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.

{49}

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.

{50}

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.

{62}

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.

{65}

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.

{66}

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]

{127}

{128}

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.

{131}

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.

{136}

{137}

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:—