish-black, and the soles of the feet brown: in L. glacialis the winter fur is pure white, except the soles of the feet and the points of the ears. Even in the variously-coloured fancy rabbits we may often observe a tendency in these same parts to be more darkly tinted than the rest of the body. Thus, as it seems to me, the appearance of the several coloured marks on the Himalayan rabbit, as it grows old, is rendered intelligible. I may add a nearly analogous case: fancy rabbits very often have a white star on their foreheads; and the common English hare, whilst young, generally has, as I have myself observed, a similar white star on its forehead.

When variously coloured rabbits are set free in Europe, and are thus placed under their natural conditions, they generally revert to the aboriginal grey colour; this may be in part due to the tendency in all crossed animals, as lately observed, to revert to their primordial state. But this tendency does not always prevail; thus silver-grey rabbits are kept in warrens, and remain true though living almost in a state of nature; but a {112}warren must not be stocked with both silver-greys and common rabbits; otherwise "in a few years there will be none but common greys surviving."^[266] When rabbits run wild in foreign countries, under different conditions of life, they by no means always revert to their aboriginal colour. In Jamaica the feral rabbits are described as "slate-coloured, deeply tinted with sprinklings of white on the neck, on the shoulders, and on the back; softening off to blue-white under the breast and belly."^[267] But in this tropical island the conditions were not favourable to their increase, and they never spread widely; and, as I hear from Mr. R. Hill, owing to a great fire which occurred in the woods, they have now become extinct. Rabbits during many years have run wild in the Falkland Islands; they are abundant in certain parts, but do not spread extensively. Most of them are of the common grey colour; a few, as I am informed by Admiral Sulivan, are hare-coloured, and many are black, often with nearly symmetrical white marks on their faces. Hence, M. Lesson described the black variety as a distinct species, under the name of Lepus magellanicus, but this, as I have elsewhere shown, is an error.^[268] Within recent times the sealers have stocked some of the small outlying islets in the Falkland group with rabbits; and on Pebble Islet, as I hear from Admiral Sulivan, a large proportion are hare-coloured, whereas on Rabbit Islet a large proportion are of a bluish colour which is not elsewhere seen. How the rabbits were coloured which were turned out on these islets is not known.

The rabbits which have become feral on the island of Porto Santo, near Madeira, deserve a fuller account. In 1418 or 1419, J. Gonzales Zarco^[269] happened to have a female rabbit on board which had produced young during the voyage, and he turned them all out on the island. These animals soon increased so {113}rapidly, that they became a nuisance, and actually caused the abandonment of the settlement. Thirty-seven years subsequently, Cada Mosto describes them as innumerable; nor is this surprising, as the island was not inhabited by any beast of prey or by any terrestrial mammal. We do not know the character of the mother-rabbit; but we have every reason to believe that it was the common domesticated kind. The Spanish peninsula, whence Zarco sailed, is known to have abounded with the common wild species at the most remote historical period. As these rabbits were taken on board for food, it is improbable that they should have been of any peculiar breed. That the breed was well domesticated is shown by the doe having littered during the voyage. Mr. Wollaston, at my request, brought home two of these feral rabbits in spirits of wine; and, subsequently, Mr. W. Haywood sent to me three more specimens in brine, and two alive. These seven specimens, though caught at different periods, closely resembled each other. They were full grown, as shown by the state of their bones. Although the conditions of life in Porto Santo are evidently highly favourable to rabbits, as proved by their extraordinarily rapid increase, yet they differ conspicuously in their small size from the wild English rabbit. Four English rabbits, measured from the incisors to the anus, varied between 17 and 17¾ inches in length; whilst two of the Porto Santo rabbits were only 14½ and 15 inches in length. But the decrease in size is best shown by weight; four wild English rabbits averaged 3 lb. 5 oz., whilst one of the Porto Santo rabbits, which had lived for four years in the Zoological Gardens, but had become thin, weighed only 1 lb. 9 oz. A fairer test is afforded by the comparison of the well-cleaned limb-bones of a P. Santo rabbit killed on the island with the same bones of a wild English rabbit of average size, and they differed in the proportion of rather less than five to nine. So that the Porto Santo rabbits have decreased nearly three inches in length, and almost half in weight of body.^[270] The head has not decreased in length {114}proportionally with the body; and the capacity of the brain-case is, as we shall hereafter see, singularly variable. I prepared four skulls, and these resembled each other more closely than do generally the skulls of wild English rabbits; but the only difference in structure which they presented was that the supra-orbital processes of the frontal bones were narrower.

In colour the Porto Santo rabbit differs considerably from the common rabbit; the upper surface is redder, and is rarely interspersed with any black or black-tipped hairs. The throat and certain parts of the under surface, instead of being pure white, are generally pale grey or leaden colour. But the most remarkable difference is in the ears and tail; I have examined many fresh English rabbits, and the large collection of skins in the British Museum from various countries, and all have the upper surface of the tail and the tips of the ears clothed with blackish-grey fur; and this is given in most works as one of the specific characters of the rabbit. Now in the seven Porto Santo rabbits the upper surface of the tail was reddish-brown, and the tips of the ears had no trace of the black edging. But here we meet with a singular circumstance: in June, 1861, I examined two of these rabbits recently sent to the Zoological Gardens, and their tails and ears were coloured as just described; but when one of their dead bodies was sent to me in February, 1865, the ears were plainly edged, and the upper surface of the tail was covered, with blackish-grey fur, and the whole body was much less red; so that under the English climate this individual rabbit had recovered the proper colour of its fur in rather less than four years!

The two little Porto Santo rabbits, whilst alive in the Zoological Gardens, had a remarkably different appearance from the common kind. They were extraordinarily wild and active, so that many persons exclaimed on seeing them that they were more like large rats than rabbits. They were nocturnal to an unusual degree in their habits, and their wildness was never in the least subdued; so that the superintendent, Mr. Bartlett, assured me that he had never had a wilder animal under his charge. This is a singular fact, considering that they are descended from a domesticated breed; I was so much surprised at it, that I requested Mr. Haywood to make inquiries on the spot, {115}whether they were much hunted by the inhabitants, or persecuted by hawks, or cats, or other animals; but this is not the case, and no cause can be assigned for their wildness. They live on the central, higher rocky land and near the sea-cliffs, and, being exceedingly shy and timid, seldom appear in the lower and cultivated districts. They are said to produce from four to six young at a birth, and their breeding season is in July and August. Lastly, and this is a highly remarkable fact, Mr. Bartlett could never succeed in getting these two rabbits, which were both males, to associate or breed with the females of several breeds which were repeatedly placed with them.

If the history of these Porto Santo rabbits had not been known, most naturalists, on observing their much reduced size, their reddish colour above and grey beneath, with neither tail nor ears tipped with black, would have ranked them as a distinct species. They would have been strongly confirmed in this view by seeing them alive in the Zoological Gardens, and hearing that they refused to couple with other rabbits. Yet this rabbit, which there can be little doubt would thus have been ranked as a distinct species, has certainly originated since the year 1420. Finally, from the three cases of the rabbits which have run wild in Porto Santo, Jamaica, and the Falkland Islands, we see that these animals do not, under new conditions of life, revert to or retain their aboriginal character, as is so generally asserted to be the case by most authors.

Osteological Characters.

When we remember, on the one hand, how frequently it is stated that important parts of the structure never vary; and, on the other hand, on what small differences in the skeleton, fossil species have often been founded, the variability of the skull and of some other bones in the domesticated rabbit well deserves attention. It must not be supposed that the more important differences immediately to be described strictly characterise any one breed; all that can be said is, that they are generally present in certain breeds. We should bear in mind that selection has not been applied to fix any character in the skeleton, and that the animals have not had to support themselves under {116}uniform habits of life. We cannot account for most of the differences in the skeleton; but we shall see that the increased size of the body, due to careful nurture and continued selection, has affected the head in a particular manner. Even the elongation and lopping of the ears have influenced in a small degree the form of the whole skull. The want of exercise has apparently modified the proportional length of the limbs in comparison with the body.

Fig. 7.—Skull of large Lop-eared Rabbit, of natural size.

Fig. 6.—Skull of Wild Rabbit, of natural size.

As a standard of comparison, I prepared skeletons of two wild rabbits from Kent, one from the Shetland Islands, and one from Antrim in Ireland. As all the bones in these four specimens from such distant localities closely resembled each other, presenting scarcely any appreciable difference, it may be concluded that the bones of the wild rabbit are generally uniform in character.

Skull.—I have carefully examined skulls of ten large lop-eared fancy rabbits, and of five common domestic rabbits, which latter differ from the lop-eared only in not having such large bodies or ears, yet both larger than in the wild rabbit. First for the ten lop-eared rabbits: in all these the skull is remarkably elongated in comparison with its breadth. In a wild rabbit the length was 3.15 inches, in a large fancy rabbit 4.30; whilst the breadth of the cranium enclosing the brain was in both almost exactly the same. Even by taking as the standard of comparison the widest part of the zygomatic arch, the skulls of the lop-eared are proportionally to their breadth three-quarters of an inch too long. The depth of the head has increased almost in the same proportion with the length; it is the breadth alone which has not increased. The parietal and occipital bones enclosing the brain are less arched, both in a longitudinal and transverse line, than in the wild rabbit, so that the shape of the cranium is somewhat different. The surface is rougher, less cleanly sculptured, and the lines of sutures are more prominent.

Although the skulls of the large lop-eared rabbits in comparison with those of the wild rabbit are much elongated relatively to their breadth, yet, relatively to the size of body, they are far from elongated. The lop-eared rabbits which I examined were, though not fat, more than twice as heavy as the wild specimens; but the skull was very far from being twice as long. Even if we take the fairer standard of the length of body, from the nose to the anus, the skull is not on an average as long as it ought to be by a third of an inch. In the small feral P. Santo rabbit, on the other hand, the head relatively to the length of body is about a quarter of an inch too long.

Fig. 8.—Part of Zygomatic Arch, showing the projecting end of the malar bone and the auditory meatus: of natural size. Upper figure, Wild Rabbit. Lower figure, Lop-eared, hare-coloured Rabbit.

This elongation of the skull relatively to its breadth, I find a universal character, not only with the large lop-eared rabbits, but in all the artificial breeds; as is well seen in the skull of the Angora. I was at first much surprised at the fact, and could not imagine why domestication should produce this uniform result; but the explanation seems to lie in the circumstance that during a number of generations the artificial races have been closely confined, and have had little occasion to exert either their senses, or intellect, or voluntary muscles; consequently the brain, as {117}we shall presently more fully see, has not increased relatively with the size of body. As the brain has not increased, the bony case enclosing it has not increased, and this has evidently affected through correlation the breadth of the entire skull from end to end.

In all the skulls of the large lop-eared rabbits, the supra-orbital plates or processes of the frontal bones ere much broader than in the wild rabbit, and they generally project more upwards. In the zygomatic arch the posterior or projecting point of the malar-bone is broader and blunter; and in the specimen, fig. 8, it is so in a remarkable degree. This point approaches nearer to the auditory meatus than in the wild rabbit, as may be best seen in fig. 8; but this circumstance mainly depends on the changed direction of the meatus. The inter-parietal bone (see fig. 9) differs much in shape in the several skulls; generally it is more oval, or has a greater width in the line of the longitudinal axis of the skull, than in the wild rabbit. The {118}posterior margin of "the square raised platform" ^[271] of the occiput, instead of being truncated, or projecting slightly as in the wild rabbit, is in most lop-eared rabbits pointed, as in fig. 9, C. The paramastoids relatively to the size of the skull are generally much thicker than in the wild rabbit.

The occipital foramen (fig. 10) presents some remarkable differences: in the wild rabbit, the lower edge between the condyles is considerably and almost angularly hollowed out, and the upper edge is deeply and squarely notched; hence the longitudinal axis exceeds the transverse axis. In the skulls of the lop-eared rabbits the transverse axis exceeds the longitudinal; for in none of these skulls was the lower edge between the condyles so deeply hollowed out; in five of them there was no upper square notch, in three there was a trace of the notch, and in two alone it was well developed. These differences in the shape of the foramen are remarkable, considering that it gives passage to so important a structure as the spinal marrow, though apparently the outline of the latter is not affected by the shape of the passage.

Fig. 9.—Posterior end of Skull, of natural size, showing the inter-parietal bone. A. Wild Rabbit. B. Feral Rabbit from island of P. Santo, near Madeira. C. Large Lop-eared Rabbit.

Fig. 10.—Occipital Foramen, of natural size, in—A. Wild Rabbit; B. Large Lop-eared Rabbit.

In all the skulls of the large lop-eared rabbits, the bony auditory meatus is conspicuously larger than in the wild rabbit. In a skull 4.3 inches in length, and which barely exceeded in breadth the skull of a wild rabbit (which was 3.15 inches in length), the longer diameter of the meatus was exactly twice as great. The orifice is more compressed, and its margin on the side nearest the skull stands up higher than the outer side. The whole meatus is directed more forwards. As in breeding lop-eared rabbits the length of the ears, and their consequent lopping and lying flat on the face, are the chief points of excellence, there can hardly be a doubt that the great change in the size, form, and direction of the bony meatus, relatively to this same part in the wild rabbit, is due to the continued selection of individuals having {119}larger and larger ears. The influence of the external ear on the bony meatus is well shown in the skulls (I have examined three) of half-lops (see fig. 5), in which one ear stands upright, and the other and longer ear hangs down; for in these skulls there was a plain difference in the form and direction of the bony meatus on the two sides. But it is a much more interesting fact, that the changed direction and increased size of the bony meatus have slightly affected on the same side the structure of the whole skull. I here give a drawing of the skull of a half-lop; and it may be observed that the suture between the parietal and frontal bones does not run strictly at right angles to the longitudinal axis of the skull; the left frontal bone projects beyond the right one; both the posterior and anterior margins of the left zygomatic arch on the side of the lopping ear stand a little in advance of the corresponding bones on the opposite side. Even the lower jaw is affected, and the condyles are not quite symmetrical, that on the left standing a little in advance of that on the right. This seems to me a remarkable case of correlation of growth. Who would have surmised that by keeping an animal during many generations under confinement, and so leading to the disuse of the muscles of the ears, and by continually selecting individuals with the longest and largest ears, he would thus indirectly have affected almost every suture in the skull and the form of the lower jaw!

Fig. 11.—Skull, of natural size, of Half-lop Rabbit, showing the different direction of the auditory meatus on the two sides, and the consequent general distortion of the skull. The left ear of the animal (or right side of figure) lopped forwards.

In the large lop-eared rabbits the only difference in the lower jaw, in comparison with that of the wild rabbit, is that the posterior margin of the ascending ramus is broader and more inflected. The teeth in neither jaw present any difference, except that the small incisors, beneath the large ones, are proportionally a little longer. The molar teeth have increased in size proportionally with the increased width of the skull, measured across the zygomatic arch, and not proportionally with its increased length. The inner line of the sockets of the molar teeth in the upper jaw of the wild rabbit forms a perfectly straight line; but in {120}some of the largest skulls of the lop-eared this line was plainly bowed inwards. In one specimen there was an additional molar tooth on each side of the upper jaw, between the molars and premolars; but these two teeth did not correspond in size; and as no rodent has seven molars, this is merely a monstrosity, though a curious one.

The five other skulls of common domestic rabbits, some of which approach in size the above-described largest skulls, whilst the others exceed but little those of the wild rabbit, are only worth notice as presenting a perfect gradation in all the above-specified differences between the skulls of the largest lop-eared and wild rabbits. In all, however, the supra-orbital plates are rather larger, and in all the auditory meatus is larger, in conformity with the increased size of the external ears, than in the wild rabbit. The lower notch in the occipital foramen in some was not so deep as in the wild, but in all five skulls the upper notch was well developed.

The skull of the Angora rabbit, like the latter five skulls, is intermediate in general proportions, and in most other characters, between those of the largest lop-eared and wild rabbits. It presents only one singular character: though considerably longer than the skull of the wild, the breadth measured within the posterior supra-orbital fissures is nearly a third less than in the wild. The skulls of the silver-grey, and chinchilla and Himalayan rabbits are more elongated than in the wild, with broader supra-orbital plates, but differ little in any other respect, excepting that the upper and lower notches of the occipital foramen are not so deep or so well developed. The skull of the Moscow rabbit scarcely differs in any respect from that of the wild rabbit. In the Porto Santo feral rabbits the supra-orbital plates are generally narrower and more pointed than in our wild rabbits.

As some of the largest lop-eared rabbits of which I prepared skeletons were coloured almost like hares, and as these latter animals and rabbits have, as it is affirmed, been recently crossed in France, it might be thought that some of the above-described characters had been derived from a cross at a remote period with the hare. Consequently I examined skulls of the hare, but no light could thus be thrown on the peculiarities of the skulls of the larger rabbits. It is, however, an interesting fact, as illustrating the law that varieties of one species often assume the characters of other species of the same genus, that I found, on comparing the skulls of ten species of hares in the British Museum, that they differed from each other chiefly in the very same points in which domestic rabbits vary,—namely, in general proportions, in the form and size of the supra-orbital plates, in the form of the free end of the malar bone, and in the line of suture separating the occipital and frontal bones. Moreover two eminently variable characters in the domestic rabbit, namely, the outline of the occipital foramen and the shape of the "raised platform" of the occiput, were likewise variable in two instances in the same species of hare.

Vertebræ.—The number is uniform in all the skeletons which I have examined, with two exceptions, namely, in one of the small feral Porto Santo rabbits and in one of the largest lop-eared kinds; both of these had as usual seven cervical, twelve dorsal with ribs, but, instead of seven lumbar, both had eight lumbar vertebræ. This is remarkable, as Gervais gives {121}seven as the number for the whole genus Lepus. The caudal vertebræ apparently differ by two or three, but I did not attend to them, and they are difficult to count with certainty.

In the first cervical vertebra, or atlas, the anterior margin of the neural arch varies a little in wild specimens, being either nearly smooth, or furnished with a small supra-median atlantoid process; I have figured a specimen with the largest process (a) which I have seen; but it will be observed how inferior this is in size and different in shape to that in a large lop-eared rabbit. In the latter, the infra-median process (b) is also proportionally much thicker and longer. The alæ are a little squarer in outline.

Fig. 12.—Atlas Vertebræ, of natural size; inferior surface viewed obliquely. Upper figure, Wild Rabbit. Lower figure, Hare-coloured, large, Lop-eared Rabbit. a, supra-median, atlantoid process; b, infra-median process.

Third cervical vertebra.—In the wild rabbit (fig. 13, A a) this vertebra, viewed on the inferior surface, has a transverse process, which is directed obliquely backwards, and consists of a single pointed bar; in the fourth vertebra this process is slightly forked in the middle. In the large lop-eared rabbits this process (B a) is forked in the third vertebra, as in the fourth of the wild rabbit. But the third cervical vertebræ of the wild and lop-eared (A b, B b) rabbits differ more conspicuously when their anterior articular surfaces are compared; for the extremities of the antero-dorsal processes in the wild rabbit are simply rounded, whilst in the lop-eared they are trifid, with a deep central pit. The canal for the spinal marrow in the lop-eared (B b) is more elongated in a transverse direction than in the wild rabbit; and the passages for the arteries are of a slightly different shape. These several differences in this vertebra seem to me well deserving attention.

Fig. 13.—Third Cervical Vertebra, of natural size, of—A. Wild Rabbit; B. Hare-coloured, large, Lop-eared Rabbit. a, a, inferior surface; b, b, anterior articular surfaces.

First dorsal vertebra.—Its neural spine varies in length in the wild rabbit; being sometimes very short, but generally more than half as long as that of the second dorsal; but I have seen it in two large lop-eared rabbits three-fourths of the length of that of the second dorsal vertebra.

Ninth and tenth dorsal vertebræ.—In the wild rabbit the neural spine of the ninth vertebra is just perceptibly thicker than that of the eighth; and {122}the neural spine of the tenth is plainly thicker and shorter than those of all the anterior vertebræ. In the large lop-cared rabbits the neural spines of the tenth, ninth, eighth, and even in a slight degree that of the seventh vertebra, are very much thicker, and of somewhat different shape, in comparison with those of the wild rabbit. So that this part of the vertebral column differs considerably in appearance from the same part in the wild rabbit, and closely resembles in an interesting manner these same vertebræ in some species of hares. In the Angora, Chinchilla, and Himalayan rabbits, the neural spines of the eighth and ninth vertebræ are in a slight degree thicker than in the wild. On the other hand, in one of the feral Porto Santo rabbits, which in most of its characters deviates in an exactly opposite manner to what the large lop-cared rabbits do from the common wild rabbit, the neural spines of the ninth and tenth vertebræ were not at all larger than those of the several anterior vertebræ. In this same Porto Santo specimen there was no trace in the ninth vertebra of the anterior lateral processes (see woodcut 14), which are plainly developed in all British wild rabbits, and still more plainly developed in the large lop-eared rabbits. In a half-wild rabbit from Sandon Park,^[272] a hæmal spine was moderately well developed on the under side of the twelfth dorsal vertebra, and I have seen this in no other specimen.

Fig. 14.—Dorsal Vertebræ, from sixth to tenth inclusive, of natural size, viewed laterally. A. Wild Rabbit. B. Large, Hare-coloured, so called Spanish Rabbit.

Fig. 15.—Terminal bone of Sternum, of natural size. A. Wild Rabbit. B. Hare-coloured, Lop-eared Rabbit. C. Hare-coloured, Spanish Rabbit. (N.B. The left-hand angle of the upper articular extremity of B was broken, and has been accidentally thus represented.)

Lumbar vertebræ.—I have stated that in two cases there were eight instead of seven lumbar vertebræ. The third lumbar vertebra in one skeleton of a wild British rabbit, and in one of the Porto Santo feral rabbits, had a hæmal spine; whilst in four skeletons of large lop-eared rabbits, and in the Himalayan rabbit, this same vertebra had a well-developed hæmal spine.

Pelvis.—In four wild specimens this bone was almost absolutely identical in shape; but in several domesticated breeds shades of differences {123}could be distinguished. In the large lop-eared rabbits the whole upper part of the ilium is straighter, or less splayed outwards, than in the wild rabbit; and the tuberosity on the inner lip of the anterior and upper part of the ilium is proportionally more prominent.

Sternum.—The posterior end of the posterior sternal bone in the wild rabbit (fig. 15, A) is thin and slightly enlarged; in some of the large lop-eared rabbits (B) it is much more enlarged towards the extremity; whilst in other specimens (C) it keeps nearly of the same breadth from end to end, but is much thicker at the extremity.

Fig. 16.—Acromion of Scapula, of natural size. A. Wild Rabbit. B, C, D; Large, Lop-eared Rabbits.

Scapula.—The acromion sends out a rectangular bar, ending in an oblique knob, which latter in the wild rabbit (fig. 16, A) varies a little in shape and size, as does the apex of the acromion in sharpness, and the part just below the rectangular bar in breadth. But the variations in these respects in the wild rabbit are very slight; whilst in the large lop-eared rabbits they are considerable. Thus in some specimens (B) the oblique terminal knob is developed into a short bar, forming an obtuse angle with the rectangular bar. In another specimen (C) these two unequal bars form nearly a straight line. The apex of the acromion varies much in breadth and sharpness, as may be seen by comparing figs. B, C, and D.

Limbs.—In these I could detect no variation; but the bones of the feet were too troublesome to compare with much care.

I have now described all the differences in the skeletons which I have observed. It is impossible not to be struck with the high degree of variability or plasticity of many of the bones. We see how erroneous the often-repeated statement is, that only the crests of the bones which give attachment to muscles vary in shape, and that only parts of slight importance {124}become modified under domestication. No one will say, for instance, that the occipital foramen, or the atlas, or the third cervical vertebra is a part of slight importance. If the several vertebræ of the wild and lop-eared rabbits, of which figures have been given, had been found fossil, palæontologists would have declared without hesitation that they had belonged to distinct species.

The effects of the use and disuse of parts.—In the large lop-eared rabbits the relative proportional lengths of the bones of the same leg, and of the front and hind legs compared with each other, have remained nearly the same as in the wild rabbit; but in weight, the bones of the hind legs apparently have not increased in due proportion with the front legs. The weight of the whole body in the large rabbits examined by me was from twice to twice and a half as great as that of the wild rabbit; and the weight of the bones of the front and hind limbs taken together (excluding the feet, on account of the difficulty of perfectly cleaning so many small bones) has increased in the large lop-eared rabbits in nearly the same proportion; consequently in due proportion to the weight of body which they have to support. If we take the length of the body as the standard of comparison, the limbs of the large rabbits have not increased in length in due proportion by one inch, or by one inch and a half. Again, if we take as the standard of comparison the length of the skull, which, as we have before seen, has not increased in length in due proportion to the length of body, the limbs will be found to be, proportionally with those of the wild rabbit, from half to three-quarters of an inch too short. Hence, whatever standard of comparison be taken, the limb-bones of the large lop-eared rabbits have not increased in length, though they have in weight, in full proportion to the other parts of the frame; and this, I presume, may be accounted for by the inactive life which during many generations they have spent. Nor has the scapula increased in length in due proportion to the increased length of the body.

The capacity of the osseous case of the brain is a more interesting point, to which I was led to attend by finding, as previously stated, that with all domesticated rabbits the length of the skull relatively to its breadth has greatly increased in comparison with that of the wild rabbit. If we had possessed a large number of domesticated rabbits of nearly the same size with the wild rabbit, it would have been a simple task to have measured and compared the capacities of their skulls. But this is not the case; almost all the domestic breeds have larger bodies than wild rabbits, and the lop-eared kinds are more than double their weight. As a small animal has to exert its senses, intellect, and instincts equally with a large animal, we ought not by any means to expect an animal twice or thrice as large as another to have a brain of double or treble the size.^[273] Now, after weighing {125}the bodies of four wild rabbits, and of four large but not fattened lop-eared rabbits, I find that on an average the wild are to the lop-eared in weight as 1 to 2.47; in average length of body as 1 to 1.41; whilst in capacity of skull (measured as hereafter to be described) they are only as 1 to 1.15. Hence we see that the capacity of the skull, and consequently the size of the brain, has increased but little, relatively to the increased size of the body; and this fact explains the narrowness of the skull relatively to its length in all domestic rabbits.

In the upper half of the following table I have given the measurements of the skulls of ten wild rabbits; and in the lower half of eleven thoroughly domesticated kinds. As these rabbits differ so greatly in size, it is necessary to have some standard by which to compare the capacities of their skulls. I have selected the length of skull as the best standard, for in the larger rabbits it has not, as already stated, increased in length so much as the body; but as the skull, like every other part, varies in length, neither it nor any other part affords a perfect standard.

In the first column of figures the extreme length of the skull is given in inches and decimals. I am aware that these measurements pretend to greater accuracy than is possible; but I have found it the least trouble to record the exact length which the compass gave. The second and third columns give the length and weight of body, whenever these measurements have been made. The fourth column gives the capacity of the skull by the weight of small shot with which the skulls had been filled; but it is not pretended that these weights are accurate within a few grains. In the fifth column the capacity is given which the skull ought to have had by calculation, according to the length of skull, in comparison with that of the wild rabbit No. 1; in the sixth column the difference between the actual and calculated capacities, and in the seventh the percentage of increase or decrease, are given. For instance, as the wild rabbit No. 5 has a shorter and lighter body than the wild rabbit No. 1, we might have expected that its skull would have had less capacity; the actual capacity, as expressed by the weight of shot, is 875 grains, which is 97 grains less than that of the first rabbit. But comparing these two rabbits by the length of their skulls, we see that in No. 1 the skull is 3.15 inches in length, and in No. 5 2.96 inches in length; according to this ratio, the brain of No. 5 ought to have had a capacity of 913 grains of shot, which is above the actual capacity, but only by 38 grains. Or, to put the case in another way (as in column VII), the brain of this small rabbit, No. 5, for every 100 grains of weight is only 4 per cent. too light,—that is, it ought, according to the standard rabbit No. 1, to have been 4 per cent. heavier. I have taken the rabbit No. 1 as the standard of comparison because, of the skulls having a full average length, this has the least capacity; so that it is the least favourable to the result which I wish to show, namely, that the brain in all long-domesticated rabbits has decreased in size, either actually, or relatively to the length of the head and body, in comparison with the brain of the wild rabbit. Had I taken the Irish rabbit, No. 3, as the standard, the following results would have been somewhat more striking.

Turning to the Table: the first four wild rabbits have skulls of the same length, and these differ but little in capacity. The Sandon rabbit {126}(No. 4) is interesting, as, though now wild, it is known to be descended from a domesticated breed, as is still shown by its peculiar colouring and longer body; nevertheless the skull has recovered its normal length and full capacity. The next three rabbits are wild, but of small size, and they all have skulls with slightly lessened capacities. The three Porto Santo feral rabbits (Nos. 8 to 10) offer a perplexing case; their bodies are greatly reduced in size, as in a lesser degree are their skulls in length and in actual capacity, in comparison with the skulls of wild English rabbits. But when we compare the capacities of the skull in the three Porto Santo rabbits, we observe a surprising difference, which does not stand in any relation to the slight difference in the length of their skulls, nor, as I believe, to any difference in the size of their bodies; but I neglected to weigh separately their bodies. I can hardly suppose that the medullary matter of the brain in these three rabbits, living under similar conditions, can differ as much as is indicated by the proportional difference of capacity in their skulls; nor do I know whether it is possible that one brain may contain considerably more fluid than another. Hence I can throw no light on this case.

Looking to the lower half of the Table, which gives the measurements of domesticated rabbits, we see that in all the capacity of the skull is less, but in very various degrees, than might have been anticipated according to the length of their skulls, relatively to that of the wild rabbit No. 1. In line 22 the average measurements of seven large lop-eared rabbits are given. Now the question arises, has the average capacity of the skull in these seven large rabbits increased as much as might have been expected from their greatly increased size of body. We may endeavour to answer this question in two ways: in the upper half of the Table we have measurements of the skulls of six small wild rabbits (Nos. 5 to 10), and we find that on an average the skulls are in length .18 of an inch shorter, and in capacity 91 grains less, than the average length and capacity of the three first wild rabbits on the list. The seven large lop-cared rabbits, on an average, have skulls 4.11 inches in length, and 1136 grains in capacity; so that these skulls have increased in length more than five times as much as the skulls of the six small wild rabbits have decreased in length; hence we might have expected that the skulls of the large lop-eared rabbits would have increased in capacity five times as much as the skulls of the six small rabbits have decreased in capacity; and this would have given an average increased capacity of 455 grains, whilst the real average increase is only 155 grains. Again, the large lop-eared rabbits have bodies of nearly the same weight and size as the common hare, but their heads are longer; consequently, if the lop-eared rabbits had been wild, it might have been expected that their skulls would have had nearly the same capacity as that of the skull of the hare. But this is far from being the case; for the average capacity of the two hare-skulls (Nos. 23, 24) is so much larger than the average capacity of the seven lop-cared skulls, that the latter would have to be increased 21 per cent. to come up to the standard of the hare.^[274]

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I have previously remarked that, if we had possessed many domestic rabbits of the same average size with the wild rabbit, it would have been easy to compare the capacity of their skulls. Now the Himalayan, Moscow, and Angora rabbits (Nos. 11, 12, 13 of Table) are only a little larger in body, and have skulls only a little longer, than the wild animal, and we see that the actual capacity of their skulls is less than in the wild animal, and considerably less by calculation (column 7), according to the difference in the length of their skulls. The narrowness of the brain-case in these three rabbits could be plainly seen and proved by external measurement. The Chinchilla rabbit (No. 14) is a considerably larger animal than the wild rabbit, yet the capacity of its skull only slightly exceeds that of the wild rabbit. The Angora rabbit, No. 13, offers the most remarkable case; this animal in its pure white colour and length of silky fur bears the stamp of long domesticity. It has a considerably longer head and body than the wild rabbit, but the actual capacity of its skull is less than that of even the little wild Porto Santo rabbits. By the standard of the length of skull the capacity (see column 7) is only half of what it ought to have been! I kept this individual animal alive, and it was not unhealthy nor idiotic. This case of the Angora rabbit so much surprised me, that I repeated all the measurements and found them correct. I have also compared the capacity of the skull of the Angora with that of the wild rabbit by other standards, namely, by the length and weight of the body, and by the weight of the limb-bones; but by all these standards the brain appears to be much too small, though in a less degree when the standard of the limb-bones was used; and this latter circumstance may probably be accounted for by the Limbs of this anciently domesticated breed having become much reduced in weight, from its long-continued inactive life. Hence I infer that in the Angora breed, which is said to differ from other breeds in being quieter and more social, the capacity of the skull has really undergone a remarkable amount of reduction.

From the several facts above given,—namely, firstly, that the actual capacity of the skull in the Himalayan, Moscow, and Angora breeds, is less than in the wild rabbit, though they are in all their dimensions rather larger animals; secondly, that the capacity of the skull of the large lop-eared rabbits has not been increased in nearly the same ratio as the capacity of the skull of the smaller wild rabbits has been decreased; and thirdly, that the capacity of the skull in these same large lop-eared rabbits is very inferior to that of the hare, an animal of nearly the same {129}size,—I conclude, notwithstanding the remarkable differences in capacity in the skulls of the small P. Santo rabbits, and likewise in the large lop-eared kinds, that in all long-domesticated rabbits the brain has either by no means increased in due proportion with the increased length of the head and increased size of the body, or that it has actually decreased in size, relatively to what would have occurred had these animals lived in a state of nature. When we remember that rabbits, from having been domesticated and closely confined during many generations, cannot have exerted their intellect, instincts, senses, and voluntary movements, either in escaping from various dangers or in searching for food, we may conclude that their brains will have been feebly exercised, and consequently have suffered in development. We thus see that the most important and complicated organ in the whole organization is subject to the law of decrease in size from disuse.

Finally, let us sum up the more important modifications which domestic rabbits have undergone, together with their causes as far as we can obscurely see them. By the supply of abundant and nutritious food, together with little exercise, and by the continued selection of the heaviest individuals, the weight of the larger breeds has been more than doubled. The bones of the limbs have increased in weight (but the hind legs less than the front legs), in due proportion with the increased weight of body; but in length they have not increased in due proportion, and this may have been caused by the want of proper exercise. With the increased size of the body the third cervical vertebra has assumed characters proper to the fourth cervical; and the eighth and ninth dorsal vertebræ have similarly assumed characters proper to the tenth and posterior vertebræ. The skull in the larger breeds has increased in length, but not in due proportion with the increased length of body; the brain has not duly increased in dimensions, or has even actually decreased, and consequently the bony case for the brain has remained narrow, and by correlation has affected the bones of the face and the entire length of the skull. The skull has thus acquired its characteristic narrowness. From unknown causes the supra-orbital processes of the frontal bones and the free end of the malar bones have increased in breadth; and in the larger breeds {130}the occipital foramen is generally much less deeply notched than in wild rabbits. Certain parts of the scapula and the terminal sternal bones have become highly variable in shape. The ears have been increased enormously in length and breadth through continued selection; their weight, conjoined probably with the disuse of their muscles, has caused them to lop downwards; and this has affected the position and form of the bony auditory meatus; and this again, by correlation, the position in a slight degree of almost every bone in the upper part of the skull, and even the position of the condyles of the lower jaw.

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CHAPTER V.

DOMESTIC PIGEONS.

ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS—INDIVIDUAL VARIABILITY—VARIATIONS OF A REMARKABLE NATURE—OSTEOLOGICAL CHARACTERS: SKULL, LOWER JAW, NUMBER OF VERTEBRÆ—CORRELATION OF GROWTH: TONGUE WITH BEAK; EYELIDS AND NOSTRILS WITH WATTLED SKIN—NUMBER OF WING-FEATHERS, AND LENGTH OF WING—COLOUR AND DOWN—WEBBED AND FEATHERED FEET—ON THE EFFECTS OF DISUSE—LENGTH OF FEET IN CORRELATION WITH LENGTH OF BEAK—LENGTH OP STERNUM, SCAPULA, AND FURCULA—LENGTH OF WINGS—SUMMARY ON THE POINTS OF DIFFERENCE IN THE SEVERAL BREEDS

I have been led to study domestic pigeons with particular care, because the evidence that all the domestic races have descended from one known source is far clearer than with any other anciently domesticated animal. Secondly, because many treatises in several languages, some of them old, have been written on the pigeon, so that we are enabled to trace the history of several breeds. And lastly, because, from causes which we can partly understand, the amount of variation has been extraordinarily great. The details will often be tediously minute; but no one who really wants to understand the progress of change in domestic animals will regret this; and no one who has kept pigeons and has marked the great difference between the breeds and the trueness with which most of them propagate their kind, will think this care superfluous. Notwithstanding the clear evidence that all the breeds are the descendants of a single species, I could not persuade myself until some years had passed that the whole amount of difference between them had arisen since man first domesticated the wild rock-pigeon.

I have kept alive all the most distinct breeds, which I could procure in England or from the Continent; and have prepared skeletons of all. I have received skins from Persia, and a large number from India and other quarters of the {132}world.^[275] Since my admission into two of the London pigeon-clubs, I have received the kindest assistance from many of the most eminent amateurs.^[276]

The races of the Pigeon which can be distinguished, and which breed true, are very numerous. MM. Boitard and Corbié^[277] describe in detail 122 kinds; and I could add several European kinds not known to them. In India, judging from the skins sent me, there are many breeds unknown here; and Sir W. Elliot informs me that a collection imported by an Indian merchant into Madras from Cairo and Constantinople included several kinds unknown in India. I have no doubt that there exist considerably above 150 kinds which breed true and have been separately named. But of these the far greater number differ from each other only in unimportant characters. Such differences will be here entirely passed over, and I shall confine myself to the more important points of structure. That many important differences exist we shall presently see. I have looked through the magnificent collection of the Columbidæ in the British Museum, and, with the exception of a few forms (such as the Didunculus, Calænas, Goura, &c), I do not hesitate to {133}affirm that some domestic races of the rock-pigeon differ fully as much from each other in external characters as do the most distinct natural genera. We may look in vain through the 288 known species^[278] for a beak so small and conical as that of the short-faced tumbler; for one so broad and short as that of the barb; for one so long, straight, and narrow, with its enormous wattles, as that of the English carrier; for an expanded upraised tail like that of the fantail; or for an œsophagus like that of the pouter. I do not for a moment pretend that the domestic races differ from each other in their whole organisation as much as the more distinct natural genera. I refer only to external characters, on which, however, it must be confessed that most genera of birds have been founded. When, in a future chapter, we discuss the principle of selection as followed by man, we shall clearly see why the differences between the domestic races are almost always confined to external, or at least to externally visible, characters.

Owing to the amount and gradations of difference between the several breeds, I have found it indispensable in the following classification to rank them under Groups, Races, and Sub-races; to which varieties and sub-varieties, all strictly inheriting their proper characters, must often be added. Even with the individuals of the same sub-variety, when long kept by different fanciers, different strains can sometimes be recognised. There can be no doubt that, if well-characterized forms of the several Races had been found wild, all would have been ranked as distinct species, and several of them would certainly have been placed by ornithologists in distinct genera. A good classification of the various domestic breeds is extremely difficult, owing to the manner in which many of the forms graduate into each other; but it is curious how exactly the same difficulties are encountered, and the same rules have to be followed, as in the classification of any natural but difficult group of organic beings. An "artificial classification" might be followed which would present fewer difficulties than a "natural classification;" but then it would interrupt many plain affinities. Extreme forms can readily be defined; but intermediate and troublesome forms {134}often destroy our definitions. Forms which may be called "aberrant" must sometimes be included within groups to which they do not accurately belong. Characters of all kinds must be used; but as with birds in a state of nature, those afforded by the beak are the best and most readily appreciated. It is not possible to weigh the importance of all the characters which have to be used so as to make the groups and sub-groups of equal value. Lastly, a group may contain only one race, and another and less distinctly defined group may contain several races and sub-races, and in this case it is difficult, as in the classification of natural species, to avoid placing too high a value on characters which are common to a large number of forms.

In my measurements I have never trusted to the eye; and when speaking of a part being large or small, I always refer to the wild rock-pigeon (Columba livia) as the standard of comparison. The measurements are given in decimals of an inch.^[279]

I will now give a brief description of all the principal breeds. The following diagram may aid the reader in learning their names and seeing their affinities. The rock-pigeon, or Columba livia (including under this name two or three closely-allied sub-species or geographical races, hereafter to be described), may be confidently viewed, as we shall see in the next chapter, as the common parent-form. The names in italics on the right-hand side of the table show us the most distinct breeds, or those which have undergone the greatest amount of modification. The lengths of the dotted lines rudely represent the degree of distinctness of each breed from the parent-stock, and the names {135}placed under each other in the columns show the more or less closely connecting links. The distances of the dotted lines from each other approximately represent the amount of difference between the several breeds.

Fig. 17.—The Rock-pigeon, or Columba livia.^[280] The parent-form of all domesticated Pigeons.

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Group I.

This group includes a single race, that of the Pouters. If the most strongly marked sub-race be taken, namely, the Improved English Pouter, this is perhaps the most distinct of all domesticated pigeons.

Fig. 18.—English Pouter.

Race I.—Pouter Pigeons. (Kropf-tauben, German. Grosses-gorges, or boulans, French.)

Œsophagus of great size, barely separated from the crop, often inflated. Body and legs elongated. Beak of moderate dimensions. {138}

Sub-race I.—The improved English Pouter, when its crop is fully inflated, presents a truly astonishing appearance. The habit of slightly inflating the crop is common to all domestic pigeons, but is carried to an extreme in the Pouter. The crop does not differ, except in size, from that of other pigeons; but is less plainly separated by an oblique construction from the œsophagus. The diameter of the upper part of the œsophagus is immense, even close up to the head. The beak in one bird which I possessed was almost completely buried when the œsophagus was fully expanded. The males, especially when excited, pout more than the females, and they glory in exercising this power. If a bird will not, to use the technical expression, "play," the fancier, as I have witnessed, by taking the beak into his mouth, blows him up like a balloon; and the bird, then puffed up with wind and pride, struts about, retaining his magnificent size as long as he can. Pouters often take flight with their crops inflated; and after one of my birds had swallowed a good meal of peas and water, as he flew up in order to disgorge them and thus feed his nearly fledged young, I have heard the peas rattling in his inflated crop as if in a bladder. When flying, they often strike the backs of their wings together, and thus make a clapping noise.

Pouters stand remarkably upright, and their bodies are thin and elongated. In connexion with this form of body, the ribs are generally broader and the vertebræ more numerous than in other breeds. From their manner of standing their legs appear longer than they really are, though, in proportion with those of C. livia, the legs and feet are actually longer. The wings appear much elongated, but by measurement, in relation to the length of body, this is not the case. The beak likewise appears longer, but it is in fact a little shorter (about .03 of an inch), proportionally with the size of the body, and relatively to the beak of the rock-pigeon. The Pouter, though not bulky, is a large bird; I measured one which was 34½ inches from tip to tip of wing, and 19 inches from tip of beak to end of tail. In a wild rock-pigeon from the Shetland Islands the same measurements gave only 28¼ and 14¾. There are many sub-varieties of the Pouter of different colours, but these I pass over.

Sub-race II. Dutch Pouter.—This seems to be the parent-form of our improved English Pouters. I kept a pair, but I suspect that they were not pure birds. They are smaller than English pouters, and less well developed in all their characters. Neumeister^[281] says that the wings are crossed over the tail, and do not reach to its extremity.

Sub-race III. The Lille Pouter—I know this breed only from description.^[282] It approaches in general form the Dutch Pouter, but the inflated œsophagus assumes a spherical form, as if the pigeon had swallowed a large orange, which had stuck close under the beak. This inflated ball is represented as rising to a level with the crown of the head. The middle toe alone is feathered. A variety of this sub-race, called the claquant, is described by MM. Boitard and Corbié; it pouts but little, and is characterised {139}by the habit of violently hitting its wings together over its back,—a habit which the English Pouter has in a slight degree.

Sub-race IV. Common German Pouter.—I know this bird only from the figures and description given by the accurate Neumeister, one of the few writers on pigeons who, as I have found, may be always trusted. This sub-race seems considerably different. The upper part of the œsophagus is much less distended. The bird stands less upright. The feet are not feathered, and the legs and beak are shorter. In these respects there is an approach in form to the common rock-pigeon. The tail-feathers are very long, yet the tips of the closed wings extend beyond the end of the tail; and the length of the wings, from tip to tip, and of the body, is greater than in the English Pouter.

Group II.

This group includes three Races, namely, Carriers, Runts, and Barbs, which are manifestly allied to each other. Indeed, certain carriers and runts pass into each other by such insensible gradations that an arbitrary line has to be drawn between them. Carriers also graduate through foreign breeds into the rock-pigeon. Yet, if well-characterised Carriers and Barbs (see figs. 19 and 20) had existed as wild species, no ornithologist would have placed them in the same genus with each other or with the rock-pigeon. This group may, as a general rule, be recognised by the beak being long, with the skin over the nostrils swollen and often carunculated or wattled, and with that round the eyes bare and likewise carunculated. The mouth is very wide, and the feet are large. Nevertheless the Barb, which must be classed in this same group, has a very short beak, and some runts have very little bare skin round their eyes.

Race II.—Carriers. (Türkische Taube: Pigeons Turcs: Dragons.)

Beak elongated, narrow, pointed; eyes surrounded by much naked, generally carunculated skin; neck and body elongated.

Fig. 19.—English Carrier.

Sub-race I. The English Carrier.—This is a fine bird, of large size, close feathered, generally dark-coloured, with an elongated neck. The beak is attenuated and of wonderful length: in one specimen it was 1.4 inch in length from the feathered base to the tip; therefore nearly twice as long as that of the rock-pigeon, which measured only .77. Whenever I compare proportionally any part in the carrier and rock-pigeon, I take the length of the body from the base of the beak to the end of the tail as the standard of comparison; and according to this standard, the beak in one {140}Carrier was nearly half an inch longer than in the rock-pigeon. The upper mandible is often slightly arched. The tongue is very long. The development of the carunculated skin or wattle round the eyes, over the nostrils, and on the lower mandible, is prodigious. The eyelids, measured longitudinally, were in some specimens exactly twice as long as in the rock-pigeon. The external orifice or furrow of the nostrils was also twice as long. The open mouth in its widest part was in one case .75 of an inch in width, whereas in the rock-pigeon it is only about .4 of an inch. This great width of mouth is shown in the skeleton by the reflexed edges of the ramus of the lower jaw. The head is flat on the summit and narrow between the orbits. The feet are large and coarse; the length, as {141}measured from end of hind toe to end of middle toe (without the claws), was in two specimens 2.6 inches; and this, proportionally with the rock-pigeon, is an excess of nearly a quarter of an inch. One very fine Carrier measured 31½ inches from tip to tip of wing. Birds of this sub-race are too valuable to be flown as carriers.

Sub-race II. Dragons; Persian Carriers.—The English Dragon differs from the improved English Carrier in being smaller in all its dimensions, and in having less wattle round the eyes and over the nostrils, and none on the lower mandible. Sir W. Elliot sent me from Madras a Bagdad Carrier (sometimes called khandési), the name of which shows its Persian origin; it would be considered here a very poor Dragon; the body was of the size of the rock-pigeon, with the beak a little longer, namely, 1 inch from the tip to the feathered base. The skin round the eyes was only slightly wattled, whilst that over the nostrils was fairly wattled. The Hon. C. Murray, also, sent me two Carriers direct from Persia; these had nearly the same character as the Madras bird, being about as large as the rock-pigeon, but the beak in one specimen was as much as 1.15 in length; the skin over the nostrils was only moderately, and that round the eyes scarcely at all wattled.

Sub-race III. Bagadotten-Tauben of Neumeister (Pavdotten or Hocker-Tauben).—I owe to the kindness of Mr. Baily, jun., a dead specimen of this singular breed imported from Germany. It is certainly allied to the Runts; nevertheless, from its close affinity with Carriers, it will be convenient here to describe it. The beak is long, and is hooked or bowed downwards in a highly remarkable manner, as will be seen in the woodcut to be hereafter given when I treat of the skeleton. The eyes are surrounded by a wide space of bright red skin, which, as well as that over the nostrils, is moderately wattled. The breast-bone is remarkably protuberant, being abruptly bowed outwards. The feet and tarsi are of great length, larger than in first-rate English Carriers. The whole bird is of large size, but in proportion to the size of the body the feathers of the wing and tail are short; a wild rock-pigeon, of considerably less size, had tail-feathers 4.6 inches in length, whereas in the large Bagadotten these feathers were scarcely over 4.1 inches in length. Riedel^[283] remarks that it is a very silent bird.

Sub-race IV. Bussorah Carrier.—Two specimens were sent me by Sir W. Elliot from Madras, one in spirits and the other skinned. The name shows its Persian origin. It is much valued in India, and is considered as a distinct breed from the Bagdad Carrier, which forms my second sub-race. At first I suspected that these two sub-races might have been recently formed by crosses with other breeds, though the estimation in which they are held renders this improbable; but in a Persian treatise,^[284] believed to have been written about 100 years ago, the Bagdad and Bussorah breeds are described as distinct. The Bussorah Carrier is of about the same size with the wild rock-pigeon. The shape of the beak, with some little carunculated skin over the nostrils,—the much elongated eyelids,—the {142}broad mouth measured internally,—the narrow head,—the feet proportionally a little longer than in the rock-pigeon,—and the general appearance, all show that this bird is an undoubted Carrier; yet in one specimen the beak was of exactly the same length as in the rock-pigeon. In the other specimen the beak (as well as the opening of the nostrils) was only a very little longer, viz. by .08 of an inch. Although there was a considerable space of bare and slightly carunculated skin round the eyes, that over the nostrils was only in a slight degree rugose. Sir W. Elliot informs me that in the living bird the eye seems remarkably large and prominent, and the same fact is noticed in the Persian treatise; but the bony orbit is barely larger than that in the rock-pigeon.

Amongst the several breeds sent to me from Madras by Sir W. Elliot there is a pair of the Kala Par, black birds with the beak slightly elongated, with the skin over the nostrils rather full, and with a little naked skin round the eyes. This breed seems more closely allied to the Carrier than to any other breed, being nearly intermediate between the Bussorah Carrier and the rock-pigeon.

The names applied in different parts of Europe and in India to the several kinds of Carriers all point to Persia or the surrounding countries as the source of this Race. And it deserves especial notice that, even if we neglect the Kala Par as of doubtful origin, we get a series broken by very small steps, from the rock-pigeon, through the Bussorah, which sometimes has a beak not at all longer than that of the rock-pigeon and with the naked skin round the eyes and over the nostrils very slightly swollen and carunculated, through the Bagdad sub-race and Dragons, to our improved English Carriers, which present so marvellous a difference from the rock-pigeon or Columba livia.

Race III.—Runts. (Scanderoons: Die Florentiner-Taube and Hinkel-Taube of Neumeister: Pigeon Bagadais, Pigeon Romain.)

Beak long, massive; body of great size.

Inextricable confusion reigns in the classification, affinities, and naming of Runts. Several characters which are generally pretty constant in other pigeons, such as the length of the wings, tail, legs, and neck, and the amount of naked skin round the eyes, are excessively variable in Runts. When the naked skin over the nostrils and round the eyes is considerably developed and wattled, and when the size of body is not very great, Runts graduate in so insensible a manner into Carriers, that the distinction is quite arbitrary. This fact is likewise shown by the names given to them in different parts of Europe. Nevertheless, taking the most distinct forms, at least five sub-races (some of them including well-marked varieties) can be distinguished, which differ in such important points of structure, that they would be considered as good species in a state of nature.

Sub-race I. Scanderoon of English writers (Die Florentiner and Hinkel-Taube of Neumeister).—Birds of this sub-race, of which I kept one alive {143}and have since seen two others, differ from the Bagadotten of Neumeister only in not haying the beak nearly so much curved downwards, and in the naked skin round the eyes and over the nostrils being hardly at all wattled. Nevertheless I have felt myself compelled to place the Bagadotten in Race II., or that of the Carriers, and the present bird in Race III., or that of the Runts. The Scanderoon has a very short, narrow, and elevated tail; wings extremely short, so that the first primary feathers were not longer than those of a small tumbler pigeon! Neck long, much bowed; breast-bone prominent. Beak long, being 1.15 inch from tip to feathered base; vertically thick; slightly curved downwards. The skin over the nostrils swollen, not wattled; naked skin round the eyes, broad, slightly carunculated. Legs long; feet very large. Skin of neck bright red, often showing a naked medial line, with a naked red patch at the distant end of the radius of the wing. My bird, as measured from the base of the beak to the root of the tail, was fully 2 inches longer than the rock-pigeon; yet the tail itself was only 4 inches in length, whereas in the rock-pigeon, which is a much smaller bird, the tail is 4⅝ inches in length.

The Hinkel or Florentiner-Taube of Neumeister (Table XIII., fig. 1) agrees with the above description in all the specified characters (for the beak is not mentioned), except that Neumeister expressly says that the neck is short, whereas in my Scanderoon it was remarkably long and bowed; so that the Hinkel forms a well-marked variety.

Sub-race II. Pigeon Cygne and Pigeon Bagadais of Boitard and Corbié (Scanderoon of French writers).—I kept two of these birds alive, imported from France. They differed from the first sub-race or true Scanderoon in the much greater length of the wing and tail, in the beak not being so long, and in the skin about the head being more carunculated. The skin of the neck is red; but the naked patches on the wings are absent. One of my birds measured 38½ inches from tip to tip of wing. By taking the length of the body as the standard of comparison, the two wings were no loss than 5 inches longer than those of the rock-pigeon! The tail was 6¼ inches in length, and therefore 2¼ inches longer than that of the Scanderoon,—a bird of nearly the same size. The beak is longer, thicker, and broader than in the rock-pigeon, proportionally with the size of body. The eyelids, nostrils, and internal gape of mouth are all proportionally very large, as in Carriers. The foot, from the end of the middle to end of hind toe, was actually 2.85 inches in length, which is an excess of .32 of an inch over the foot of the rock-pigeon, relatively to the size of the two birds.

Sub-race III. Spanish and Roman Runts.—I am not sure that I am right in placing these Runts in a distinct sub-race; yet, if we take well-characterized birds, there can be no doubt of the propriety of the separation. They are heavy, massive birds, with shorter necks, legs, and beaks than in the foregoing races. The skin over the nostrils is swollen, but not carunculated; the naked skin round the eyes is not very wide, and only slightly carunculated; and I have seen a fine so-called Spanish Runt with hardly any naked skin round the eyes. Of the two varieties to be seen in England, one, which is the rarer, has very long wings and tail, {144}and agrees pretty closely with the last sub-race; the other, with shorter wings and tail, is apparently the Pigeon Romain ordinaire of Boitard and Corbié. These Runts are apt to tremble like Fantails. They are bad flyers. A few years ago Mr. Gulliver^[285] exhibited a Runt which weighed 1 lb. 14 oz.; and, as I am informed by Mr. Tegetmeier, two Runts from the south of France were lately exhibited at the Crystal Palace, each of which weighed 2 lbs. 2½ oz. A very fine rock-pigeon from the Shetland Islands weighed only 14½ oz.

Sub-race IV. Tronfo of Aldrovandi (Leghorn Runt?).—In Aldrovandi's work published in 1600 there is a coarse woodcut of a great Italian pigeon, with an elevated tail, short legs, massive body, and with the beak short and thick. I had imagined that this latter character, so abnormal in the group, was merely a false representation from bad drawing; but Moore, in his work published in 1735, says that he possessed a Leghorn Runt of which "the beak was very short for so large a bird." In other respects Moore's bird resembled the first sub-race or Scanderoon, for it had a long bowed neck, long legs, short beak, and elevated tail, and not much wattle about the head. So that Aldrovandi's and Moore's birds must have formed distinct varieties, both of which seem to be now extinct in Europe. Sir W. Elliot, however, informs me that he has seen in Madras a short-beaked Runt imported from Cairo.

Sub-race V. Murassa (adorned Pigeon) of Madras.—Skins of these handsome chequered birds were sent me from Madras by Sir W. Elliot. They are rather larger than the largest rock-pigeon, with longer and more massive beaks. The skin over the nostrils is rather full and very slightly carunculated, and they have some naked skin round the eyes: feet large. This breed is intermediate between the rock-pigeon and a very poor variety of Runt or Carrier.

From these several descriptions we see that with Runts, as with Carriers, we have a fine gradation from the rock-pigeon (with the Tronfo diverging as a distinct branch) to our largest and most massive Runts. But the chain of affinities, and many points of resemblance, between Runts and Carriers, make me believe that these two races have not descended by independent lines from the rock-pigeon, but from some common parent, as represented in the Table, which had already acquired a moderately long beak, with slightly swollen skin over the nostrils, and with some slightly carunculated naked skin round the eyes.

Race IV.—Barbs. (Indische-Taube: Pigeons Polonais.)

Beak short, broad, deep; naked skin round the eyes, broad and carunculated; skin over nostrils slightly swollen.

Fig. 20.—English Barb.

Misled by the extraordinary shortness and form of the beak, I did not at first perceive the near affinity of this Race to that of Carriers until the fact was pointed out to me by Mr. Brent. Subsequently, after examining {145}the Bussorah Carrier, I saw that no very great amount of modification would be requisite to convert it into a Barb. This view of the affinity of Barbs to Carriers is supported by the analogical difference between the short and long-beaked Runts; and still more strongly by the fact, that young Barbs and Dragons, within 24 hours after being hatched, resemble each other much more closely than do young pigeons of other and equally distinct breeds. At this early age, the length of beak, the swollen skin over the rather open nostrils, the gape of the mouth, and the size of the feet, are the same in both; although these parts afterwards become widely different. We thus see that embryology (as the comparison of very young animals {146}may perhaps be called) comes into play in the classification of domestic varieties, as with species in a state of nature.

Fanciers, with some truth, compare the head and beak of the Barb to that of a bullfinch. The Barb, if found in a state of nature, would certainly have been placed in a new genus formed for its reception. The body is a little larger than that of the rock-pigeon, but the beak is more than .2 of an inch shorter; although shorter, it is both vertically and horizontally thicker. From the outward flexure of the rami of the lower jaw, the mouth internally is very broad, in the proportion of .6 to .4 to that of the rock-pigeon. The whole head is broad. The skin over the nostrils is swollen, but not carunculated, except slightly in first-rate birds when old; whilst the naked skin round the eye is broad and much carunculated. It is sometimes so much developed, that a bird belonging to Mr. Harrison Weir could hardly see to pick up food from the ground. The eyelids in one specimen were nearly twice as long as those of the rock-pigeon. The feet are coarse and strong, but proportionally rather shorter than in the rock-pigeon. The plumage is generally dark and uniform. Barbs, in short, may be called short-beaked Carriers, bearing the same relation to Carriers that the Tronfo of Aldrovandi does to the common Runt.

Group III.

This group is artificial, and includes a heterogeneous collection of distinct forms. It may be defined by the beak, in well-characterised specimens of the several races, being shorter than in the rock-pigeon, and by the skin round the eyes not being much developed.

Race V.—Fantails.

Sub-race I. European Fantails (Pfauen-Taube; Trembleurs). Tail expanded, directed upwards, formed of many feathers; oil-gland aborted; body and beak rather short.

Fig. 21.—English Fantail.

The normal number of tail-feathers in the genus Columba is 12; but Fantails have from only 12 (as has been asserted) up to, according to MM. Boitard and Corbié, 42. I have counted in one of my own birds 33, and at Calcutta Mr. Blyth^[286] has counted in an imperfect tail 34 feathers. In Madras, as I am informed by Sir W. Elliot, 32 is the standard number; but in England number is much less valued than the position and expansion of the tail. The feathers are arranged in an irregular double row; their permanent expansion, like a fan, and their upward direction, are more remarkable characters than their increased number. The tail is capable of the same movements as in other pigeons, and can be depressed so as to sweep the ground. It arises from a more expanded basis than in {147}other pigeons; and in three skeletons there were one or two extra coccygeal vertebræ. I have examined many specimens of various colours from different countries, and there was no trace of the oil-gland; this is a curious case of abortion.^[287] The neck is thin and bowed {148}backwards. The breast is broad and protuberant. The feet are small. The carriage of the bird is very different from that of other pigeons; in good birds the head touches the tail-feathers, which consequently often become crumpled. They habitually tremble much; and their necks have an extraordinary, apparently convulsive, backward and forward movement. Good birds walk in a singular manner, as if their small feet were stiff. Owing to their large tails, they fly badly on a windy day. The dark-coloured varieties are generally larger than white Fantails.

Although between the best and common Fantails, now existing in England, there is a vast difference in the position and size of the tail, in the carriage of the head and neck, in the convulsive movements of the neck, in the manner of walking, and in the breadth of the breast, the differences so graduate away, that it is impossible to make more than one sub-race. Moore, however, an excellent old authority,^[288] says, that in 1735 there were two sorts of broad-tailed shakers (i.e. fantails), "one having a neck much longer and more slender than the other;" and I am informed by Mr. B. P. Brent that there is an existing German Fantail with a thicker and shorter beak.

Sub-race II. Java Fantail.—Mr. Swinhoe sent me from Amoy, in China, the skin of a Fantail belonging to a breed known to have been imported from Java. It was coloured in a peculiar manner, unlike any European Fantail, and, for a Fantail, had a remarkably short beak. Although a good bird of the kind, it had only 14 tail-feathers; but Mr. Swinhoe has counted in other birds of this breed from 18 to 24 tail-feathers. From a rough sketch sent to me, it is evident that the tail is not so much expanded or so much upraised as in even second-rate European Fantails. The bird shakes its neck like our Fantails. It had a well-developed oil-gland. Fantails were known in India, as we shall hereafter see, before the year 1600; and we may suspect that in the Java Fantail we see the breed in its earlier and less improved condition.

Race VI.—Turbit and Owl. (Möven-Taube: Pigeons à cravate.)

Feathers divergent along the front of the neck and breast; beak very short, vertically rather thick; œsophagus somewhat enlarged.

Fig. 22.—African Owl.

Turbits and Owls differ from each other slightly in the shape of the head, in the former having a crest, and in the curvature of the beak, but they may be here conveniently grouped together. These pretty birds, some of which are very small, can be recognised at once by the feathers irregularly diverging, like a frill, along the front of the neck, in the same manner, but in a less degree, as along the back of the neck in the Jacobin. This bird has the remarkable habit of continually, and momentarily inflating the upper part of the œsophagus, which causes a movement in the frill. {149}When the œsophagus of a dead bird was inflated, it was seen to be larger than in other breeds, and not so distinctly separated from the crop. The Pouter inflates both its true crop and œsophagus; the Turbit inflates in a much less degree the œsophagus alone. The beak of the Turbit is very short, being .28 of an inch shorter than that of the rock-pigeon, proportionally with the size of their bodies; and in some owls brought by Mr. E. Vernon Harcourt from Tunis, it was even shorter. The beak is vertically thicker, and perhaps a little broader, in proportion to that of the rock-pigeon.

{150}

Race VII.—Tumblers. (Tümmler, or Burzel-Tauben: Culbutants.)

During flight, tumble backwards; body generally small; beak generally short, sometimes excessively short and conical.

This Race may be divided into four sub-races, namely, Persian, Lotan, Common, and Short-faced Tumblers. These sub-races include many varieties which breed true. I have examined eight skeletons of various kinds of Tumblers: excepting in one imperfect and doubtful specimen, the ribs are only seven in number, whereas the rock-pigeon has eight ribs.

Sub-race I. Persian Tumblers.—I have received a pair direct from Persia, from the Hon. C. Murray. They were rather smaller birds than the wild rock-pigeon, being about the size of the common dovecot-pigeon, white and mottled, slightly feathered on the feet, with the beak just perceptibly shorter than in the rock-pigeon. H.M. Consul, Mr. Keith Abbott, informs me that the difference in the length of beak is so slight, that only practised Persian fanciers can distinguish these Tumblers from the common pigeon of the country. He informs me that they fly in flocks high up in the air and tumble well. Some of them occasionally appear to become giddy and tumble to the ground, in which respect they resemble some of our Tumblers.

Sub-race II. Lotan, or Lowtun: Indian Ground Tumblers.—These birds present one of the most remarkable inherited habits or instincts which have ever been recorded. The specimens sent to me from Madras by Sir W. Elliot are white, slightly feathered on the feet, with the feathers on the head reversed; and they are rather smaller than the rock or dovecot pigeon. The beak is proportionally only slightly shorter and rather thinner than in the rock-pigeon. These birds when gently shaken and placed on the ground immediately begin tumbling head over heels, and they continue thus to tumble until taken up and soothed,—the ceremony being generally to blow in their faces, as in recovering a person from a state of hypnotism or mesmerism. It is asserted that they will continue to roll over till they die, if not taken up. There is abundant evidence with respect to these remarkable peculiarities; but what makes the case the more worthy of attention is, that the habit has been strictly inherited since before the year 1600, for the breed is distinctly described in the 'Ayeen Akbery.'^[289] Mr. Evans kept a pair in London, imported by Captain Vigne; and he assures me that he has seen them tumble in the air, as well as in the manner above described on the ground. Sir W. Elliot, however, writes to me from Madras, that he is informed that they tumble exclusively on the ground, or at a very small height above it. He also {151}mentions another sub-variety, called the Kalmi Lotan, which begins to roll over if only touched on the neck with a rod or wand.

Sub-race III. Common English Tumblers.—These birds have exactly the same habits as the Persian Tumbler, but tumble better. The English bird is rather smaller than the Persian, and the beak is plainly shorter. Compared with the rock-pigeon, and proportionally with the size of body, the beak is from .16 to nearly .2 of an inch shorter, but it is not thinner. There are several varieties of the common Tumbler, namely, Baldheads, Beards, and Dutch Rollers. I have kept the latter alive; they have differently shaped heads, longer necks, and are feather-footed. They tumble to an extraordinary degree; as Mr. Brent remarks,^[290] "Every few seconds over they go; one, two, or three summersaults at a time. Here and there a bird gives a very quick and rapid spin, revolving like a wheel, though they sometimes lose their balance, and make a rather ungraceful fall, in which they occasionally hurt themselves by striking some object." From Madras I have received several specimens of the common Tumbler of India, differing slightly from each other in the length of their beaks. Mr. Brent sent me a dead specimen of a "House-tumbler,"^[291] which is a Scotch variety, not differing in general appearance and form of beak from the common Tumbler. Mr. Brent states that these birds generally begin to tumble "almost as soon as they can well fly; at three months old they tumble well, but still fly strong; at five or six months they tumble excessively; and in the second year they mostly give up flying, on account of their tumbling so much and so close to the ground. Some fly round with the flock, throwing a clean summersault every few yards, till they are obliged to settle from giddiness and exhaustion. These are called Air Tumblers, and they commonly throw from twenty to thirty summersaults in a minute, each clear and clean. I have one red cock that I have on two or three occasions timed by my watch, and counted forty summersaults in the minute. Others tumble differently. At first they throw a single summersault, then it is double, till it becomes a continuous roll, which puts an end to flying, for if they fly a few yards over they go, and roll till they reach the ground. Thus I had one kill herself, and another broke his leg. Many of them turn over only a few inches from the ground, and will tumble two or three times in flying across their loft. These are called House-tumblers, from tumbling in the house. The act of tumbling seems to be one over which they have no control, an involuntary movement which they seem to try to prevent. I have seen a bird sometimes in his struggles fly a yard or two straight upwards, the impulse forcing him backwards while he struggles to go forwards. If suddenly startled, or in a strange place, they seem less able to fly than if quiet in their accustomed loft." These House-tumblers differ from the Lotan or Ground {152}Tumbler of India, in not requiring to be shaken in order to begin tumbling. The breed has probably been formed merely by selecting the best common Tumblers, though it is possible that they may have been crossed at some former period with Lotans.

Fig. 23.—Short-faced English Tumbler.

Sub-race IV. Short-faced Tumblers.—These are marvellous birds, and are the glory and pride of many fanciers. In their extremely short, sharp, and conical beaks, with the skin over the nostrils but little developed, they almost depart from the type of the Columbidæ. Their heads are nearly globular {153}and upright in front, so that some fanciers say^[292] "the head should resemble a cherry with a barley-corn stuck in it." These are the smallest kind of pigeons. Mr. Esquilant possessed a blue Baldhead, two years old, which when alive weighed, before feeding-time, only 6 oz. 5 drs.; two others, each weighed 7 oz. We have seen that a wild rock-pigeon weighed 14 oz. 2 drs., and a Runt 34 oz. 4 drs. Short-faced Tumblers have a remarkably erect carriage, with prominent breasts, drooping wings, and very small feet. The length of the beak from the tip to the feathered base was in one good bird only .4 of an inch; in a wild rock-pigeon it was exactly double this length. As these Tumblers have shorter bodies than the wild rock-pigeon, they ought of course to have shorter beaks; but proportionally with the size of body, the beak is .28 of an inch too short. So, again, the feet of this bird were actually .45 shorter, and proportionally .21 of an inch shorter, than the feet of the rock-pigeon. The middle toe has only twelve or thirteen, instead of fourteen or fifteen scutellæ. The primary wing-feathers are not rarely only nine instead of ten in number. The improved short-faced Tumblers have almost lost the power of tumbling; but there are several authentic accounts of their occasionally tumbling. There are several sub-varieties, such as Baldheads, Beards, Mottles, and Almonds; the latter are remarkable from not acquiring their perfectly-coloured plumage until they have moulted three or four times. There is good reason to believe that most of these sub-varieties, some of which breed truly, have arisen since the publication of Moore's treatise in 1735.^[293]

Finally, in regard to the whole group of Tumblers, it is impossible to conceive a more perfect gradation than I have now lying before me, from the rock-pigeon, through Persian, Lotan, and Common Tumblers, up to the marvellous short-faced birds; which latter, no ornithologist, judging from mere external structure, would place in the same genus with the rock-pigeon. The differences between the successive steps in this series are not greater than those which may be observed between common dovecot-pigeons (C. livia) brought from different countries.

Race VIII—Indian Frill-back.

Beak very short; feathers reversed.

A specimen of this bird, in spirits, was sent to me from Madras by Sir W. Elliot. It is wholly different from the Frill-back often exhibited in England. It is a smallish bird, about the size of the common Tumbler, but has a beak in all its proportions like our short-faced Tumblers. The beak, measured from the tip to the feathered base, was only .46 of an inch in length. The feathers over the whole body are reversed or curl backwards. Had this bird occurred in Europe, I should have thought it only a monstrous variety of our improved Tumbler; but as short-faced Tumblers are not known in India, I think it must rank as a distinct breed. Probably {154}this is the breed seen by Hasselquist in 1757 at Cairo, and said to have been imported from India.

Race IX.—Jacobin. (Zopf or Perücken-Taube: Nonnains.)

Feathers of the neck forming a hood; wings and tail long; beak moderately short.

This pigeon can at once be recognised by its hood, almost enclosing the head and meeting in front of the neck. The hood seems to be merely an exaggeration of the crest of reversed feathers on the back of the head, which is common to many sub-varieties, and which in the Latz-taube^[294] is in a nearly intermediate state between a hood and a crest. The feathers of the hood are elongated. Both the wings and tail are likewise much elongated; thus the folded wing of the Jacobin, though a somewhat smaller bird, is fully 1¼ inch longer than in the rock-pigeon. Taking the length of the body without the tail as the standard of comparison, the folded wing, proportionally with the wings of the rock-pigeon, is 2¼ inches too long, and the two wings, from tip to tip, 5¼ inches too long. In disposition this bird is singularly quiet, seldom flying or moving about, as Bechstein and Riedel have likewise remarked in Germany.^[295] The latter author also notices the length of the wings and tail. The beak is nearly .2 of an inch shorter in proportion to the size of the body than in the rock-pigeon; but the internal gape of the mouth is considerably wider.

Group IV.

The birds of this group may be characterised by their resemblance in all important points of structure, especially in the beak, to the rock-pigeon. The Trumpeter forms the only well-marked race. Of the numerous other sub-races and varieties I shall specify only a few of the most distinct, which I have myself seen and kept alive.

Race X.—Trumpeter. (Trommel-Taube; Pigeon tambour; glougou.)

A tuft of feathers at the base of the beak curling forward; feet much feathered; voice very peculiar; size exceeding that of the rock-pigeon.

This is a well-marked breed, with a peculiar voice, wholly unlike that of any other pigeon. The coo is rapidly repeated, and is continued for {155}several minutes; hence their name of Trumpeters. They are also characterised by a tuft of elongated feathers, which curls forward over the base of the beak, and which is possessed by no other breed. Their feet are so heavily feathered, that they almost appear like little wings. They are larger birds than the rock-pigeon, but their beak is of very nearly the same proportional size. Their feet are rather small. This breed was perfectly characterised in Moore's time, in 1735. Mr. Brent says that two varieties exist, which differ in size.

Race XI.—Scarcely differing in structure from the wild Columba livia.

Sub-race 1. Laughers. Size less than the Rock-pigeon; voice very peculiar.—As this bird agrees in nearly all its proportions with the rock-pigeon, though of smaller size, I should not have thought it worthy of mention, had it not been for its peculiar voice—a character supposed seldom to vary with birds. Although the voice of the Laugher is very different from that of the Trumpeter, yet one of my Trumpeters used to utter a single note like that of the Laugher. I have kept two varieties of Laughers, which differed only in one variety being turn-crowned; the smooth-headed kind, for which I am indebted to the kindness of Mr. Brent, besides its peculiar note, used to coo in a singular and pleasing manner, which, independently, struck both Mr. Brent and myself as resembling that of the turtle-dove. Both varieties come from Arabia. This breed was known by Moore in 1735. A pigeon which seems to say Yak-roo is mentioned in 1600 in the 'Ayeen Akbery,' and is probably the same breed. Sir W. Elliot has also sent me from Madras a pigeon called Yahui, said to have come from Mecca, which does not differ in appearance from the Laugher; it has "a deep melancholy voice, like Yahu, often repeated." Yahu, yahu, means Oh God, Oh God; and Sayzid Mohammed Musari, in the treatise written about 100 years ago, says that these birds "are not flown, because they repeat the name of the Most High God." Mr. Keith Abbott, however, informs me that the common pigeon is called Yahoo in Persia.

Sub-race II. Common Frill-back (Die Strupp-Taube). Beak rather longer than in the Rock-pigeon; feathers reversed.—This is a considerably larger bird than the rock-pigeons and with the beak, proportionally with the size of body, a little (viz. by .04 of an inch) longer. The feathers, especially on the wing-coverts, have their points curled upwards or backwards.

Sub-race III. Nuns (Pigeons-coquilles).—These elegant birds are smaller than the rock-pigeon. The beak is actually .17, and proportionally with the size of the body .1 of an inch shorter than in the rock-pigeons, although of the same thickness. In young birds the scutellæ on the tarsi and toes are generally of a leaden-black colour; and this is a remarkable character (though observed in a lesser degree in some other breeds), as the colour of the legs in the adult state is subject to very little variation in any breed. I have on two or three occasions counted thirteen or fourteen feathers in the tail; this likewise occurs in the barely distinct breed called Helmets. {156}Nuns are symmetrically coloured, with the head, primary wing-feathers, tail, and tail-coverts of the same colour, namely, black or red, and with the rest of the body white. This breed has retained the same character since Aldrovandi wrote in 1600. I have received from Madras almost similarly coloured birds.

Sub-race IV. Spots (Die Blass-Taube: Pigeons heurtés).—These birds are a very little larger than the rock-pigeon, with the beak a trace smaller in all its dimensions, and with the feet decidedly smaller. They are symmetrically coloured, with a spot on the forehead, with the tail and tail-coverts of the same colour, the rest of the body being white. This breed existed in 1676;^[296] and in 1735 Moore remarks that they breed truly, as is the case at the present day.

Sub-race V. Swallows.—These birds, as measured from tip to tip of wing, or from the end of the beak to the end of the tail, exceed in size the rock-pigeon; but their bodies are much less bulky; their feet and legs are likewise smaller. The beak is of about the same length, but rather slighter. Altogether their general appearance is considerably different from that of the rock-pigeon. Their heads and wings are of the same colour, the rest of the body being white. Their flight is said to be peculiar. This seems to be a modern breed, which, however, originated before the year 1795 in Germany, for it is described by Bechstein.

---

Besides the several breeds now described, three or four other very distinct kinds existed lately, or perhaps still exist, in Germany and France. Firstly, the Karmeliten, or Carme Pigeon, which I have not seen; it is described as of small size, with very short legs, and with an extremely short beak. Secondly, the Finnikin, which is now extinct in England. It had, according to Moore's^[297] treatise, published in 1735, a tuft of feathers on the hinder part of the head, which ran down its back not unlike a horse's mane. "When it is salacious it rises over the hen and turns round three or four times, flapping its wings, then reverses and turns as many times the other way." The Turner, on the other hand, when it "plays to the female, turns only one way." Whether these extraordinary statements may be trusted I know not; but the inheritance of any habit may be believed, after what we have seen with respect to the Ground-tumbler of India. MM. Boitard and Corbié describe a pigeon^[298] which has the singular habit of sailing for a considerable time through the air, without flapping its wings, like a bird of prey. The confusion is inextricable, from the time of Aldrovandi in 1600 to the present day, in the accounts published of the Draijers, Smiters, Finnikins, Turners, Claquers, &c., which are all remarkable from their manner of flight. Mr. Brent informs me that he has seen one of these breeds in Germany with its wing-feathers injured from having been so often struck together; but he did not see it flying. An old stuffed specimen of a Finnikin in the British Museum presents no well-marked character. Thirdly, a singular pigeon {157}with a forked tail is mentioned in some treatises; and as Bechstein^[299] briefly describes and figures this bird, with a tail "having completely the structure of that of the house-swallow," it must once have, existed, for Bechstein was far too good a naturalist to have confounded any distinct species with the domestic pigeon. Lastly, an extraordinary pigeon imported from Belgium has lately been exhibited at the Philoperisteron Society in London,^[300] which "conjoins the colour of an archangel with the head of an owl or barb, its most striking peculiarity being the extraordinary length of the tail and wing-feathers, the latter crossing beyond the tail, and giving to the bird the appearance of a gigantic swift (Cypselus), or long-winged hawk." Mr. Tegetmeier informs me that this bird weighed only 10 ounces, but in length was 15½ inches from tip of beak to end of tail, and 32½ inches from tip to tip of wing; now the wild rock-pigeon weighs 14½ ounces, and measures from tip of beak to end of tail 15 inches, and from tip to tip of wing only 26¾ inches.

I have now described all the domestic pigeons known to me, and have added a few others on reliable authority. I have classed them under four Groups, in order to mark their affinities and degrees of difference; but the third group is artificial. The kinds examined by me form eleven races, which include several sub-races; and even these latter present differences that would certainly have been thought of specific value if observed in a state of nature. The sub-races likewise include many strictly inherited varieties; so that altogether there must exist, as previously stated, above 150 kinds which can be distinguished, though generally by characters of extremely slight importance. Many of the genera of the Columbidæ, which are admitted by ornithologists, do not differ in any great degree from each other; taking this into consideration, there can be no doubt that several of the most strongly characterised domestic forms, if found wild, would have been placed in at least five new genera. Thus, a new genus would have been formed for the reception of the improved English Pouter: a second genus for Carriers and Runts; and this would have been a wide or comprehensive genus, for it would have admitted common Spanish Runts without any wattle, short-beaked Runts like the Tronfo, and the improved English Carrier: a third genus would have been termed for the Barb: a fourth for the Fantail: and lastly, a fifth for the short-beaked, not-wattled pigeons, such as Turbits {158}and short-faced Tumblers. The remaining domestic forms might have been included in the same genus with the wild rock-pigeon.

Individual Variability; Variations of a remarkable nature.

The differences which we have as yet considered are characteristic of distinct breeds; but there are other differences, either confined to individual birds, or often observed in certain breeds but not characteristic of them. These individual differences are of importance, as they might in most cases be secured and accumulated by man's power of selection; and thus an existing breed might be greatly modified or a new one formed. Fanciers notice and select only those slight differences which are externally visible; but the whole organisation is so tied together by correlation of growth, that a change in one part is frequently accompanied by other changes. For our purpose, modifications of all kinds are equally important, and, if affecting a part which does not commonly vary, are of more importance than a modification in some conspicuous part. At the present day any visible deviation of character in a well-established breed is rejected as a blemish; but it by no means follows that at an early period, before well-marked breeds had been formed, such deviations would have been rejected; on the contrary, they would have been eagerly preserved as presenting a novelty, and would then have been slowly augmented, as we shall hereafter more clearly see, by the process of unconscious selection.

I have made numerous measurements of the various parts of the body in the several breeds, and have hardly ever found them quite the same in birds of the same breed,—the differences being greater than we commonly meet with in wild species. To begin with the primary feathers of the wing and tail; but I may first mention, as some readers may not be aware of the fact, that the number of the primary wing and tail feathers in wild birds is generally constant, and characterises, not only whole genera, but even whole families. When the tail-feathers are unusually numerous, as for instance in the swan, they are apt to be variable in number; but this does not apply to the several species and genera of the Columbidæ, which never (as far as I can hear) have less than twelve or more than sixteen tail-feathers; and these numbers characterise, with rare exception, whole sub-families.^[301] The wild rock-pigeon has twelve tail-feathers. With {159}Fantails, as we have seen, the number varies from fourteen to forty-two. In two young birds in the same nest I counted twenty-two and twenty-seven feathers. Pouters are very liable to have additional tail-feathers, and I have seen on several occasions fourteen or fifteen in my own birds, Mr. Bult had a specimen, examined by Mr. Yarrell, with seventeen tail-feathers. I had a Nun with thirteen, and another with fourteen tail-feathers; and in a Helmet, a breed barely distinguishable from the Nun, I have counted fifteen, and have heard of other such instances. On the other hand, Mr. Brent possessed a Dragon, which during its whole life never had more than ten tail-feathers; and one of my Dragons, descended from Mr. Brent's, had only eleven. I have seen a Baldhead-Tumbler with only ten; and Mr. Brent had an Air-Tumbler with the same number, but another with fourteen tail-feathers. Two of these latter Tumblers, bred by Mr. Brent, were remarkable,—one from having the two central tail-feathers a little divergent, and the other from having the two outer feathers longer by three-eighths of an inch than the others; so that in both cases the tail exhibited a tendency, but in different ways, to become forked. And this shows us how a swallow-tailed breed, like that described by Bechstein, might have been formed by careful selection.

With respect to the primary wing-feathers, the number in the Columbidæ, as far as I can find out, is always nine or ten. In the rock-pigeon it is ten; but I have seen no less than eight short-faced Tumblers with only nine primaries, and the occurrence of this number has been noticed by fanciers, owing to ten flight-feathers of a white colour being one of the points in Short-faced Baldhead-Tumblers. Mr. Brent, however, had an Air-Tumbler (not short-faced) which had in both wings eleven primaries. Mr. Corker, the eminent breeder of prize Carriers, assures me that some of his birds had eleven primaries in both wings. I have seen eleven in one wing in two Pouters. I have been assured by three fanciers that they have seen twelve in Scanderoons; but as Neumeister asserts that in the allied Florence Runt the middle flight-feather is often double, the number twelve may have been caused by two of the ten primaries having each two shafts to a single feather. The secondary wing-feathers are difficult to count, but the number seems to vary from twelve to fifteen. The length of the wing and tail relatively to the body, and of the wings to the tail, certainly varies; I have especially noticed this in Jacobins. In Mr. Bult's magnificent collection of Pouters, the wings and tail varied greatly in length; and were sometimes so much elongated that the birds could hardly play upright. In the relative length of the few first primaries I have observed only a slight degree of variability. Mr. Brent informs me that he has observed the shape of the first feather to vary very slightly. But the variation in these latter points is extremely slight compared with what may often be observed in the natural species of the Columbidæ.

In the beak I have observed very considerable differences in birds of the {160}same breed, as in carefully bred Jacobins and Trumpeters. In Carriers there is often a conspicuous difference in the degree of attenuation and curvature of the beak. So it is indeed in many breeds: thus I had two strains of black Barbs, which evidently differed in the curvature of the upper mandible. In width of mouth I have found a great difference in two Swallows. In Fantails of first-rate merit I have seen some birds with much longer and thinner necks than in others. Other analogous facts could be given. We have seen that the oil-gland is aborted in all Fantails (with the exception of the sub-race from Java), and, I may add, so hereditary is this tendency to abortion, that some, although not all, of the mongrels from the Fantail and Pouter had no oil-gland; in one Swallow out of many which I have examined, and in two Nuns, there was no oil-gland.

The number of the scutellæ on the toes often varies in the same breed, and sometimes even differs on the two feet of the same individual; the Shetland rock-pigeon has fifteen on the middle, and six on the hinder toe; whereas I have seen a Runt with sixteen on the middle and eight on the hind toe; and a short-faced Tumbler with only twelve and five on these same toes. The rock-pigeon has no sensible amount of skin between its toes; but I possessed a Spot and a Nun with the skin extending for a space of a quarter of an inch from the fork, between the two inner toes. On the other hand, as will hereafter be more fully shown, pigeons with feathered feet very generally have the bases of their outer toes connected by skin. I had a red Tumbler, which had a coo unlike that of its fellows, approaching in tone to that of the Laugher: this bird had the habit, to a degree which I never saw equalled in any other pigeon, of often walking with its wings raised and arched in an elegant manner. I need say nothing on the great variability, in almost every breed, in size of body, in colour, in the feathering of the feet, and in the feathers on the back of the head being reversed. But I may mention a remarkable Tumbler^[302] exhibited at the Crystal Palace, which had an irregular crest of feathers on its head, somewhat like the tuft on the head of the Polish fowl. Mr. Bult reared by accident a hen Jacobin with the feathers on the thigh so long as to reach the ground, and a cock having, but in a lesser degree, the same peculiarity: from these two birds he bred others similarly characterised, which were exhibited at the Philoperisteron Club. I bred a mongrel pigeon which had fibrous feathers, and the wing and tail-feathers so short and imperfect that the bird could not fly even a foot in height.

There are many singular and inherited peculiarities in the plumage of pigeons: thus Almond-Tumblers do not acquire their perfect mottled feathers until they have moulted three or four times: the Kite-Tumbler is at first brindled black and red with a barred appearance, but when "it throws its nest feathers it becomes almost black, generally with a bluish tail, and a reddish colour on the inner webs of the primary wing feathers."^[303] {161}Neumeister describes a breed of a black colour with white bars on the wing and a white crescent-shaped mark on the breast; these marks are generally rusty-red before the first moult, but after the third or fourth moult they undergo a change; the wing-feathers and the crown of the head likewise then become white or grey.^[304]

It is an important fact, and I believe there is hardly an exception to the rule, that the especial characters for which each breed is valued are eminently variable: thus, in the Fantail, the number and direction of the tail-feathers, the carriage of the body, and the degree of trembling are all highly variable points; in Pouters, the degree to which they pout, and the shape of their inflated crops; in the Carrier, the length, narrowness, and curvature of the beak, and the amount of wattle; in Short-faced Tumblers, the shortness of the beak, the prominence of the forehead, and general carriage,^[305] and in the Almond Tumbler the colour of the plumage; in common Tumblers, the manner of tumbling; in the Barb, the breadth and shortness of the beak and the amount of eye-wattle; in Runts, the size of body; in Turbits, the frill; and lastly in Trumpeters, the cooing, as well as the size of the tuft of feathers over the nostrils. These, which are the distinctive and selected characters of the several breeds, are all eminently variable.

There is another interesting fact with respect to the character of the different breeds, namely, that they are often most strongly displayed in the male bird. In Carriers, when the males and females are exhibited in separate pens, the wattle is plainly seen to be much more developed in the males, though I have seen a hen Carrier belonging to Mr. Haynes heavily wattled. Mr. Tegetmeier informs me that, in twenty Barbs in Mr. P. H. Jones's possession, the males had generally the largest eye-wattles; Mr. Esquilant also believes in this rule, but Mr. H. Weir, a first-rate judge, entertains some doubt on the subject. Hale Pouters distend their crops to a much greater size than do the females; I have, however, seen a hen in the possession of Mr. Evans which pouted excellently; but this is an unusual circumstance. Mr. Harrison Weir, a successful breeder of prize {162}Fantails, informs me that his cock birds often have a greater number of tail-feathers than the hens. Mr. Eaton asserts^[306] that, if a cock and hen Tumbler were of equal merit, the hen would be worth double the money; and as pigeons always pair, so that an equal number of both sexes is necessary for reproduction, this seems to show that high merit is rarer in the female than in the male. In the development of the frill in Turbits, of the hood in Jacobins, of the tuft in Trumpeters, of tumbling in Tumblers, there is no difference between the males and females. I may here add a rather different case, namely, the existence in France^[307] of a wine-coloured variety of the Pouter, in which the male is generally chequered with black, whilst the female is never so chequered. Dr. Chapuis also remarks^[308] that in certain light-coloured pigeons the males have their feathers striated with black, and these striæ increase in size at each moult, so that the male ultimately becomes spotted with black. With Carriers, the wattle, both on the beak and round the eyes, and with Barbs that round the eyes, goes on increasing with age. This augmentation of character with advancing age, and more especially the difference between the males and females in the above-mentioned several respects, are highly remarkable facts, for there is no sensible difference at any age between the two sexes in the aboriginal rock-pigeon; and rarely any such difference throughout the whole family of the Columbidæ.^[309]

Fig. 24.—Skulls of Pigeons, viewed laterally, of natural size. A. Wild Rock-pigeon, Columba livia. B. Short-faced Tumbler. C. English Carrier. D. Bagadotten Carrier.

Osteological Characters.

In the skeletons of the various breeds there is much variability; and though certain differences occur frequently, and others rarely, in certain breeds, yet none can be said to be absolutely characteristic of any breed. Considering that strongly-marked domestic races have been formed chiefly by man's power {163}of selection, we ought not to expect to find great and constant differences in the skeleton; for fanciers can neither see, nor do they care for, modifications of structure in the internal framework. Nor ought we to expect changes in the skeletons from hanged habits of life; as every facility is given to the most distinct breeds to follow the same habits, and the much modified races are never allowed to wander abroad and procure their own food in various ways. Moreover, I find, on comparing the skeletons of Columba livia, œnas, palumbus, and turtur, which are ranked by all systematists in two or three distinct though allied genera, that the differences are extremely slight, certainly less than between the skeletons of some of the most distinct domestic breeds. How far the skeleton of the wild rock-pigeon is constant I have no means of judging, as I have examined only two.

Skull.—The individual bones, especially those at the base, do not differ in shape. But the whole skull, in its proportions, outline, and relative direction of the bones, differs greatly in some of the breeds, as may be seen by comparing the figures of (A) the wild rock-pigeon, (B) the {164}shortfaced tumbler, (C) the English carrier, and (D) the Bagadotten carrier (of Neumeister), all drawn of the natural size and viewed laterally. In the carrier, besides the elongation of the bones of the face, the space between the orbits is proportionally a little narrower than in the rock-pigeon. In the Bagadotten the upper mandible is remarkably arched, and the premaxillary bones are proportionally broader. In the short-faced tumbler the skull is more globular; all the bones of the face are much shortened, and the front of the skull and descending nasal bones are almost perpendicular; the maxillo-jugal arch and premaxillary bones form an almost straight line; the space between the prominent edges of the eye-orbits is depressed. In the barb the premaxillary bones are much shortened, and their anterior portion is thicker than in the rock-pigeon, as is the lower part of the nasal bone. In two nuns the ascending branches of the premaxillaries, near their tips, were somewhat attenuated, and in these birds, as well as in some others, for instance in the spot, the occipital crest over the foramen was considerably more prominent than in the rock-pigeon.

Fig. 25.—Lower jaws, seen from above, of natural size. A. Rock-pigeon. B. Runt. C. Barb.

Fig. 26.—Skull of Runt, seen from above, of natural size, showing the reflexed margin of the distal portion of the lower jaw.

In the lower jaw, the articular surface is proportionally smaller in many breeds than in the rock-pigeon; and the vertical diameter more especially of the outer part of the articular surface is considerably shorter. May not this be accounted for by the lessened use of the jaws, owing to nutritious food having been given during a long period to all highly improved pigeons? In runts, carriers, and barbs (and in a lesser degree in several breeds), the whole side of the jaw near the articular end is bent inwards in a highly remarkable manner; and the superior margin of the ramus, beyond the middle, is reflexed in an equally remarkable manner, as may be seen in the accompanying figures, in comparison with the jaw of the rock-pigeon. This reflexion of the upper margin of the lower jaw is plainly connected with the singularly wide gape of the mouth, as has been described in runts, carriers, and barbs. The reflexion is well shown in fig. 26 of the head of a runt seen from above; here a wide open space may be observed on each side, between the edges of the lower jaw and of the premaxillary {165}bones. In the rock-pigeon, and in several domestic breeds, the edges of the lower jaw on each side come close up to the premaxillary bones, so that no open space is left. The degree of downward curvature of the distal half of the lower jaw also differs to an extraordinary degree in some breeds, as may be seen in the drawings (fig. A) of the rock-pigeon, (B) of the short-faced tumbler, and (C) of the Bagadotten carrier of Neumeister. In some runts the symphysis of the lower jaw is remarkably solid. No one would readily have believed that jaws differing so greatly in the several above-specified points could have belonged to the same species.

Fig. 27.—Lateral view of jaws, of natural size. A. Rock-pigeon. B. Short-faced Tumbler. C. Bagadotten Carrier.

Vertebræ.— All the breeds have twelve cervical vertebræ.^[310] But in a Bussorah carrier from India, the twelfth vertebra carried a small rib, a quarter of an inch in length, with a perfect double articulation.

The dorsal vertebræ are always eight. In the rock-pigeon all eight bear ribs; the eighth rib being very thin, and the seventh having no process. In pouters all the ribs are extremely broad, and, in three out of four skeletons examined by me, the eighth rib was twice or even thrice as broad as in the rock-pigeon; and the seventh pair had distinct processes. In many breeds there are only seven ribs, as in seven out of eight skeletons of various tumblers, and in several skeletons of fantails, turbits, and nuns. In all these breeds the seventh pair was very small, and was destitute of processes, in which respect it differed from the same rib in the rock-pigeon. In one tumbler, and in the Bussorah carrier, even the sixth pair had no process. The hypapophysis of the second dorsal vertebra varies much in development; being sometimes (as in several, but {166}not all tumblers) nearly as prominent as that of the third dorsal vertebra; and the two hypapophyses together tend to form an ossified arch. The development of the arch, formed by the hypapophyses of the third and fourth dorsal vertebræ, also varies considerably, as does the size of the hypapophysis of the fifth vertebra.

The rock-pigeon has twelve sacral vertebræ; but these vary in number, relative size, and distinctness in the different breeds. In pouters, with their elongated bodies, there are thirteen or even fourteen, and, as we shall immediately see, an additional number of caudal vertebræ. In runts and carriers there is generally the proper number, namely twelve; but in one runt, and in the Bussorah carrier, there were only eleven. In tumblers there are either eleven, twelve, or thirteen sacral vertebræ.

The caudal vertebræ are seven in number in the rock-pigeon. In fantails, which have their tails so largely developed, there are either eight or nine, and apparently in one case ten, and they are a little longer than in the rock-pigeon, and their shape varies considerably. Pouters, also, have eight or nine caudal vertebræ. I have seen eight in a nun and jacobin. Tumblers, though such small birds, always have the normal number seven; as have carriers, with one exception, in which there were only six.

The following table will serve as a summary, and will show the most remarkable deviations in the number of the vertebræ and ribs which I have observed:—

The pelvis differs very little in any breed. The anterior margin of the ilium, however, is sometimes a little more equally rounded on both sides than in the rock-pigeon, The ischium is also frequently rather more elongated. The obturator-notch is sometimes, as in many tumblers, less developed than in the rock-pigeon. The ridges on the ilium are very prominent in most runts.

In the bones of the extremities I could detect no difference, except in their proportional lengths; for instance, the metatarsus in a pouter was 1.65 inch, and in a short-faced tumbler only .95 in length; and this is a greater difference than would naturally follow from their differently-sized bodies; but long legs in the pouter, and small feet in the tumbler, are selected points. In some pouters the scapula is rather straighter, and in some {167}tumblers it is straighter, with the apex less elongated, than in the rock-pigeon: in the woodcut, fig. 28, the scapulæ of the rock-pigeon (A), and of a short-faced tumbler (B), are given. The processes at the summit of the coracoid, which receive the extremities of the furcula, form a more perfect cavity in some tumblers than in the rock-pigeon: in pouters these processes are larger and differently shaped, and the exterior angle of the extremity of the coracoid, which is articulated to the sternum, is squarer.

Fig. 29.—Furculæ, of natural size. A. Short-faced Tumbler B and C. Fantails. D. Pouter.

The two arms of the furcula in pouters diverge less, proportionally to their length, than in the rock-pigeon; and the symphysis is more solid and pointed. In fantails the degree of divergence of the two arms varies in a remarkable mariner. In fig. 29, B and C represent the furculæ of two fantails; and it will be seen that the divergence in B is rather less even than in the furcula of the short-faced, small-sized tumbler (A); whereas the divergence in C equals that in a rock-pigeon, or in the pouter (D), though the latter is a much larger bird. The extremities of the furcula, where articulated to the coracoids, vary considerably in outline.

In the sternum the differences in form are slight, except in the size and outline of the perforations, which, both in the larger and lesser sized breeds, are sometimes small. These perforations, also, are sometimes either nearly circular, or elongated, as is often the case with carriers. The posterior perforations occasionally are not complete, being left open posteriorly. The marginal apophyses forming the anterior perforations vary greatly in development. The degree of convexity of the posterior part of the sternum differs much, being sometimes almost perfectly flat. The manubrium is rather more prominent in some individuals than in others, and the pore immediately under it varies greatly in size.

Correlation of Growth.—By this term I mean that the whole organisation is so connected, that when one part varies, other {168}parts vary; but which of two correlated variations ought to be looked at as the cause and which as the effect, or whether both result from some common cause, we can seldom or never tell. The point of interest for us is that, when fanciers, by the continued selection of slight variations, have largely modified one part, they often unintentionally produce other modifications. For instance, the beak is readily acted on by selection, and, with its increased or diminished length, the tongue increases or diminishes, but not in due proportion; for, in a barb and short-faced tumbler, both of which have very short beaks, the tongue, taking the rock-pigeon as the standard of comparison, was proportionally not shortened enough, whilst in two carriers and in a runt the tongue, proportionally with the beak, was not lengthened enough. Thus, in a first-rate English carrier, in which the beak from the tip to the feathered base was exactly thrice as long as in a first-rate short-faced tumbler, the tongue was only a little more than twice as long. But the tongue varies in length independently of the beak: thus, in a carrier with a beak 1.2 inch in length, the tongue was .67 in length; whilst in a runt which equalled the carrier in length of body and in stretch of wings from tip to tip, the beak was .92 whilst the tongue was .73 of an inch in length, so that the tongue was actually longer than in the carrier with its long beak. The tongue of the runt was also very broad at the root. Of two runts, one had its beak longer by .23 of an inch, whilst its tongue was shorter by .14 than in the other.

With the increased or diminished length of the beak the length of the slit forming the external orifice of the nostrils varies, but not in due proportion, for, taking the rock-pigeon as the standard, the orifice in a short-faced tumbler was not shortened in due proportion with its very short beak. On the other hand (and this could not have been anticipated), the orifice in three English carriers, in the Bagadotten carrier, and in a runt (pigeon cygne), was longer by above the tenth of an inch than would follow from the length of the beak proportionally with that of the rock-pigeon. In one carrier the orifice of the nostrils was thrice as long as in the rock-pigeon, though in body and length of beak this bird was not nearly double the size of the {169}rock-pigeon. This greatly increased length of the orifice of the nostrils seems to stand partly in correlation with the enlargement of the wattled skin on the upper mandible and over the nostrils; and this is a character which is selected by fanciers. So again, the broad, naked, and wattled skin round the eyes of carriers and barbs is a selected character; and in obvious correlation with this, the eyelids, measured longitudinally, are proportionally more than double the length of those of the rock-pigeon.

The great difference (see woodcut No. 27) in the curvature of the lower jaw in the rock-pigeon, the tumbler, and Bagadotten carrier, stands in obvious relation to the curvature of the upper jaw, and more especially to the angle formed by the maxillo-jugal arch with the premaxillary bones. But in carriers, runts, and barbs the singular reflexion of the upper margin of the middle part of the lower jaw (see woodcut No. 25) is not strictly correlated with the width or divergence (as may be clearly seen in woodcut No. 26) of the premaxillary bones, but with the breadth of the horny and soft parts of the upper mandible, which are always overlapped by the edges of the lower mandible.

In pouters, the elongation of the body is a selected character, and the ribs, as we have seen, have generally become very broad, with the seventh pair furnished with processes; the sacral and caudal vertebræ have been augmented in number; the sternum has likewise increased in length (but not in the depth of the crest) by .4 of an inch more than would follow from the greater bulk of the body in comparison with that of the rock-pigeon. In fantails, the length and number of the caudal vertebræ have increased. Hence, during the gradual progress of variation and selection, the internal bony frame-work and the external shape of the body have been, to a certain extent, modified in a correlated manner.

Although the wings and tail often vary in length independently of each other, it is scarcely possible to doubt that they generally tend to become elongated or shortened in correlation. This is well seen in jacobins, and still more plainly in runts, some varieties of which have their wings and tail of great length, whilst others have both very short. With jacobins, the remarkable length of the tail and {170}wing-feathers is not a character which is intentionally selected by fanciers; but fanciers have been trying for centuries, at least since the year 1600, to increase the length of the reversed feathers on the neck, so that the hood may more completely enclose the head; and it may be suspected that the increased length of the wing and tail-feathers stands in correlation with the increased length of the neck-feathers. Short-faced tumblers have short wings in nearly due proportion with the reduced size of their bodies; but it is remarkable, seeing that the number of the primary wing-feathers is a constant character in most birds, that these tumblers generally have only nine instead of ten primaries. I have myself observed this in eight birds; and the Original Columbarian Society^[311] reduced the standard for bald-head tumblers from ten to nine white flight-feathers, thinking it unfair that a bird which had only nine feathers should be disqualified for a prize because it had not ten white flight-feathers. On the other hand, in carriers and runts, which have large bodies and long wings, eleven primary feathers have occasionally been observed.

Mr. Tegetmeier has informed me of a curious and inexplicable case of correlation, namely, that young pigeons of all breeds, which when mature become white, yellow, silver (i.e. extremely pale blue), or dun-coloured, are born almost naked; whereas other coloured pigeons are born well clothed with down. Mr. Esquilant, however, has observed that young dun carriers are not so bare as young dun barbs and tumblers. Mr. Tegetmeier has seen two young birds in the same nest, produced from differently coloured parents, which differed greatly in the degree to which they were at first clothed with down.

I have observed another case of correlation which at first sight appears quite inexplicable, but on which, as we shall see in a future chapter, some light can be thrown by the law of homologous parts varying in the same manner. The case is, that, when the feet are much feathered, the roots of the feathers are connected by a web of skin, and apparently in correlation with this the two outer toes become connected for a considerable space by skin. I have observed this in very many {171}specimens of pouters, trumpeters, swallows, roller-tumblers (likewise observed in this breed by Mr. Brent), and in a lesser degree in other feather-footed pigeons.

The feet of the smaller and larger breeds are of course much smaller or larger than those of the rock-pigeon; but the scutellæ or scales covering the toes and tarsi have not only decreased or increased in size, but likewise in number. To give a single instance, I have counted eight scutellæ on the hind toe of a runt, and only five on that of a short-faced tumbler. With birds in a state of nature the number of the scutellæ on the feet is usually a constant character. The length of the feet and the length of the beak apparently stand in correlation; but as disuse apparently has affected the size of the feet, this case may come under the following discussion.

On the Effects of Disuse.—In the following discussion on the relative proportions of the feet, sternum, furcula, scapulæ, and wings, I may premise, in order to give some confidence to the reader, that my measurements were all made in the same manner, and that all the measurements of the external parts were made without the least intention of applying them to the following purpose.

I measured most of the birds which came into my possession, from the feathered base of the beak (the length of beak itself being so variable) to the end of the tail, and to the oil-gland, but unfortunately (except in a few cases) not to the root of the tail; I measured each bird from the extreme tip to tip of wing; and the length of the terminal folded part of the wing, from the extremity of the primaries to the joint of the radius. I measured the feet without the claws, from the end of the middle toe to the end of the hind toe; and the tarsus together with the middle toe. I have taken in every case the mean measurement of two wild rock-pigeons from the Shetland Islands, as the standard of comparison. The following table shows the actual length of the feet in each bird; and the difference between the length which the feet ought to have had according to the size of body of each, in comparison with the size of body and length of feet of the rock-pigeon, calculated (with a few specified exceptions) by the standard of the length of the body from the base of the beak to the oil-gland. I have preferred this standard, owing to the variability of the length of tail. But I have made similar calculations, taking as the standard the length from tip to tip of wing, and likewise in most cases from the base of the beak to the end of the tail; and the result has always been closely similar. To give an example: the first bird in the table, being a short-faced tumbler, {172}is much smaller than the rock-pigeon, and would naturally have shorter feet; but it is found on calculation to have feet too short by .11 of an inch, in comparison with the feet of the rock-pigeon, relatively to the size of the body in these two birds, as measured from the base of beak to the oil-gland. So again, when this same tumbler and the rock-pigeon were compared by the length of their wings, or by the extreme length of their bodies, the feet of the tumbler were likewise found to be too short in very nearly the same proportion. I am well aware that the measurements pretend to greater accuracy than is possible, but it was less trouble to write down the actual measurements given by the compasses in each case than an approximation.

Table I.

Pigeons with their beaks generally shorter than that of the Rock-pigeon, proportionally with the size of their bodies.

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Table II.

Pigeons with their beaks longer than that of the Rock-pigeon, proportionally with the size of their bodies.

In these two tables we see in the first column the actual length of the feet in thirty-six birds belonging to various breeds, and in the two other columns we see by how much the feet are too short or too long, according to the size of bird, in comparison with the rock-pigeon. In the first table twenty-two specimens have their feet too short, on an average by a little above the tenth of an inch (viz. .107); and five specimens have their feet on an average a very little too long, namely, by .07 of an inch. But some of these latter and exceptional cases can be explained; for instance, with pouters the legs and feet are selected for length, and thus any natural tendency to a diminution in the length of the feet will have been counteracted. In the swallow and barb, when the calculation was made on any standard of comparison excepting the one above used (viz. length of body from base of beak to oil-gland), the feet were found to be too small.

In the second table we have eight birds, with their beaks much longer than in the rock-pigeon, both actually and proportionally with the size of body, and their feet are in an equally marked manner longer, namely, in proportion, on an average by .29 of an inch. I should here state that in Table I. there are a few partial exceptions to the beak being proportionally shorter than in the rock-pigeon: thus the beak of the English frill-back is just perceptibly longer, and that of the Bussorah carrier of the same length or slightly longer, than in the rock-pigeon. The beaks of spots, swallows, and laughers are only a very little shorter, or of the same proportional length, but slenderer. Nevertheless, these two tables, taken conjointly, indicate pretty plainly some kind of correlation between the length of the beak and the size of the feet. Breeders of cattle and horses believe that there is an analogous connection between the length of the limbs and head; they assert that a race-horse with the head of a dray-horse, or a {174}greyhound with the head of a bulldog, would be a monstrous production. As fancy pigeons are generally kept in small aviaries, and are abundantly supplied with food, they must walk about much less than the wild rock-pigeon; and it may be admitted as highly probable that the reduction in the size of the feet in the twenty-two birds in the first table has been caused by disuse,^[312] and that this reduction has acted by correlation on the beaks of the great majority of the birds in Table I. When, on the other hand, the beak has been much elongated by the continued selection of successive slight increments of length, the feet by correlation have likewise become much elongated in comparison with those of the wild rock-pigeon, notwithstanding their lessened use.

As I had taken measures from the end of the middle toe to the heel of the tarsus in the rock-pigeon and in the above thirty-six birds, I have made calculations analogous with those above given, and the result is the same,—namely, that in the short-beaked breeds, with equally few exceptions as in the former case, the middle toe conjointly with the tarsus has decreased in length; whereas in the long-beaked breeds it has increased in length, though not quite so uniformly as in the former case, for the leg in some varieties of the runt varies much in length.

As fancy pigeons are generally confined in aviaries of moderate size, and as even when not confined they do not search for their own food, they must during many generations have used their wings incomparably less than the wild rock-pigeon. Hence it seemed to me probable that all the parts of the skeleton subservient to flight would be found to be reduced in size. With respect to the sternum, I have carefully measured its extreme length in twelve birds of different breeds, and in two wild rock-pigeons from the Shetland Islands. For the proportional comparison I have tried with all twelve birds three standards of measurement, namely, the length from the base of the beak to the oil-gland, to the end of the tail, and from the extreme tip to tip of wings. The result has been in each case nearly the same, the sternum being invariably found to be shorter than in the wild rock-pigeon. I will give only a single table, as calculated by the standard from the base of the beak to the oil-gland; for the result in this case is nearly the mean between the results obtained by the two other standards.

Length of Sternum.

{175}

This table shows that in these twelve breeds the sternum is on an average one-third of an inch (exactly .332) shorter than in the rock-pigeon, proportionally with the size of their bodies; so that the sternum has been reduced by between one-seventh and one-eighth of its entire length; and this is a considerable reduction.

I have also measured in twenty-one birds, including the above dozen, the prominence of the crest of the sternum relatively to its length, independently of the size of the body. In two of the twenty-one birds the crest was prominent in the same relative degree as in the rock-pigeon; in seven it was more prominent; but in five out of these seven, namely, in a fantail, two scanderoons, and two English carriers, this greater prominence may to a certain extent be explained, as a prominent breast is admired and selected by fanciers; in the remaining twelve birds the prominence was less. Hence it follows that the crest exhibits a slight, though uncertain, tendency to become reduced in prominence in a greater degree than does the length of the sternum relatively to the size of body, in comparison with the rock-pigeon.

I have measured the length of the scapula in nine different large and small-sized breeds, and in all the scapula is proportionally shorter (taking the same standard as before) than in the wild rock-pigeon. The reduction in length on an average is very nearly one-fifth of an inch, or about one-ninth of the length of the scapula in the rock-pigeon.

The arms of the furcula in all the specimens which I compared, diverged less, proportionally with the size of body, than in the rock-pigeon; and the whole furcula was proportionally shorter. Thus in a runt, which measured from tip to tip of wings 38½ inches, the furcula was only a very little longer (with the arms hardly more divergent) than in a rock-pigeon which measured from tip to tip 26½ inches. In a barb, which in all its measurements was a little larger than the same rock-pigeon, the furcula was a quarter of an inch shorter. In a pouter, the furcula had not been lengthened proportionally with the increased length of the body. In a short-faced tumbler, which measured from tip to tip of wings 24 inches, therefore only 2½ inches less than the rock-pigeon, the furcula was barely two-thirds of the length of that of the rock-pigeon.

We thus clearly see that the sternum, scapulæ, and furcula are all reduced in proportional length; but when we turn to the wings we find what at first appears a wholly different and unexpected result. I may here remark that I have not picked out specimens, but have used every measurement made by me. Taking the length from the base of beak to the end of the tail as the standard of comparison, I find that, out of thirty-five birds of various breeds, twenty-five have wings of greater, and ten have them of less proportional length, than in the rock-pigeon. But from the frequently correlated length of the tail and wing-feathers, it is better to take as the standard {176}of comparison the length from the base of the beak to the oil-gland; and by this standard, out of twenty-six of the same birds which had been thus measured, twenty-one had wings too long, and only five had them too short. In the twenty-one birds the wings exceeded in length those of the rock-pigeon, on an average, by 1⅓ inch; whilst in the five birds they were less in length by only .8 of an inch. As I was much surprised that the wings of closely confined birds should thus so frequently have been increased in length, it occurred to me that it might be solely due to the greater length of the wing-feathers; for this certainly is the case with the jacobin, which has wings of unusual length. As in almost every case I had measured the folded wings, I subtracted the length of this terminal part from that of the expanded wings, and thus I obtained, with a moderate degree of accuracy, the length of the wings from the ends of the two radii, answering from wrist to wrist in our arms. The wings, thus measured in the same twenty-five birds, now gave a widely different result; for they were proportionally with those of the rock-pigeon too short in seventeen birds, and in only eight too long. Of these eight birds, five were long-beaked,^[313] and this fact perhaps indicates that there is some correlation between the length of the beak and the length of the bones of the wings, in the same manner as with the feet and tarsi. The shortening of the humerus and radius in the seventeen birds may probably be attributed to disuse, as in the case of the scapulæ and furcula to which the wing-bones are attached;—the lengthening of the wing-feathers, and consequently the expansion of the wings from tip to tip, being, on the other hand, as completely independent of use and disuse as is the growth of the hair or wool on our long-haired dogs or long-woolled sheep.

To sum up: we may confidently admit that the length of the sternum, and frequently the prominence of its crest, the length of the scapulæ and furcula, have all been reduced in size in comparison with the same parts in the rock-pigeon. And I {177}presume that this may be safely attributed to disuse or lessened exercise. The wings, as measured from the ends of the radii, have likewise been generally reduced in length; but, owing to the increased growth of the wing-feathers, the wings, from tip to tip, are commonly longer than in the rock-pigeon. The feet, as well as the tarsi conjointly with the middle toe, have likewise in most cases become reduced; and this it is probable has been caused by their lessened use; but the existence of some sort of correlation between the feet and beak is shown more plainly than the effects of disuse. We have also some faint indication of a similar correlation between the main bones of the wing and the beak.

Summary on the Points of Difference between the several Domestic Races, and between the individual Birds.—The beak, together with the bones of the face, differ remarkably in length, breadth, shape, and curvature. The skull differs in shape, and greatly in the angle formed by the union of the premaxillary, nasal, and maxillo-jugal bones. The curvature of the lower jaw and the reflexion of its upper margin, as well as the gape of the mouth, differ in a highly remarkable manner. The tongue varies much in length, both independently and in correlation with the length of the beak. The development of the naked, wattled skin over the nostrils and round the eyes varies in an extreme degree. The eyelids and the external orifices of the nostrils vary in length, and are to a certain extent correlated with the degree of development of the wattle. The size and form of the œsophagus and crop, and their capacity for inflation, differ immensely. The length of the neck varies. With the varying shape of the body, the breadth and number of the ribs, the presence of processes, the number of the sacral vertebræ, and the length of the sternum, all vary. The number and size of the coccygeal vertebræ vary, apparently in correlation with the increased size of the tail. The size and shape of the perforations in the sternum, and the size and divergence of the arms of the furcula, differ. The oil-gland varies in development, and is sometimes quite aborted. The direction and length of certain feathers have been much modified, as in the hood of the Jacobin and the frill of the Turbit. The wing and tail feathers generally vary in {178}length together, but sometimes independently of each other and of the size of the body. The number and position of the tail-feathers vary to an unparalleled degree. The primary and secondary wing-feathers occasionally vary in number, apparently in correlation with the length of the wing. The length of the leg and the size of the feet, and, in connection with the latter, the number of the scutellæ, all vary. A web of skin sometimes connects the bases of the two inner toes, and almost invariably the two outer toes when the feet are feathered.

The size of the body differs greatly: a runt has been known to weigh more than five times as much as a short-faced tumbler. The eggs differ in size and shape. According to Parmentier,^[314] some races use much straw in building their nests, and others use little; but I cannot hear of any recent corroboration of this statement. The length of time required for hatching the eggs is uniform in all the breeds. The period at which the characteristic plumage of some breeds is acquired, and at which certain changes of colour supervene, differs. The degree to which the young birds are clothed with down when first hatched is different, and is correlated in a singular manner with the future colour of the plumage. The manner of flight, and certain inherited movements, such as clapping the wings, tumbling either in the air or on the ground, and the manner of courting the female, present the most singular differences. In disposition the several races differ. Some races are very silent; others coo in a highly peculiar manner.

Although many different races have kept true in character during several centuries, as we shall hereafter more fully see, yet there is far more individual variability in the truest breeds than in birds in a state of nature. There is hardly any exception to the rule that those characters vary most which are now most valued and attended to by fanciers, and which consequently are now being improved by continued selection. This is indirectly admitted by fanciers when they complain that it is much more difficult to breed high fancy pigeons up to the proper standard of excellence than the so-called toy pigeons, which differ from {179}each other merely in colour; for particular colours when once acquired are not liable to continued improvement or augmentation. Some characters become attached, from quite unknown causes, more strongly to the male than to the female sex; so that we have, in certain races, a tendency towards the appearance of secondary sexual characters,^[315] of which the aboriginal rock-pigeon displays not a trace.

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CHAPTER VI.

PIGEONS—continued.

ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES—HABITS OF LIFE—WILD RACES OF THE ROCK-PIGEON—DOVECOT-PIGEONS—PROOFS OF THE DESCENT OF THE SEVERAL RACES FROM COLUMBA LIVIA—FERTILITY OF THE RACES WHEN CROSSED—REVERSION TO THE PLUMAGE OF THE WILD ROCK-PIGEON—CIRCUMSTANCES FAVOURABLE TO THE FORMATION OF THE RACES—ANTIQUITY AND HISTORY OF THE PRINCIPAL RACES—MANNER OF THEIR FORMATION—SELECTION—UNCONSCIOUS SELECTION—CARE TAKEN BY FANCIERS IN SELECTING THEIR BIRDS—SLIGHTLY DIFFERENT STRAINS GRADUALLY CHANGE INTO WELL-MARKED BREEDS—EXTINCTION OF INTERMEDIATE FORMS—CERTAIN BREEDS REMAIN PERMANENT, WHILST OTHERS CHANGE—SUMMARY.

The differences described in the last chapter between the eleven chief domestic races and between individual birds of the same race, would be of little significance, if they had not all descended from a s