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

Author: Charles Darwin

Release date: March 27, 2008 [eBook #24923]

Language: English

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*** START OF THE PROJECT GUTENBERG EBOOK THE
VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION,
VOL. I. ***

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THE VARIATION
OF

ANIMALS AND PLANTS

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

IN TWO VOLUMES.—Vol. I.

WITH ILLUSTRATIONS.

LONDON:

JOHN MURRAY, ALBEMARLE STREET.

1868.

The right of Translation is reserved.

BY THE SAME AUTHOR.

ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL
SELECTION; or The Preservation of Favoured Races in the Struggle for
Life. Fourth Edition (Eighth Thousand), with Additions and Corrections.
1866. ... Murray.
A NATURALIST'S VOYAGE ROUND THE WORLD; or, A Journal of
Researches into the Natural History and Geology of the Countries visited
during the Voyage of H.M.S. Beagle, under the Command of Capt. Fitz-
Roy, R.N. Tenth Thousand. ... Murray.
ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. ...
Smith, Elder, & Co.

GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. ... Smith,
Elder, & Co.
GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. ... Smith,
Elder, & Co.
A MONOGRAPH OF THE CIRRIPEDIA. With numerous Illustrations. 2
vols. 8vo. ... Hardwicke.
ON THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND
FOREIGN ORCHIDS ARE FERTILISED BY INSECTS; and on the Good
Effects of Crossing. With numerous Woodcuts. ... Murray.
ON THE MOVEMENTS and HABITS of CLIMBING PLANTS. With
Woodcuts. ... Williams & Norgate.

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

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.

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

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

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

LIST OF ILLUSTRATIONS.

1. Dun Devonshire Pony, with shoulder, spinal, and leg
stripes ... PAGE 56
2. Head of Japan or Masked Pig ... 69
3. Head of Wild Boar, and of "Golden Days," a pig of the
Yorkshire large breed ... 72
4. Old Irish Pig, with jaw-appendages ... 75
5. Half-lop Rabbit ... 108
6. Skull of Wild Rabbit ... 117
7. Skull of large Lop-eared Rabbit ... 117
8. Part of Zygomatic Arch, showing the projecting end of
the malar-bone, and the auditory meatus, of
Rabbits ... 118
9. Posterior end of Skull, showing the inter-parietal bone,
of Rabbits ... 118
10. Occipital Foramen of Rabbits ... 118
11. Skull of Half-lop Rabbit ... 119
12. Atlas Vertebræ of Rabbits ... 121
13. Third Cervical Vertebræ of Rabbits ... 121
14. Dorsal Vertebræ, from sixth to tenth inclusive, of
Rabbits ... 122

15. Terminal Bone of Sternum of Rabbits ... 123
16. Acromion of Scapula of Rabbits ... 123
17. The Rock-Pigeon, or Columbia Livia ... 135
18. English Pouter ... 137
19. English Carrier ... 140
20. English Barb ... 145
21. English Fantail ... 147
22. African Owl ... 149
23. Short-faced English Tumbler ... 152
24. Skulls of Pigeons, viewed laterally ... 163
25. Lower Jaws of Pigeons, seen from above ... 164
26. Skull of Runt, seen from above ... 165
27. Lateral view of Jaws of Pigeons ... 165
28. Scapulæ of Pigeons ... 167
29. Furculæ of Pigeons ... 167
30. Spanish Fowl ... 226
31. Hamburgh Fowl ... 228
32. Polish Fowl ... 229
33. Occipital Foramen of the Skulls of Fowls ... 261
34. Skulls of Fowls, viewed from above, a little obliquely
... 262
35. Longitudinal sections of Skulls of Fowls, viewed
laterally ... 263
36. Skull of Horned Fowl, viewed from above, a little
obliquely ... 265
37. Sixth Cervical Vertebræ of Fowls, viewed laterally ...
267
38. Extremity of the Furcula of Fowls, viewed laterally
... 268
39. Skulls of Ducks, viewed laterally, reduced to two-
thirds of the natural size ... 282
40. Cervical Vertebræ of Ducks, of natural size ... 283
41. Pods of the Common Pea ... 328
42. Peach and Almond Stones, of natural size, viewed
edgeways ... 337
43. Plum Stones, of natural size, viewed laterally ... 345

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 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, 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 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 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 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.
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 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 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 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 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 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 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, because it
collects under one point of view, and gives a rational explanation of, many
apparently independent classes of facts.[4]

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 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 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, 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 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 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
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 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]
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 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 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 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 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.
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 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 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 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,
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 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 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 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 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 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 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 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 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 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 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 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 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 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 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 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, 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.

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.
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 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 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 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 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, 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, 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 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 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 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 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 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.

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 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 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.

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.
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 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 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 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 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 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] 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.
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 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 Fig. 3.—Head of Wild Boar, and of
"Golden Days," a pig of the Yorkshire
the result of selection has been divergence of character, here it Large Breed; the latter from a
has been convergence.[162] photograph. (Copied from Sidney's
edit. of 'The Pig,' by Youatt.)
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 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.

French
English Domestic
Long- Wild Boar, Boar,
legged African Chinese from from
Male. Female. Male. Cuvier. Cuvier.

Dorsal vertebræ 15 13 15 14 14

Lumbar 6 6 4 5 5

Dorsal and lumbar
21 19 19 19 19
together

Sacral 5 5 4 4 4

Total number of vertebræ 26 24 23 23 23

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 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 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
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 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
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 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 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 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 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 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 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 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 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 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
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 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 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, 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 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 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 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 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:—

Merinos 150.3 days.
Southdowns 144.2 "
Half-bred Merinos and Southdowns 146.3 "
¾ blood of Southdown 145.5 "
⅞ blood of Southdown 144.2 "

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 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 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 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 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
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!

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. 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 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 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 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 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. 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 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.
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 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 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 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, 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 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.

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 Fig. 6.—Skull of Wild
examined skulls of ten large Rabbit, of natural size.
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 Fig. 7.—Skull of large Lop-
ears, yet both larger than in the wild rabbit. First for the eared Rabbit, of natural size.
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.

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 Fig. 8.—Part of
muscles; consequently the brain, as we shall presently more Zygomatic Arch, showing
fully see, has not increased relatively with the size of body. the projecting end of the
malar bone and the
As the brain has not increased, the bony case enclosing it has auditory meatus: of
not increased, and this has evidently affected through natural size. Upper figure,
correlation the breadth of the entire skull from end to end. Wild Rabbit. Lower
figure, Lop-eared, hare-
coloured Rabbit.
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 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.

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
Fig. 9.—Posterior end of
4.3 inches in length, and which Fig. 10.—Occipital
Foramen, of natural size, in—
Skull, of natural size, barely exceeded in breadth the A. Wild Rabbit; B. Large Lop-
showing the inter-parietal skull of a wild rabbit (which eared Rabbit.
bone. A. Wild Rabbit. B.
Feral Rabbit from island of P. was 3.15 inches in length), the
Santo, near Madeira. C. longer diameter of the meatus was exactly twice as great.
Large Lop-eared Rabbit. 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 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!

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 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.
Fig. 11.—Skull, of natural
The five other skulls of common domestic rabbits, some of size, of Half-lop Rabbit,
which approach in size the above-described largest skulls, showing the different
direction of the auditory
whilst the others exceed but little those of the wild rabbit, meatus on the two sides, and
are only worth notice as presenting a perfect gradation in the consequent general
all the above-specified differences between the skulls of distortion of the skull. The
left ear of the animal (or
the largest lop-eared and wild rabbits. In all, however, the right side of figure) lopped
supra-orbital plates are rather larger, and in all the auditory forwards.
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 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.
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 Fig. 12.—Atlas
of the antero-dorsal processes in the wild rabbit are simply Vertebræ, of natural
size; inferior surface
rounded, whilst in the lop-eared they are trifid, with a deep viewed obliquely.
central pit. The canal for the spinal marrow in the lop-eared (B Upper figure, Wild
b) is more elongated in a transverse direction than in the wild Rabbit. Lower figure,
Hare-coloured, large,
rabbit; and the passages for the arteries are of a slightly different Lop-eared Rabbit. a,
shape. These several differences in this vertebra seem to me supra-median,
well deserving attention. atlantoid process; b,
infra-median process.
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 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
Fig. 13.—Third Cervical thicker, and of somewhat different shape, in comparison
Vertebra, of natural size, of—A. with those of the wild rabbit. So that this part of the
Wild Rabbit; B. Hare-coloured, vertebral column differs considerably in appearance
large, Lop-eared Rabbit. a, a,
inferior surface; b, b, anterior from the same part in the wild rabbit, and closely
articular surfaces. 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.
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
Fig. 15.— Porto Santo feral rabbits, had a
Terminal bone of hæmal spine; whilst in four
Sternum, of natural
size. A. Wild
skeletons of large lop-eared
Rabbit. B. Hare- rabbits, and in the Himalayan Fig. 14.—Dorsal Vertebræ, from
coloured, Lop- rabbit, this same vertebra had a sixth to tenth inclusive, of natural size,
eared Rabbit. C. viewed laterally. A. Wild Rabbit. B.
Hare-coloured,
well-developed hæmal spine.
Large, Hare-coloured, so called
Spanish Rabbit. Spanish Rabbit.
(N.B. The left- Pelvis.—In four wild specimens
hand angle of the this bone was almost absolutely
upper articular
extremity of B was
identical in shape; but in several domesticated breeds shades of
broken, and has differences could be distinguished. In the large lop-eared rabbits
been accidentally the whole upper part of the ilium is straighter, or less splayed
thus represented.)
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.
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 Fig. 16.—Acromion of
Scapula, of natural size. A.
acromion varies much in breadth and sharpness, as may Wild Rabbit. B, C, D; Large,
be seen by comparing figs. B, C, and D. Lop-eared Rabbits.

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
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 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 (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]

Name of I. II. III. IV. V. VI. VII.
Breed. Length Length Weight Capacity Capacity Difference Showing
Wild and of of of of Skull calculated between how much
Semi-wild Skull. Body whole measured according actual and per cent.
Rabbits. from Body. by Small to Length calculated the Brain,
Incisors Shot. of Skull capacities by
to relatively of Skulls. calculation,
Anus. to that of according
No. 1. to the
length of
the Skull,
is too light
or too
heavy,
relatively
to the
Brain of
the Wild
Rabbit No.
1.

lbs.
inches. inches. grains. grains. grains.
ozs.
1. Wild
rabbit, 3.15 17.4 3 5 972 .. ..
Kent
2. "
3.15 .. .. 979 .. ..
Shetland
Islands
3. " 3.15 .. .. 992 .. .. [2 per cent.
too heavy
Ireland in
comparison

with No.
1.]
4.
Domestic
rabbit, run 3.15 18.5 .. 977 .. ..
wild,
Sandon
5. Wild,
common
4 per
variety,
2.96 17.0 2 14 875 913 38 cent. too
small
light.
Specimen,
Kent
6. Wild,
fawn-
3 "
coloured 3.1 .. .. 918 950 32
"
variety,
Scotland
7. Silver-
grey, small
3 "
specimen, 2.95 15.5 2 11 938 910 28
too heavy.
Thetford
warren
8. Feral
rabbit, 2 "
2.83 .. .. 893 873 20
Porto "
Santo
9. " 16 "
2.85 .. .. 756 879 123
" too light.
10. " 9 "
2.95 .. .. 835 910 75
" "

Average
of the
three 7 "
2.88 .. .. 828 888 60
Porto "
Santo
Rabbits

Domestic
Rabbits

11. 12 "
3.5 20.5 .. 963 1080 117
Himalayan "

12. 24 "
3.25 17.0 3 8 803 1002 199
Moscow "
13. 54 "
3.5 19.5 3 1 697 1080 383
Angora "
14. 13 "
3.65 22.0 .. 995 1126 131
Chinchilla "
15. Large 18 "
4.1 24.5 7 0 1065 1265 200
lop-eared "
16. " 9 "
4.1 25.0 7 13 1153 1265 112
" "
17. " 21 "
4.07 .. .. 1037 1255 218
" "
18. " 4 "
4.1 25.0 7 4 1208 1265 57
" "
19. " 7 "
4.3 .. .. 1232 1326 94
" "
20. " 16 "
4.25 .. .. 1124 1311 187
" "
21. Large
5 "
hare- 3.86 24.0 6 14 1131 1191 60
"
coloured
22.
Average
of above
11 "
seven 4.11 24.62 7 4 1136 1268 132
"
large lop-
eared
rabbits
23. Hare
(L.
timidus) 3.61 7 0 1315
English
specimen
24. "
"
3.82 7 0 1455
German
specimen

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 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 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.

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 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 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 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 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.

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.

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 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 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
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 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 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, 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 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 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 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 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. 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.

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
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 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 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 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 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. 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 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 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
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 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] 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 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]

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 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 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 Fig. 24.—Skulls of Pigeons, viewed
the rock-pigeon. In the Bagadotten the laterally, of natural size. A. Wild Rock-
upper mandible is remarkably arched, pigeon, Columba livia. B. Short-faced
Tumbler. C. English Carrier. D.
and the premaxillary bones are Bagadotten Carrier.
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.

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 Fig. 26.—
remarkable manner, as may be seen in the accompanying Skull of Runt,
figures, in comparison with the jaw of the rock-pigeon. seen from
above, of natural
This reflexion of the upper margin of the lower jaw is size, showing
plainly connected with the singularly wide gape of the the reflexed
mouth, as has been described in runts, carriers, and barbs. margin of the
distal portion of
The reflexion is well shown in fig. 26 of the head of a the lower jaw.
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 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.

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
Fig. 27.—Lateral view of being very thin, and the seventh having no
jaws, of natural size. A.
Rock-pigeon. B. Short-faced process. In pouters all the ribs are extremely
Tumbler. C. Bagadotten broad, and, in three out of four skeletons
Carrier.
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 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:—

Pouter, Tumbler,
Rock Bussorah
from Mr. Dutch
Pigeon. Carrier.
Bult. Roller.

Cervical 12 12 12 12
Vertebræ The 12th
bore a
small rib.

Dorsal 8 8 8 8
Vertebræ
" 8 8 7 7
Ribs The 6th The 6th The 6th The 6th
pair with & 7th & 7th & 7th
processes, pair with pair pair
the 7th processes. without without
pair processes. processes.
without a
process.

Sacral 12 14 11 11
Vertebræ
Caudal 7 8 or 9 7 7
Vertebræ

Total
Vertebræ 39 42 or 43 38 38

The pelvis differs very little in any breed. The anterior margin of the
ilium, however, is sometimes a little more equally rounded on both
sides than in the rock-pigeon, The ischium is also frequently rather
more elongated. The obturator-notch is sometimes, as in many
tumblers, less developed than in the rock-pigeon. The ridges on the
ilium are very prominent in most runts.
In the bones of the extremities I could detect no difference, except in
their proportional lengths; for instance, the metatarsus in a pouter
was 1.65 inch, and in a short-faced tumbler only .95 in length; and
this is a greater difference than would naturally follow from their
differently-sized bodies; but long legs in the pouter, and small feet in
the tumbler, are selected points. In some pouters the scapula is rather
straighter, and in some tumblers it is straighter, with the apex less
elongated, than in the rock-pigeon: in the woodcut, fig. 28, the
scapulæ of the rock-pigeon (A), and of a short-faced tumbler (B), are
given. The processes at the summit of the coracoid, which receive
the extremities of the furcula, form a more perfect cavity in some
tumblers than in the rock-pigeon: in pouters these processes are
larger and differently shaped, and the exterior angle of the extremity
of the coracoid, which is articulated to the sternum, is squarer.
The two arms of the furcula in pouters diverge less, proportionally to
their length, than in the rock-pigeon; and the symphysis is more solid
and pointed. In fantails the degree of divergence of the two arms
varies in a remarkable mariner. In fig. 29, B and C represent the
furculæ of two fantails; and it will be seen that the divergence in B is
rather less even than in the furcula of the short-faced, small-sized
tumbler (A); whereas the divergence in C equals that in a rock-
pigeon, or in the pouter (D), though the latter is a much larger bird.
The extremities of the furcula, where articulated to the coracoids,
vary considerably in outline.
In the sternum the differences in form are slight, except in the size
and outline of the perforations, which, both in the larger and lesser
sized breeds, are sometimes small. These perforations, also, are

sometimes either nearly circular, or elongated, as is
often the case with carriers. The posterior perforations
occasionally are not complete, being left open
posteriorly. The marginal apophyses forming the
anterior perforations vary greatly in development. The
degree of convexity of the posterior part of the
sternum differs much, being sometimes almost
perfectly flat. The manubrium is rather more
prominent in some individuals than in others, and the
pore immediately under it varies greatly in size.

Correlation of Growth.—By this term I mean that the
whole organisation is so connected, that when one part
varies, other parts vary; but which of two correlated
variations ought to be looked at as the cause and which as
the effect, or whether both result from some common cause,
we can seldom or never tell. The point of interest for us is
that, when fanciers, by the continued selection of slight Fig. 29.—
Furculæ, of
variations, have largely modified one part, they often natural size.
unintentionally produce other modifications. For instance, A. Short-
the beak is readily acted on by selection, and, with its faced
increased or diminished length, the tongue increases or Tumbler and C.
B

diminishes, but not in due proportion; for, in a barb and Fantails. D.
short-faced tumbler, both of which have very short beaks, Pouter.
the tongue, taking the rock-pigeon as the standard of
comparison, was proportionally not shortened enough, whilst in two carriers
and in a runt the tongue, proportionally with the beak, was not lengthened
enough. Thus, in a first-rate English carrier, in which the beak from the tip to
the feathered base was exactly thrice as long as in a first-rate short-faced
tumbler, the tongue was only a little more than twice as long. But the tongue
varies in length independently of the beak: thus, in a carrier with a beak 1.2
inch in length, the tongue was .67 in length; whilst in a runt which equalled the
carrier in length of body and in stretch of wings from tip to tip, the beak was
.92 whilst the tongue was .73 of an inch in length, so that the tongue was
actually longer than in the carrier with its long beak. The tongue of the runt
was also very broad at the root. Of two runts, one had its beak longer by .23 of
an inch, whilst its tongue was shorter by .14 than in the other.

With the increased or diminished length of the beak the length of the slit
forming the external orifice of the nostrils varies, but not in due proportion,
for, taking the rock-pigeon as the standard, the orifice in a short-faced tumbler
was not shortened in due proportion with its very short beak. On the other
hand (and this could not have been anticipated), the orifice in three English
carriers, in the Bagadotten carrier, and in a runt (pigeon cygne), was longer by
above the tenth of an inch than would follow from the length of the beak
proportionally with that of the rock-pigeon. In one carrier the orifice of the
nostrils was thrice as long as in the rock-pigeon, though in body and length of
beak this bird was not nearly double the size of the rock-pigeon. This greatly
increased length of the orifice of the nostrils seems to stand partly in
correlation with the enlargement of the wattled skin on the upper mandible and
over the nostrils; and this is a character which is selected by fanciers. So again,
the broad, naked, and wattled skin round the eyes of carriers and barbs is a
selected character; and in obvious correlation with this, the eyelids, measured
longitudinally, are proportionally more than double the length of those of the
rock-pigeon.
The great difference (see woodcut No. 27) in the curvature of the lower jaw in
the rock-pigeon, the tumbler, and Bagadotten carrier, stands in obvious relation
to the curvature of the upper jaw, and more especially to the angle formed by
the maxillo-jugal arch with the premaxillary bones. But in carriers, runts, and
barbs the singular reflexion of the upper margin of the middle part of the lower
jaw (see woodcut No. 25) is not strictly correlated with the width or
divergence (as may be clearly seen in woodcut No. 26) of the premaxillary
bones, but with the breadth of the horny and soft parts of the upper mandible,
which are always overlapped by the edges of the lower mandible.
In pouters, the elongation of the body is a selected character, and the ribs, as
we have seen, have generally become very broad, with the seventh pair
furnished with processes; the sacral and caudal vertebræ have been augmented
in number; the sternum has likewise increased in length (but not in the depth of
the crest) by .4 of an inch more than would follow from the greater bulk of the
body in comparison with that of the rock-pigeon. In fantails, the length and
number of the caudal vertebræ have increased. Hence, during the gradual
progress of variation and selection, the internal bony frame-work and the
external shape of the body have been, to a certain extent, modified in a
correlated manner.

Although the wings and tail often vary in length independently of each other, it
is scarcely possible to doubt that they generally tend to become elongated or
shortened in correlation. This is well seen in jacobins, and still more plainly in
runts, some varieties of which have their wings and tail of great length, whilst
others have both very short. With jacobins, the remarkable length of the tail
and wing-feathers is not a character which is intentionally selected by fanciers;
but fanciers have been trying for centuries, at least since the year 1600, to
increase the length of the reversed feathers on the neck, so that the hood may
more completely enclose the head; and it may be suspected that the increased
length of the wing and tail-feathers stands in correlation with the increased
length of the neck-feathers. Short-faced tumblers have short wings in nearly
due proportion with the reduced size of their bodies; but it is remarkable,
seeing that the number of the primary wing-feathers is a constant character in
most birds, that these tumblers generally have only nine instead of ten
primaries. I have myself observed this in eight birds; and the Original
Columbarian Society[311] reduced the standard for bald-head tumblers from ten
to nine white flight-feathers, thinking it unfair that a bird which had only nine
feathers should be disqualified for a prize because it had not ten white flight-
feathers. On the other hand, in carriers and runts, which have large bodies and
long wings, eleven primary feathers have occasionally been observed.
Mr. Tegetmeier has informed me of a curious and inexplicable case of
correlation, namely, that young pigeons of all breeds, which when mature
become white, yellow, silver (i.e. extremely pale blue), or dun-coloured, are
born almost naked; whereas other coloured pigeons are born well clothed with
down. Mr. Esquilant, however, has observed that young dun carriers are not so
bare as young dun barbs and tumblers. Mr. Tegetmeier has seen two young
birds in the same nest, produced from differently coloured parents, which
differed greatly in the degree to which they were at first clothed with down.
I have observed another case of correlation which at first sight appears quite
inexplicable, but on which, as we shall see in a future chapter, some light can
be thrown by the law of homologous parts varying in the same manner. The
case is, that, when the feet are much feathered, the roots of the feathers are
connected by a web of skin, and apparently in correlation with this the two
outer toes become connected for a considerable space by skin. I have observed
this in very many specimens of pouters, trumpeters, swallows, roller-tumblers
(likewise observed in this breed by Mr. Brent), and in a lesser degree in other
feather-footed pigeons.

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

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

I measured most of the birds which came into my possession, from
the feathered base of the beak (the length of beak itself being so
variable) to the end of the tail, and to the oil-gland, but unfortunately
(except in a few cases) not to the root of the tail; I measured each
bird from the extreme tip to tip of wing; and the length of the
terminal folded part of the wing, from the extremity of the primaries
to the joint of the radius. I measured the feet without the claws, from
the end of the middle toe to the end of the hind toe; and the tarsus
together with the middle toe. I have taken in every case the mean
measurement of two wild rock-pigeons from the Shetland Islands, as
the standard of comparison. The following table shows the actual
length of the feet in each bird; and the difference between the length
which the feet ought to have had according to the size of body of
each, in comparison with the size of body and length of feet of the
rock-pigeon, calculated (with a few specified exceptions) by the
standard of the length of the body from the base of the beak to the
oil-gland. I have preferred this standard, owing to the variability of
the length of tail. But I have made similar calculations, taking as the
standard the length from tip to tip of wing, and likewise in most
cases from the base of the beak to the end of the tail; and the result
has always been closely similar. To give an example: the first bird in

the table, being a short-faced tumbler, is much smaller than the rock-
pigeon, and would naturally have shorter feet; but it is found on
calculation to have feet too short by .11 of an inch, in comparison
with the feet of the rock-pigeon, relatively to the size of the body in
these two birds, as measured from the base of beak to the oil-gland.
So again, when this same tumbler and the rock-pigeon were
compared by the length of their wings, or by the extreme length of
their bodies, the feet of the tumbler were likewise found to be too
short in very nearly the same proportion. I am well aware that the
measurements pretend to greater accuracy than is possible, but it was
less trouble to write down the actual measurements given by the
compasses in each case than an approximation.

Table I.

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

Difference
between
actual and
calculated
length of
feet, in
proportion
to length of
feet and
Actual size of
length body in the
of Rock-
Name of Breed. Feet pigeon

Too Too
Wild rock-pigeon short long
(mean measurement) 2.02 by by

Short-faced Tumbler,
1.57 0.11 ..
bald-head

" "
1.60 0.16 ..
almond
Tumbler, red magpie 1.75 0.19 ..
" red common
(by standard to end of 1.85 0.07 ..
tail)
" common
1.85 0.18 ..
bald-head
" roller 1.80 0.06 ..
Turbit 1.75 0.17 ..
" 1.80 0.01 ..
" 1.84 0.15 ..
Jacobin 1.90 0.02 ..
Trumpeter, white 2.02 0.06 ..
" mottled 1.95 0.18 ..
Fantail (by standard to
1.85 0.15 ..
end of tail)
" "
1.95 0.15 ..
"
" crested var.
1.95 0.0 0.0
"
Indian Frill-back
1.80 0.19 ..
"
English Frill-back 2.10 0.03 ..
Nun 1.82 0.02 ..
Laugher 1.65 0.16 ..
Barb 2.00 0.03 ..
" 2.00 .. 0.03
Spot 1.90 0.02 ..
" 1.90 0.07 ..
Swallow, red 1.85 0.18 ..
" blue 2.00 .. 0.03
Pouter 2.42 .. 0.11
" German 2.30 .. 0.09
Bussorah Carrier 2.17 .. 0.09

Number of specimens 28 22 5

Table II.

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

Difference
between
actual and
calculated
length of
feet, in
proportion
to length of
feet and
Actual size of
length body in the
of Rock-
Name of Breed. Feet pigeon

Too Too
Wild rock-pigeon short long
(mean measurement) 2.02 by by

Carrier 2.60 .. 0.31
" 2.60 .. 0.25
" 2.40 .. 0.21
" Dragon 2.25 .. 0.06
Bagadotten Carrier 2.80 .. 0.56
Scanderoon, white 2.80 .. 0.37
" Pigeon
2.85 .. 0.29
cygne
Runt 2.75 .. 0.27

Number of specimens 8 .. 8

In these two tables we see in the first column the actual length of the
feet in thirty-six birds belonging to various breeds, and in the two
other columns we see by how much the feet are too short or too long,
according to the size of bird, in comparison with the rock-pigeon. In
the first table twenty-two specimens have their feet too short, on an
average by a little above the tenth of an inch (viz. .107); and five
specimens have their feet on an average a very little too long,
namely, by .07 of an inch. But some of these latter and exceptional
cases can be explained; for instance, with pouters the legs and feet
are selected for length, and thus any natural tendency to a diminution
in the length of the feet will have been counteracted. In the swallow
and barb, when the calculation was made on any standard of
comparison excepting the one above used (viz. length of body from
base of beak to oil-gland), the feet were found to be too small.
In the second table we have eight birds, with their beaks much
longer than in the rock-pigeon, both actually and proportionally with
the size of body, and their feet are in an equally marked manner
longer, namely, in proportion, on an average by .29 of an inch. I
should here state that in Table I. there are a few partial exceptions to
the beak being proportionally shorter than in the rock-pigeon: thus
the beak of the English frill-back is just perceptibly longer, and that
of the Bussorah carrier of the same length or slightly longer, than in
the rock-pigeon. The beaks of spots, swallows, and laughers are only
a very little shorter, or of the same proportional length, but slenderer.
Nevertheless, these two tables, taken conjointly, indicate pretty
plainly some kind of correlation between the length of the beak and
the size of the feet. Breeders of cattle and horses believe that there is
an analogous connection between the length of the limbs and head;
they assert that a race-horse with the head of a dray-horse, or a
greyhound with the head of a bulldog, would be a monstrous
production. As fancy pigeons are generally kept in small aviaries,
and are abundantly supplied with food, they must walk about much
less than the wild rock-pigeon; and it may be admitted as highly
probable that the reduction in the size of the feet in the twenty-two

birds in the first table has been caused by disuse,[312] and that this
reduction has acted by correlation on the beaks of the great majority
of the birds in Table I. When, on the other hand, the beak has been
much elongated by the continued selection of successive slight
increments of length, the feet by correlation have likewise become
much elongated in comparison with those of the wild rock-pigeon,
notwithstanding their lessened use.
As I had taken measures from the end of the middle toe to the heel of
the tarsus in the rock-pigeon and in the above thirty-six birds, I have
made calculations analogous with those above given, and the result
is the same,—namely, that in the short-beaked breeds, with equally
few exceptions as in the former case, the middle toe conjointly with
the tarsus has decreased in length; whereas in the long-beaked breeds
it has increased in length, though not quite so uniformly as in the
former case, for the leg in some varieties of the runt varies much in
length.
As fancy pigeons are generally confined in aviaries of moderate size,
and as even when not confined they do not search for their own food,
they must during many generations have used their wings
incomparably less than the wild rock-pigeon. Hence it seemed to me
probable that all the parts of the skeleton subservient to flight would
be found to be reduced in size. With respect to the sternum, I have
carefully measured its extreme length in twelve birds of different
breeds, and in two wild rock-pigeons from the Shetland Islands. For
the proportional comparison I have tried with all twelve birds three
standards of measurement, namely, the length from the base of the
beak to the oil-gland, to the end of the tail, and from the extreme tip
to tip of wings. The result has been in each case nearly the same, the
sternum being invariably found to be shorter than in the wild rock-
pigeon. I will give only a single table, as calculated by the standard
from the base of the beak to the oil-gland; for the result in this case
is nearly the mean between the results obtained by the two other
standards.

Length of Sternum.

Actual Too Actual Too
Name of Name of
Length. Short Length. Short
Breed. Breed.
Inches. by Inches. by

Wild Rock-
2.55 .. Barb 2.35 0.34
pigeon
Pied
2.80 0.60 Nun 2.27 0.15
Scanderoon
Bagadotten German
2.80 0.17 2.36 0.54
Carrier Pouter
Dragon 2.45 0.41 Jacobin 2.33 0.22
English
Carrier 2.75 0.35 Frill- 2.40 0.43
back
Short-faced
2.05 0.28 Swallow 2.45 0.17
Tumbler

This table shows that in these twelve breeds the sternum is on an
average one-third of an inch (exactly .332) shorter than in the rock-
pigeon, proportionally with the size of their bodies; so that the
sternum has been reduced by between one-seventh and one-eighth of
its entire length; and this is a considerable reduction.
I have also measured in twenty-one birds, including the above dozen,
the prominence of the crest of the sternum relatively to its length,
independently of the size of the body. In two of the twenty-one birds
the crest was prominent in the same relative degree as in the rock-
pigeon; in seven it was more prominent; but in five out of these
seven, namely, in a fantail, two scanderoons, and two English
carriers, this greater prominence may to a certain extent be
explained, as a prominent breast is admired and selected by fanciers;
in the remaining twelve birds the prominence was less. Hence it
follows that the crest exhibits a slight, though uncertain, tendency to
become reduced in prominence in a greater degree than does the
length of the sternum relatively to the size of body, in comparison
with the rock-pigeon.

I have measured the length of the scapula in nine different large and
small-sized breeds, and in all the scapula is proportionally shorter
(taking the same standard as before) than in the wild rock-pigeon.
The reduction in length on an average is very nearly one-fifth of an
inch, or about one-ninth of the length of the scapula in the rock-
pigeon.
The arms of the furcula in all the specimens which I compared,
diverged less, proportionally with the size of body, than in the rock-
pigeon; and the whole furcula was proportionally shorter. Thus in a
runt, which measured from tip to tip of wings 38½ inches, the
furcula was only a very little longer (with the arms hardly more
divergent) than in a rock-pigeon which measured from tip to tip 26½
inches. In a barb, which in all its measurements was a little larger
than the same rock-pigeon, the furcula was a quarter of an inch
shorter. In a pouter, the furcula had not been lengthened
proportionally with the increased length of the body. In a short-faced
tumbler, which measured from tip to tip of wings 24 inches,
therefore only 2½ inches less than the rock-pigeon, the furcula was
barely two-thirds of the length of that of the rock-pigeon.

We thus clearly see that the sternum, scapulæ, and furcula are all reduced in
proportional length; but when we turn to the wings we find what at first
appears a wholly different and unexpected result. I may here remark that I
have not picked out specimens, but have used every measurement made by me.
Taking the length from the base of beak to the end of the tail as the standard of
comparison, I find that, out of thirty-five birds of various breeds, twenty-five
have wings of greater, and ten have them of less proportional length, than in
the rock-pigeon. But from the frequently correlated length of the tail and wing-
feathers, it is better to take as the standard of comparison the length from the
base of the beak to the oil-gland; and by this standard, out of twenty-six of the
same birds which had been thus measured, twenty-one had wings too long, and
only five had them too short. In the twenty-one birds the wings exceeded in
length those of the rock-pigeon, on an average, by 1⅓ inch; whilst in the five
birds they were less in length by only .8 of an inch. As I was much surprised
that the wings of closely confined birds should thus so frequently have been
increased in length, it occurred to me that it might be solely due to the greater
length of the wing-feathers; for this certainly is the case with the jacobin,
which has wings of unusual length. As in almost every case I had measured the

folded wings, I subtracted the length of this terminal part from that of the
expanded wings, and thus I obtained, with a moderate degree of accuracy, the
length of the wings from the ends of the two radii, answering from wrist to
wrist in our arms. The wings, thus measured in the same twenty-five birds,
now gave a widely different result; for they were proportionally with those of
the rock-pigeon too short in seventeen birds, and in only eight too long. Of
these eight birds, five were long-beaked,[313] and this fact perhaps indicates
that there is some correlation between the length of the beak and the length of
the bones of the wings, in the same manner as with the feet and tarsi. The
shortening of the humerus and radius in the seventeen birds may probably be
attributed to disuse, as in the case of the scapulæ and furcula to which the
wing-bones are attached;—the lengthening of the wing-feathers, and
consequently the expansion of the wings from tip to tip, being, on the other
hand, as completely independent of use and disuse as is the growth of the hair
or wool on our long-haired dogs or long-woolled sheep.
To sum up: we may confidently admit that the length of the sternum, and
frequently the prominence of its crest, the length of the scapulæ and furcula,
have all been reduced in size in comparison with the same parts in the rock-
pigeon. And I presume that this may be safely attributed to disuse or lessened
exercise. The wings, as measured from the ends of the radii, have likewise
been generally reduced in length; but, owing to the increased growth of the
wing-feathers, the wings, from tip to tip, are commonly longer than in the
rock-pigeon. The feet, as well as the tarsi conjointly with the middle toe, have
likewise in most cases become reduced; and this it is probable has been caused
by their lessened use; but the existence of some sort of correlation between the
feet and beak is shown more plainly than the effects of disuse. We have also
some faint indication of a similar correlation between the main bones of the
wing and the beak.
Summary on the Points of Difference between the several Domestic Races, and
between the individual Birds.—The beak, together with the bones of the face,
differ remarkably in length, breadth, shape, and curvature. The skull differs in
shape, and greatly in the angle formed by the union of the premaxillary, nasal,
and maxillo-jugal bones. The curvature of the lower jaw and the reflexion of
its upper margin, as well as the gape of the mouth, differ in a highly
remarkable manner. The tongue varies much in length, both independently and
in correlation with the length of the beak. The development of the naked,
wattled skin over the nostrils and round the eyes varies in an extreme degree.

The eyelids and the external orifices of the nostrils vary in length, and are to a
certain extent correlated with the degree of development of the wattle. The size
and form of the œsophagus and crop, and their capacity for inflation, differ
immensely. The length of the neck varies. With the varying shape of the body,
the breadth and number of the ribs, the presence of processes, the number of
the sacral vertebræ, and the length of the sternum, all vary. The number and
size of the coccygeal vertebræ vary, apparently in correlation with the
increased size of the tail. The size and shape of the perforations in the sternum,
and the size and divergence of the arms of the furcula, differ. The oil-gland
varies in development, and is sometimes quite aborted. The direction and
length of certain feathers have been much modified, as in the hood of the
Jacobin and the frill of the Turbit. The wing and tail feathers generally vary in
length together, but sometimes independently of each other and of the size of
the body. The number and position of the tail-feathers vary to an unparalleled
degree. The primary and secondary wing-feathers occasionally vary in number,
apparently in correlation with the length of the wing. The length of the leg and
the size of the feet, and, in connection with the latter, the number of the
scutellæ, all vary. A web of skin sometimes connects the bases of the two inner
toes, and almost invariably the two outer toes when the feet are feathered.
The size of the body differs greatly: a runt has been known to weigh more than
five times as much as a short-faced tumbler. The eggs differ in size and shape.
According to Parmentier,[314] some races use much straw in building their
nests, and others use little; but I cannot hear of any recent corroboration of this
statement. The length of time required for hatching the eggs is uniform in all
the breeds. The period at which the characteristic plumage of some breeds is
acquired, and at which certain changes of colour supervene, differs. The degree
to which the young birds are clothed with down when first hatched is different,
and is correlated in a singular manner with the future colour of the plumage.
The manner of flight, and certain inherited movements, such as clapping the
wings, tumbling either in the air or on the ground, and the manner of courting
the female, present the most singular differences. In disposition the several
races differ. Some races are very silent; others coo in a highly peculiar manner.
Although many different races have kept true in character during several
centuries, as we shall hereafter more fully see, yet there is far more individual
variability in the truest breeds than in birds in a state of nature. There is hardly
any exception to the rule that those characters vary most which are now most
valued and attended to by fanciers, and which consequently are now being

improved by continued selection. This is indirectly admitted by fanciers when
they complain that it is much more difficult to breed high fancy pigeons up to
the proper standard of excellence than the so-called toy pigeons, which differ
from each other merely in colour; for particular colours when once acquired
are not liable to continued improvement or augmentation. Some characters
become attached, from quite unknown causes, more strongly to the male than
to the female sex; so that we have, in certain races, a tendency towards the
appearance of secondary sexual characters,[315] of which the aboriginal rock-
pigeon displays not a trace.

CHAPTER VI.

PIGEONS—continued.

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

The differences described in the last chapter between the eleven chief domestic
races and between individual birds of the same race, would be of little
significance, if they had not all descended from a single wild stock. The
question of their origin is therefore of fundamental importance, and must be
discussed at considerable length. No one will think this superfluous who
considers the great amount of difference between the races, who knows how
ancient many of them are, and how truly they breed at the present day.
Fanciers almost unanimously believe that the different races are descended
from several wild stocks, whereas most naturalists believe that all are
descended from the Columba livia or rock-pigeon.

Temminck[316] has well observed, and Mr. Gould has made the same remark to
me, that the aboriginal parent must have been a species which roosted and built
its nest on rocks; and I may add that it must have been a social bird. For all the
domestic races are highly social, and none are known to build or habitually to
roost on trees. The awkward manner in which some pigeons, kept by me in a
summer-house near an old walnut-tree, occasionally alighted on the barer
branches, was evident.[317] Nevertheless, Mr. R. Scot Skirving informs me that
he often saw crowds of pigeons in Upper Egypt settling on the low trees, but
not on the palms, in preference to the mud hovels of the natives. In India Mr.
Blyth[318] has been assured that the wild C. livia, var. intermedia, sometimes
roosts in trees. I may here give a curious instance of compulsion leading to
changed habits: the banks of the Nile above lat. 28° 30' are perpendicular for a
long distance, so that when the river is full the pigeons cannot alight on the
shore to drink, and Mr. Skirving repeatedly saw whole flocks settle on the
water, and drink whilst they floated down the stream. These flocks seen from a
distance resembled flocks of gulls on the surface of the sea.
If any domestic race had descended from a species which was not social, or
which built its nest or roosted in trees,[319] the sharp eyes of fanciers would
assuredly have detected some vestige of so different an aboriginal habit. For
we have reason to believe that aboriginal habits are long retained under
domestication. Thus with the common ass we see signs of its original desert
life in its strong dislike to cross the smallest stream of water, and in its pleasure
in rolling in the dust. The same strong dislike to cross a stream is common to
the camel, which has been domesticated from a very ancient period. Young
pigs, though so tame, sometimes squat when frightened, and thus try to
conceal themselves even on an open and bare place. Young turkeys, and
occasionally even young fowls, when the hen gives the danger-cry, run away
and try to hide themselves, like young partridges or pheasants, in order that
their mother may take flight, of which she has lost the power. The musk-duck
(Dendrocygna viduata) in its native country often perches and roosts on trees,
[320]
and our domesticated musk-ducks, though such sluggish birds, "are fond
of perching on the tops of barns, walls, &c., and, if allowed to spend the night
in the hen-house, the female will generally go to roost by the side of the hens,
but the drake is too heavy to mount thither with ease."[321] We know that the
dog, however well and regularly fed, often buries, like the fox, any superfluous
food; and we see him turning round and round on a carpet, as if to trample
down grass to form a bed; we see him on bare pavements scratching

backwards as if to throw earth over his excrement, although, as I believe, this
is never effected even where there is earth. In the delight with which lambs and
kids crowd together and frisk on the smallest hillock, we see a vestige of their
former alpine habits.
We have therefore good reason to believe that all the domestic races of the
pigeon are descended either from some one or from several species which both
roosted and built their nests on rocks, and were social in disposition. As only
five or six wild species with these habits and making any near approach in
structure to the domesticated pigeon are known to exist, I will enumerate them.

Firstly, the Columba leuconota resembles certain domestic varieties
in its plumage, with the one marked and never-failing difference of a
white band which crosses the tail at some distance from the
extremity. This species, moreover, inhabits the Himalaya, close to
the limit of perpetual snow; and therefore, as Mr. Blyth has
remarked, is not likely to have been the parent of our domestic
breeds, which thrive in the hottest countries. Secondly, the C.
rupestris, of Central Asia, which is intermediate[322] between the C.
leuconota and livia; but has nearly the same coloured tail with the
former species. Thirdly, the Columba littoralis builds and roosts,
according to Temminck, on rocks in the Malayan archipelago; it is
white, excepting parts of the wing and the tip of the tail, which are
black; its legs are livid-coloured, and this is a character not observed
in any adult domestic pigeon; but I need not have mentioned this
species or the closely-allied C. luctuosa, as they in fact belong to the
genus Carpophaga. Fourthly, Columba Guinea, which ranges from
Guinea[323] to the Cape of Good Hope, and roosts either on trees or
rocks, according to the nature of the country. This species belongs to
the genus Strictœnas of Reichenbach, but is closely allied to true
Columba; it is to some extent coloured like certain domestic races,
and has been said to be domesticated in Abyssinia; but Mr.
Mansfield Parkyns, who collected the birds of that country and
knows the species, informs me that this is a mistake. Moreover the
C. Guinea is characterized by the feathers of the neck having
peculiar notched tips,—a character not observed in any domestic
race. Fifthly, the Columba œnas of Europe, which roosts on trees,
and builds its nest in holes, either in trees or the ground; this species,
as far as external characters go, might be the parent of several

domestic races; but, though it crosses readily with the true rock-
pigeon, the offspring, as we shall presently see, are sterile hybrids,
and of such sterility there is not a trace when the domestic races are
intercrossed. It should also be observed that if we were to admit,
against all probability, that any of the foregoing five or six species
were the parents of some of our domestic pigeons, not the least light
would be thrown on the chief differences between the eleven most
strongly-marked races.
We now come to the best known rock-pigeon, the Columba livia,
which is often designated in Europe pre-eminently as the Rock-
pigeon, and which naturalists believe to be the parent of all the
domesticated breeds. This bird agrees in every essential character
with the breeds which have been only slightly modified. It differs
from all other species in being of a slaty-blue colour, with two black
bars on the wings, and with the croup (or loins) white. Occasionally
birds are seen in Faroe and the Hebrides with the black bars replaced
by two or three black spots; this form has been named by Brehm[324]
C. amaliæ, but this species has not been admitted as distinct by other
ornithologists. Graba[325] even found a difference between the wing-
bars of the same bird in Faroe. Another and rather more distinct form
is either truly wild or has become feral on the cliffs of England, and
was doubtfully named by Mr. Blyth[326] as C. affinis, but is now no
longer considered by him as a distinct species. C. affinis is rather
smaller than the rock-pigeon of the Scottish islands, and has a very
different appearance owing to the wing-coverts being chequered
with black, with similar marks often extending over the back. The
chequering consists of a large black spot on the two sides, but chiefly
on the outer side, of each feather. The wing-bars in the true rock-
pigeon and in the chequered variety are, in fact, due to similar
though larger spots symmetrically crossing the secondary wing-
feather and the larger coverts. Hence the chequering arises merely
from an extension of these marks to other parts of the plumage.
Chequered birds are not confined to the coasts of England; for they
were found by Graba at Faroe; and W. Thompson[327] says that at
Islay fully half the wild rock-pigeons were chequered. Colonel King,
of Hythe, stocked his dovecot with young wild birds which he
himself procured from nests at the Orkney Islands; and several

specimens, kindly sent to me by him, were all plainly chequered. As
we thus see that chequered birds occur mingled with the true rock-
pigeon at three distinct sites, namely, Faroe, the Orkney Islands, and
Islay, no importance can be attached to this natural variation in the
plumage.

Prince C. L. Bonaparte,[328] a great divider of species, enumerates,
with a mark of interrogation, as distinct from C. livia, the C.
turricola of Italy, the C. rupestris of Daouria, and the C. Schimperi
of Abyssinia; but these birds differ from C. livia in characters of the
most trifling value. In the British Museum there is a chequered
pigeon, probably the C. Schimperi of Bonaparte, from Abyssinia. To
these may be added the C. gymnocyclus of G. R. Gray from W.
Africa, which is slightly more distinct, and has rather more naked
skin round the eyes than the rock-pigeon; but from information given
me by Dr. Daniell, it is doubtful whether this is a wild bird, for
dovecot-pigeons (which I have examined) are kept on the coast of
Guinea.
The wild rock-pigeon of India (C. intermedia of Strickland) has been
more generally accepted as a distinct species. It chiefly differs in the
croup being blue instead of snow-white; but as Mr. Blyth informs
me, the tint varies, being sometimes albescent. When this form is
domesticated chequered birds appear, just as occurs in Europe with
the truly wild C. livia. Moreover we shall immediately have proof
that the blue and white croup is a highly variable character; and
Bechstein[329] asserts that with dovecot-pigeons in Germany this is
the most variable of all the characters of the plumage. Hence it may
be concluded that C. intermedia cannot be ranked as specifically
distinct from C. livia.
In Madeira there is a rock-pigeon which a few ornithologists have
suspected to be distinct from C. livia. I have examined numerous
specimens collected by Mr. E. V. Harcourt and Mr. Mason. They are
rather smaller than the rock-pigeon from the Shetland Islands, and
their beaks are plainly thinner; but the thickness of the beak varied in
the several specimens. In plumage there is remarkable diversity;
some specimens are identical in every feather (I speak after actual
comparison) with the rock-pigeon of the Shetland Islands; others are

chequered, like C. affinis from the cliffs of England, but generally to
a greater degree, being almost black over the whole back; others are
identical with the so-called C. intermedia of India in the degree of
blueness of the croup; whilst others have this part very pale or very
dark blue, and are likewise chequered. So much variability raises a
strong suspicion that these birds are domestic pigeons which have
become feral.
From these facts it can hardly be doubted that C. livia, affinis,
intermedia, and the forms marked with an interrogation by
Bonaparte, ought all to be included under a single species. But it is
quite immaterial whether or not they are thus ranked, and whether
some one of these forms or all are the progenitors of the various
domestic kinds, as far as any light is thus thrown on the differences
between the more strongly-marked races. That common dovecot-
pigeons, which are kept in various parts of the world, are descended
from one or from several of the above-mentioned wild varieties of C.
livia, no one who compares them will doubt. But before making a
few remarks on dovecot-pigeons, it should be stated that the wild
rock-pigeon has been found easy to tame in several countries. We
have seen that Colonel King at Hythe stocked his dovecot more than
twenty years ago with young wild birds taken at the Orkney Islands,
and since this time they have greatly multiplied. The accurate
Macgillivray[330] asserts that he completely tamed a wild rock-
pigeon in the Hebrides; and several accounts are on record of these
pigeons having bred in dovecots in the Shetland Islands. In India, as
Captain Hutton informs me, the wild rock-pigeon is easily tamed,
and breeds readily with the domestic kind; and Mr. Blyth[331] asserts
that wild birds come frequently to the dovecots and mingle freely
with their inhabitants. In the ancient 'Ayeen Akbery' it is written that,
if a few wild pigeons be taken, "they are speedily joined by a
thousand others of their kind."
Dovecot-pigeons are those which are kept in dovecots in a semi-
domesticated state; for no special care is taken of them, and they
procure their own food, except during the severest weather. In
England, and, judging from MM. Boitard and Corbié's work, in
France, the common dovecot-pigeon exactly resembles the
chequered variety of C. livia; but I have seen dovecots brought from

Yorkshire, without any trace of chequering, like the wild rock-pigeon
of the Shetland Islands. The chequered dovecots from the Orkney
Islands, after having been domesticated by Colonel King for more
than twenty years, differed slightly from each other in the darkness
of their plumage, and in the thickness of their beaks; the thinnest
beak being rather thicker than the thickest one in the Madeira birds.
In Germany, according to Bechstein, the common dovecot-pigeon is
not chequered. In India they often become chequered, and
sometimes pied with white; the croup also, as I am informed by Mr.
Blyth, becomes nearly white. I have received from Sir J. Brooke
some dovecot-pigeons, which originally came from the S. Natunas
Islands in the Malay archipelago, and which had been crossed with
the Singapore dovecots; they were small, and the darkest variety was
extremely like the dark chequered variety with a blue croup from
Madeira; but the beak was not so thin, though decidedly thinner than
in the rock-pigeon from the Shetland Islands. A dovecot-pigeon sent
to me by Mr. Swinhoe from Foochow, in China, was likewise rather
small, but differed in no other respect. I have also received, through
the kindness of Dr. Daniell, four living dovecot-pigeons from Sierra
Leone;[332] these were fully as large as the Shetland rock-pigeon,
with even bulkier bodies. In plumage some of them were identical
with the Shetland rock-pigeon, but with the metallic tints apparently
rather more brilliant; others had a blue croup and resembled the
chequered variety of C. intermedia of India; and some were so much
chequered as to be nearly black. In these four birds the beak differed
slightly in length, but in all it was decidedly shorter, more massive,
and stronger than in the wild rock-pigeon from the Shetland Islands,
or in the English dovecot. When the beaks of these African pigeons
were compared with the thinnest beaks of the wild Madeira
specimens, the contrast was great; the former being fully one-third
thicker in a vertical direction than the latter; so that any one at first
would have felt inclined to rank these birds as specifically distinct;
yet-so perfectly graduated a series could be formed between the
above-mentioned varieties, that it was obviously impossible to
separate them.

To sum up: the wild Columba livia, including under this name C. affinis,
intermedia, and the other still more closely-affined geographical races, has a
vast range from the southern coast of Norway and the Faroe Islands to the

shores of the Mediterranean, to Madeira and the Canary Islands, to Abyssinia,
India, and Japan. It varies greatly in plumage, being in many places chequered
with black, and having either a white or blue croup or loins: it varies also
slightly in the size of the beak and body. Dovecot-pigeons, which no one
disputes are descended from one or more of the above wild forms, present a
similar but greater range of variation in plumage, in the size of body, and in the
length and thickness of the beak. There seems to be some relation between the
croup being blue or white, and the temperature of the country inhabited by
both wild and dovecot pigeons; for nearly all the dovecot-pigeons in the
northern parts of Europe have a white croup, like that of the wild European
rock-pigeon; and nearly all the dovecot-pigeons of India have a blue croup like
that of the wild C. intermedia of India. As in various countries the wild rock-
pigeon has been found easy to tame, it seems extremely probable that the
dovecot-pigeons throughout the world are the descendants of at least two and
perhaps more wild stocks, but these, as we have just seen, cannot be ranked as
specifically distinct.
With respect to the variation of C. livia, we may without fear of contradiction
go one step further. Those pigeon-fanciers who believe that all the chief races,
such as Carriers, Pouters, Fantails, &c., are descended from distinct aboriginal
stocks, yet admit that the so-called toy-pigeons, which differ from the rock-
pigeon in little except in colour, are descended from this bird. By toy-pigeons
are meant such birds as Spots, Nuns, Helmets, Swallows, Priests, Monks,
Porcelains, Swabians, Archangels, Breasts, Shields, and others in Europe, and
many others in India. It would indeed be as puerile to suppose that all these
birds are descended from so many distinct wild stocks as to suppose this to be
the case with the many varieties of the gooseberry, heartsease, or dahlia. Yet
these pigeons all breed true, and many of them present sub-varieties which
likewise truly transmit their character. They differ greatly from each other and
from the rock-pigeon in plumage, slightly in size and proportions of body, in
size of feet, and in the length and thickness of their beaks. They differ from
each other in these respects more than do dovecot-pigeons. Although we may
safely admit that the latter, which vary slightly, and that the toy-pigeons, which
vary in a greater degree in accordance with their more highly-domesticated
condition, are descended from C. livia, including under this name the above-
enumerated wild geographical races; yet the question becomes far more
difficult when we consider the eleven principal races, most of which have been
so profoundly modified. It can, however, be shown, by indirect evidence of a
perfectly conclusive nature, that these principal races are not descended from

so many wild stocks; and if this be once admitted, few will dispute that they
are the descendants of C. livia, which agrees with them so closely in habits and
in most characters, which varies in a state of nature, and which has certainly
undergone a considerable amount of variation, as in the toy-pigeons. We shall
moreover presently see how eminently favourable circumstances have been for
a great amount of modification in the more carefully tended breeds.
The reasons for concluding that the several principal races have not descended
from so many aboriginal and unknown stocks may be grouped under the
following six heads:—Firstly, if the eleven chief races have not arisen from
the variation of some one species, together with its geographical races, they
must be descended from several extremely distinct aboriginal species; for no
amount of crossing between only six or seven wild forms could produce races
so distinct as pouters, carriers, runts, fantails, turbits, short-faced tumblers,
jacobins, and trumpeters. How could crossing produce, for instance, a pouter
or a fantail, unless the two supposed aboriginal parents possessed the
remarkable characters of these breeds? I am aware that some naturalists,
following Pallas, believe that crossing gives a strong tendency to variation,
independently of the characters inherited from either parent. They believe that
it would be easier to raise a pouter or fantail pigeon from crossing two distinct
species, neither of which possessed the characters of these races, than from any
single species. I can find few facts in support of this doctrine, and believe in it
only to a limited degree; but in a future chapter I shall have to recur to this
subject. For our present purpose the point is not material. The question which
concerns us is, whether or not many new and important characters have arisen
since man first domesticated the pigeon. On the ordinary view, variability is
due to changed conditions of life; on the Pallasian doctrine, variability, or the
appearance of new characters, is due to some mysterious effect from the
crossing of two species, neither of which possess the characters in question. In
some few instances it is credible, though for several reasons not probable, that
well-marked races have been formed by crossing; for instance, a barb might
perhaps have been formed by a cross between a long-beaked carrier, having
large eye-wattles, and some short-beaked pigeon. That many races have been
in some degree modified by crossing, and that certain varieties which are
distinguished only by peculiar tints have arisen from crosses between
differently-coloured varieties, may be admitted as almost certain. On the
doctrine, therefore, that the chief races owe their differences to their descent
from distinct species, we must admit that at least eight or nine, or more
probably a dozen species, all having the same habit of breeding and roosting

on rocks and living in society, either now exist somewhere, or formerly existed
but have become extinct as wild birds. Considering how carefully wild pigeons
have been collected throughout the world, and what conspicuous birds they
are, especially when frequenting rocks, it is extremely improbable that eight or
nine species, which were long ago domesticated and therefore must have
inhabited some anciently known country, should still exist in the wild state and
be unknown to ornithologists.
The hypothesis that such species formerly existed, but have become extinct, is
in some slight degree more probable. But the extinction of so many species
within the historical period is a bold hypothesis, seeing how little influence
man has had in exterminating the common rock-pigeon, which agrees in all its
habits of life with the domestic races. The C. livia now exists and flourishes on
the small northern islands of Faroe, on many islands off the coast of Scotland,
on Sardinia and the shores of the Mediterranean, and in the centre of India.
Fanciers have sometimes imagined that the several supposed parent-species
were originally confined to small islands, and thus might readily have been
exterminated; but the facts just given do not favour the probability of their
extinction, even on small islands. Nor is it probable, from what is known of the
distribution of birds, that the islands near Europe should have been inhabited
by peculiar species of pigeons; and if we assume that distant oceanic islands
were the homes of the supposed parent-species, we must remember that
ancient voyages were tediously slow, and that ships were then ill-provided
with fresh food, so that it would not have been easy to bring home living birds.
I have said ancient voyages, for nearly all the races of the pigeon were known
before the year 1600, so that the supposed wild species must have been
captured and domesticated before that date.
Secondly.—The doctrine that the chief domestic races have descended from
several aboriginal species, implies that several species were formerly so
thoroughly domesticated as to breed readily when confined. Although it is easy
to tame most wild birds, experience shows us that it is difficult to get them to
breed freely under confinement; although it must be owned that this is less
difficult with pigeons than with most other birds. During the last two or three
hundred years, many birds have been kept in aviaries, but hardly one has been
added to our list of thoroughly reclaimed species; yet on the above doctrine we
must admit that in ancient times nearly a dozen kinds of pigeons, now
unknown in the wild state, were thoroughly domesticated.

Thirdly.—Most of our domesticated animals have run wild in various parts of
the world; but birds, owing apparently to their partial loss of the power of
flight, less often than quadrupeds. Nevertheless I have met with accounts
showing that the common fowl has become feral in South America and
perhaps in West Africa, and on several islands: the turkey was at one time
almost feral on the banks of the Parana; and the Guinea-fowl has become
perfectly wild at Ascension and in Jamaica. In this latter island the peacock,
also, "has become a maroon bird." The common duck wanders from its home
and becomes almost wild in Norfolk. Hybrids between the common and musk-
duck which have become wild have been shot in North America, Belgium, and
near the Caspian Sea. The goose is said to have run wild in La Plata. The
common dovecot-pigeon has become wild at Juan Fernandez, Norfolk Island,
Ascension, probably at Madeira, on the shores of Scotland, and, as is asserted,
on the banks of the Hudson in North America.[333] But how different is the
case, when we turn to the eleven chief domestic races of the pigeon, which are
supposed by some authors to be descended from so many distinct species! no
one has ever pretended that any one of these races has been found wild in any
quarter of the world; yet they have been transported to all countries, and some
of them must have been carried back to their native homes. On the view that
all the races are the product of variation, we can understand why they have not
become feral, for the great amount of modification which they have undergone
shows how long and how thoroughly they have been domesticated; and this
would unfit them for a wild life.
Fourthly.—If it be assumed that the characteristic differences between the
various domestic races are due to descent from several aboriginal species, we
must conclude that man chose for domestication in ancient times, either
intentionally or by chance, a most abnormal set of pigeons; for that species
resembling such birds as pouters, fantails, carriers, barbs, short-faced tumblers,
turbits, &c., would be in the highest degree abnormal, as compared with all the
existing members of the great pigeon-family, cannot be doubted. Thus we
should have to believe that man not only formerly succeeded in thoroughly
domesticating several highly abnormal species, but that these same species
have since all become extinct, or are at least now unknown. This double
accident is so extremely improbable that the assumed existence of so many
abnormal species would require to be supported by the strongest evidence. On
the other hand, if all the races are descended from C. livia, we can understand,
as will hereafter be more fully explained, how any slight deviation in structure
which first appeared would continually be augmented by the preservation of

the most strongly marked individuals; and as the power of selection would be
applied according to man's fancy, and not for the bird's own good, the
accumulated amount of deviation would certainly be of an abnormal nature in
comparison with the structure of pigeons living in a state of nature.
I have already alluded to the remarkable fact, that the characteristic differences
between the chief domestic races are eminently variable: we see this plainly in
the great difference in the number of the tail-feathers in the fantail, in the
development of the crop in pouters, in the length of the beak in tumblers, in the
state of the wattle in carriers, &c. If these characters are the result of
successive variations added together by selection, we can understand why they
should be so variable: for these are the very parts which have varied since the
domestication of the pigeon, and therefore would be likely still to vary; these
variations moreover have been recently, and are still being accumulated by
man's selection; therefore they have not as yet become firmly fixed.
Fifthly.—All the domestic races pair readily together, and, what is equally
important, their mongrel offspring are perfectly fertile. To ascertain this fact I
made many experiments, which are given in the note below; and recently Mr.
Tegetmeier has made similar experiments with the same result.[334] The
accurate Neumeister[335] asserts that when dovecots are crossed with pigeons
of any other breed, the mongrels are extremely fertile and hardy. MM. Boitard
and Corbié[336] affirm, after their great experience, that with crossed pigeons
the more distinct the breeds, the more productive are their mongrel offspring. I
admit that the doctrine first broached by Pallas is highly probable, if not
actually proved, namely, that closely allied species, which in a state of nature
or when first captured would have been in some degree sterile when crossed,
lose this sterility after a long course of domestication; yet when we consider
the great difference between such races as pouters, carriers, runts, fantails,
turbits, tumblers, &c., the fact of their perfect, or even increased, fertility when
intercrossed in the most complicated manner becomes a strong argument in
favour of their having all descended from a single species. This argument is
rendered much stronger when we hear (I append in a note[337] all the cases
which I have collected) that hardly a single well-ascertained instance is known
of hybrids between two true species of pigeons being fertile, inter se, or even
when crossed with one of their pure parents.
Sixthly.—Excluding certain important characteristic differences, the chief races
agree most closely both with each other and with C. livia in all other respects.

As previously observed, all are eminently sociable; all dislike to perch or roost,
and refuse to build in trees; all lay two eggs, and this is not a universal rule
with the Columbidæ; all, as far as I can hear, require the same time for
hatching their eggs; all can endure the same great range of climate; all prefer
the same food, and are passionately fond of salt; all exhibit (with the asserted
exception of the finnikin and turner, which do not differ much in any other
character) the same peculiar gestures when courting the females; and all (with
the exception of trumpeters and laughers, which likewise do not differ much in
any other character) coo in the same peculiar manner, unlike the voice of any
other wild pigeon. All the coloured breeds display the same peculiar metallic
tints on the breast, a character far from general with pigeons. Each race
presents nearly the same range of variation in colour; and in most of the races
we have the same singular correlation between the development of down in the
young and the future colour of plumage. All have the proportional length of
their toes, and of their primary wing-feathers, nearly the same,—characters
which are apt to differ in the several members of the Columbidæ. In those
races which present some remarkable deviation of structure, such as in the tail
of fantails, crop of pouters, beak of carriers and tumblers, &c., the other parts
remain nearly unaltered. Now every naturalist will admit that it would be
scarcely possible to pick out a dozen natural species in any Family, which
should agree closely in habits and in general structure, and yet should differ
greatly in a few characters alone. This fact is explicable through the doctrine of
natural selection; for each successive modification of structure in each natural
species is preserved, solely because it is of service; and such modifications
when largely accumulated imply a great change in the habits of life, and this
will almost certainly lead to other changes of structure throughout the whole
organisation. On the other hand, if the several races of the pigeon have been
produced by man through selection and variation, we can readily understand
how it is that they should still all resemble each other in habits and in those
many characters which man has not cared to modify, whilst they differ to so
prodigious a degree in those parts which have struck his eye or pleased his
fancy.
Besides the points above enumerated, in which all the domestic races resemble
C. livia and each other, there is one which deserves special notice. The wild
rock-pigeon is of a slaty-blue colour; the wings are crossed by two black bars;
the croup varies in colour, being generally white in the pigeon of Europe, and
blue in that of India; the tail has a black bar close to the end, and the outer
webs of the outer tail-feathers are edged with white, except near the tips. These

combined characters are not found in any wild pigeon besides C. livia. I have
looked carefully through the great collection of pigeons in the British Museum,
and I find that a dark bar at the end of the tail is common; that the white edging
to the outer tail-feathers is not rare; but that the white croup is extremely rare,
and the two black bars on the wings occur in no other pigeon, excepting the
alpine C. leuconota and C. rupestris of Asia. Now if we turn to the domestic
races, it is highly remarkable, as an eminent fancier, Mr. Wicking, observed to
me, that, whenever a blue bird appears in any race, the wings almost invariably
show the double black bars.[338] The primary wing-feathers may be white or
black, and the whole body may be of any colour, but if the wing-coverts alone
are blue, the two black bars surely appear. I have myself seen, or acquired
trustworthy evidence, as given below,[339] of blue birds with black bars on the
wing, with the croup either white or very pale or dark blue, with the tail having
a terminal black bar, and with the outer feathers externally edged with white or
very pale coloured, in the following races, which, as I carefully observed in
each case, appeared to be perfectly pure: namely, in Pouters, Fantails,
Tumblers, Jacobins, Turbits, Barbs, Carriers, Runts of three distinct varieties,
Trumpeters, Swallows, and in many other toy-pigeons, which, as being closely
allied to C. livia, are not worth enumerating. Thus we see that, in purely-bred
races of every kind known in Europe, blue birds occasionally appear having all
the marks which characterise C. livia, and which concur in no other wild
species. Mr. Blyth, also, has made the same observation with respect to the
various domestic races known in India.
Certain variations in the plumage are equally common in the wild C. livia, in
dovecot-pigeons, and in all the most highly modified races. Thus, in all, the
croup varies from white to blue, being most frequently white in Europe, and
very generally blue in India.[340] We have seen that the wild C. livia in Europe,
and dovecots in all parts of the world, often have the upper wing-coverts
chequered with black; and all the most distinct races, when blue, are
occasionally chequered in precisely the same manner. Thus I have seen
Pouters, Fantails, Carriers, Turbits, Tumblers (Indian and English), Swallows,
Bald-pates, and other toy-pigeons blue and chequered; and Mr. Esquilant has
seen a chequered Runt. I bred from two pure blue Tumblers a chequered bird.

The facts hitherto given refer to the occasional appearance in pure races of
blue birds with black wing-bars, and likewise of blue and chequered birds; but
it will now be seen that when two birds belonging to distinct races are crossed,

neither of which have, nor probably have had during many generations, a trace
of blue in their plumage, or a trace of wing-bars and the other characteristic
marks, they very frequently produce mongrel offspring of a blue colour,
sometimes chequered, with black wing-bars, &c.; or if not of a blue colour, yet
with the several characteristic marks more or less plainly developed. I was led
to investigate this subject from MM. Boitard and Corbié[341] having asserted
that from crosses between certain breeds it is rare to get anything but bisets or
dovecot-pigeons, which, as we know, are blue birds with the usual
characteristic marks. We shall hereafter see that this subject possesses,
independently of our present object, considerable interest, so that I will give
the results of my own trials in full. I selected for experiment races which, when
pure, very seldom produce birds of a blue colour, or have bars on their wings
and tail.
The nun is white, with the head, tail, and primary wing-feathers black; it is a
breed which was established as long ago as the year 1600. I crossed a male nun
with a female red common tumbler, which latter variety generally breeds true.
Thus neither parent had a trace of blue in the plumage, or of bars on the wing
and tail. I should premise that common tumblers are rarely blue in England.
From the above cross I reared several young: one was red over the whole back,
but with the tail as blue as that of the rock-pigeon; the terminal bar, however,
was absent, but the outer feathers were edged with white: a second and third
nearly resembled the first, but the tail in both presented a trace of the bar at the
end: a fourth was brownish, and the wings showed a trace of the double bar: a
fifth was pale blue over the whole breast, back, croup, and tail, but the neck
and primary wing-feathers were reddish; the wings presented two distinct bars
of a red colour; the tail was not barred, but the outer feathers were edged with
white. I crossed this last curiously coloured bird with a black mongrel of
complicated descent, namely, from a black barb, a spot, and almond tumbler,
so that the two young birds produced from this cross included the blood of five
varieties, none of which had a trace of blue or of wing and tail bars: one of the
two young birds was brownish-black, with black wing-bars; the other was
reddish-dun, with reddish wing-bars, paler than the rest of the body, with the
croup pale blue, the tail bluish, with a trace of the terminal bar.

Mr. Eaton[342] matched two short-faced tumblers, namely, a splash cock and
kite hen (neither of which are blue or barred), and from the first nest he got a
perfect blue bird, and from the second a silver or pale blue bird, both of which,

in accordance with all analogy, no doubt presented the usual characteristic
marks.
I crossed two male black barbs with two female red spots. These latter have the
whole body and wings white, with a spot on the forehead, the tail and tail-
coverts red; the race existed at least as long ago as 1676, and now breeds
perfectly true, as was known to be the case in the year 1735.[343] Barbs are
uniformly-coloured birds, with rarely even a trace of bars on the wing or tail;
they are known to breed very true. The mongrels thus raised were black or
nearly black, or dark or pale brown, sometimes slightly piebald with white: of
these birds no less than six presented double wing-bars; in two the bars were
conspicuous and quite black; in seven some white feathers appeared on the
croup; and in two or three there was a trace of the terminal bar to the tail, but
in none were the outer tail-feathers edged with white.
I crossed black barbs (of two excellent strains) with purely-bred, snow-white
fantails. The mongrels were generally quite black, with a few of the primary
wing and tail-feathers white: others were dark reddish-brown, and others
snow-white: none had a trace of wing-bars or of the white croup. I then paired
together two of these mongrels, namely, a brown and black bird, and their
offspring displayed wing-bars, faint, but of a darker brown than the rest of
body. In a second brood from the same parents a brown bird was produced,
with several white feathers confined to the croup.
I crossed a male dun dragon belonging to a family which had been dun-
coloured without wing-bars during several generations, with a uniform red
barb (bred from two black barbs); and the offspring presented decided but faint
traces of wing-bars. I crossed a uniform red male runt with a white trumpeter;
and the offspring had a slaty-blue tail, with a bar at the end, and with the outer
feathers edged with white. I also crossed a female black and white chequered
trumpeter (of a different strain from the last) with a male almond-tumbler,
neither of which exhibited a trace of blue, or of the white croup, or of the bar
at end of tail: nor is it probable that the progenitors of these two birds had for
many generations exhibited any of these characters, for I have never even
heard of a blue trumpeter in this country, and my almond-tumbler was purely
bred; yet the tail of this mongrel was bluish, with a broad black bar at the end,
and the croup was perfectly white. It may be observed in several of these
cases, that the tail first shows a tendency to become by reversion blue; and this

fact of the persistency of colour in the tail and tail-coverts[344] will surprise no
one who has attended to the crossing of pigeons.
The last case which I will give is the most curious. I paired a mongrel female
barb-fantail with a mongrel male barb-spot; neither of which mongrels had the
least blue about them. Let it be remembered that blue barbs are excessively
rare; that spots, as has been already stated, were perfectly characterized in the
year 1676, and breed perfectly true; this likewise is the case with white
fantails, so much so that I have never heard of white fantails throwing any
other colour. Nevertheless the offspring from the above two mongrels was of
exactly the same blue tint as that of the wild rock-pigeon from the Shetland
Islands over the whole back and wings; the double black wing-bars were
equally conspicuous; the tail was exactly alike in all its characters, and the
croup was pure white; the head, however, was tinted with a shade of red,
evidently derived from the spot, and was of a paler blue than in the rock-
pigeon, as was the stomach. So that two black barbs, a red spot, and a white
fantail, as the four purely-bred grandparents, produced a bird of the same
general blue colour, together with every characteristic mark, as in the wild
Columba livia.
With respect to crossed breeds frequently producing blue birds chequered with
black, and resembling in all respects both the dovecot-pigeon and the
chequered wild variety of the rock-pigeon, the statement before referred to by
MM. Boitard and Corbié would almost suffice; but I will give three instances
of the appearance of such birds from crosses in which one alone of the parents
or great-grandparents was blue, but not chequered. I crossed a male blue turbit
with a snow-white trumpeter, and the following year with a dark, leaden-
brown, short-faced tumbler; the offspring from the first cross were as perfectly
chequered as any dovecot-pigeon; and from the second, so much so as to be
nearly as black as the most darkly chequered rock-pigeon from Madeira.
Another bird, whose great-grandparents were a white trumpeter, a white
fantail, a white red-spot, a red runt, and a blue pouter, was slaty-blue and
chequered exactly like a dovecot-pigeon. I may here add a remark made to me
by Mr. Wicking, who has had more experience than any other person in
England in breeding pigeons of various colours: namely, that when a blue, or a
blue and chequered bird, having black wing-bars, once appears in any race and
is allowed to breed, these characters are so strongly transmitted that it is
extremely difficult to eradicate them.

What, then, are we to conclude from this tendency in all the chief domestic
races, both when purely bred and more especially when intercrossed, to
produce offspring of a blue colour, with the same characteristic marks, varying
in the same manner, as in Columba livia? If we admit that these races have all
descended from C. livia, no breeder will doubt that the occasional appearance
of blue birds thus characterised is accounted for on the well-known principle
of "throwing back" or reversion. Why crossing should give so strong a
tendency to reversion, we do not with certainty know; but abundant evidence
of this fact will be given in the following chapters. It is probable that I might
have bred even for a century pure black barbs, spots, nuns, white fantails,
trumpeters, &c., without obtaining a single blue or barred bird; yet by crossing
these breeds I reared in the first and second generation, during the course of
only three or four years, a considerable number of young birds, more or less
plainly coloured blue, and with most of the characteristic marks. When black
and white, or black and red birds, are crossed, it would appear that a slight
tendency exists in both parents to produce blue offspring, and that this, when
combined, overpowers the separate tendency in either parent to produce black,
or white, or red offspring.
If we reject the belief that all the races of the pigeon are the modified
descendants of C. livia, and suppose that they are descended from several
aboriginal stocks, then we must choose between the three following
assumptions: firstly, that at least eight or nine species formerly existed which
were aboriginally coloured in various ways, but have since varied in so exactly
the same manner as to assume the colouring of C. livia; but this assumption
throws not the least light on the appearance of such colours and marks when
the races are crossed. Or secondly, we may assume that the aboriginal species
were all coloured blue, and had the wing-bars and other characteristic marks of
C. livia,—a supposition which is highly improbable, as besides this one
species no existing member of the Columbidæ presents these combined
characters; and it would not be possible to find any other instance of several
species identical in plumage, yet as different in important points of structure as
are pouters, fantails, carriers, tumblers, &c. Or lastly, we may assume that all
the races, whether descended from C. livia or from several aboriginal species,
although they have been bred with so much care and are so highly valued by
fanciers, have all been crossed within a dozen or score of generations with C.
livia, and have thus acquired their tendency to produce blue birds with the
several characteristic marks. I have said that it must be assumed that each race
has been crossed with C. livia within a dozen, or, at the utmost, within a score

of generations; for there is no reason to believe that crossed offspring ever
revert to one of their ancestors when removed by a greater number of
generations. In a breed which has been crossed only once, the tendency to
reversion will naturally become less and less in the succeeding generations, as
in each there will be less and less of the blood of the foreign breed; but when
there has been no cross with a distinct breed, and there is a tendency in both
parents to revert to some long-lost character, this tendency, for all that we can
see to the contrary, may be transmitted undiminished for an indefinite number
of generations. These two distinct cases of reversion are often confounded
together by those who have written on inheritance.
Considering, on the one hand, the improbability of the three assumptions
which have just been discussed, and, on the other hand, how simply the facts
are explained on the principle of reversion, we may conclude that the
occasional appearance in all the races, both when purely bred and more
especially when crossed, of blue birds, sometimes chequered, with double
wing-bars, with white or blue croups, with a bar at the end of the tail, and with
the outer tail-feathers edged with white, affords an argument of the greatest
weight in favour of the view that all are descended from Columba livia,
including under this name the three or four wild varieties or sub-species before
enumerated.
To sum up the six foregoing arguments, which are opposed to the belief that
the chief domestic races are the descendants of at least eight or nine or perhaps
a dozen species; for the crossing of any less number would not yield the
characteristic differences between the several races. Firstly, the improbability
that so many species should still exist somewhere, but be unknown to
ornithologists, or that they should have become within the historical period
extinct, although man has had so little influence in exterminating the wild C.
livia. Secondly, the improbability of man in former times having thoroughly
domesticated and rendered fertile under confinement so many species. Thirdly,
these supposed species having nowhere become feral. Fourthly, the
extraordinary fact that man should, intentionally or by chance, have chosen for
domestication several species, extremely abnormal in character; and
furthermore, the points of structure which render these supposed species so
abnormal being now highly variable. Fifthly, the fact of all the races, though
differing in many important points of structure, producing perfectly fertile
mongrels; whilst all the hybrids which have been produced between even
closely allied species in the pigeon-family are sterile. Sixthly, the remarkable

statements just given on the tendency in all the races, both when purely bred
and when crossed, to revert in numerous minute details of colouring to the
character of the wild rock-pigeon, and to vary in a similar manner. To these
arguments may be added the extreme improbability that a number of species
formerly existed, which differed greatly from each other in some few points,
but which resembled each other as closely as do the domestic races in other
points of structure, in voice, and in all their habits of life. When these several
facts and arguments are fairly taken into consideration, it would require an
overwhelming amount of evidence to make us admit that the chief domestic
races are descended from several aboriginal stocks; and of such evidence there
is absolutely none.
The belief that the chief domestic races are descended from several wild stocks
no doubt has arisen from the apparent improbability of such great
modifications of structure having been effected since man first domesticated
the rock-pigeon. Nor am I surprised at any degree of hesitation in admitting
their common origin: formerly, when I went into my aviaries and watched such
birds as pouters, carriers, barbs, fantails, and short-faced tumblers, &c., I could
not persuade myself that they had all descended from the same wild stock, and
that man had consequently in one sense created these remarkable
modifications. Therefore I have argued the question of their origin at great,
and, as some will think, superfluous length.
Finally, in favour of the belief that all the races are descended from a single
stock, we have in Columba livia a still existing and widely distributed species,
which can be and has been domesticated in various countries. This species
agrees in most points of structure and in all its habits of life, as well as
occasionally in every detail of plumage, with the several domestic races. It
breeds freely with them, and produces fertile offspring. It varies in a state of
nature,[345] and still more so when semi-domesticated, as shown by comparing
the Sierra Leone pigeons with those of India, or with those which apparently
have run wild in Madeira. It has undergone a still greater amount of variation
in the case of the numerous toy-pigeons, which no one supposes to be
descended from distinct species; yet some of these toy-pigeons have
transmitted their character truly for centuries. Why, then, should we hesitate to
believe in that greater amount of variation which is necessary for the
production of the eleven chief races? It should be borne in mind that in two of
the most strongly-marked races, namely, carriers and short-faced tumblers, the
extreme forms can be connected with the parent-species by graduated

differences not greater than those which may be observed between the
dovecot-pigeons inhabiting different countries, or between the various kinds of
toy-pigeons,—gradations which must certainly be attributed to variation.
That circumstances have been eminently favourable for the modification of the
pigeon through variation and selection will now be shown. The earliest record,
as has been pointed out to me by Professor Lepsius, of pigeons in a
domesticated condition, occurs in the fifth Egyptian dynasty, about 3000 B.C.;
[346] but Mr. Birch, of the British Museum, informs me that the pigeon appears

in a bill of fare in the previous dynasty. Domestic pigeons are mentioned in
Genesis, Leviticus, and Isaiah.[347] In the time of the Romans, as we hear from
Pliny,[348] immense prices were given for pigeons; "nay, they are come to this
pass, that they can reckon up their pedigree and race." In India, about the year
1600, pigeons were much valued by Akber Khan: 20,000 birds were carried
about with the court, and the merchants brought valuable collections. "The
monarchs of Iran and Turan sent him some very rare breeds. His Majesty,"
says the courtly historian, "by crossing the breeds, which method was never
practised before, has improved them astonishingly."[349] Akber Khan possessed
seventeen distinct kinds, eight of which were valuable for beauty alone. At
about this same period of 1600 the Dutch, according to Aldrovandi, were as
eager about pigeons as the Romans had formerly been. The breeds which were
kept during the fifteenth century in Europe and in India apparently differed
from each other. Tavernier, in his Travels in 1677, speaks, as does Chardin in
1735, of the vast number of pigeon-houses in Persia; and the former remarks
that, as Christians were not permitted to keep pigeons, some of the vulgar
actually turned Mahometans for this sole purpose. The Emperor of Morocco
had his favourite keeper of pigeons, as is mentioned in Moore's treatise,
published 1737. In England, from the time of Willughby in 1678 to the present
day, as well as in Germany and in France, numerous treatises have been
published on the pigeon. In India, about a hundred years ago, a Persian treatise
was written; and the writer thought it no light affair, for he begins with a
solemn invocation, "in the name of God, the gracious and merciful." Many
large towns, in Europe and the United States, now have their societies of
devoted pigeon-fanciers: at present there are three such societies in London. In
India, as I hear from Mr. Blyth, the inhabitants of Delhi and of some other
great cities are eager fanciers. Mr. Layard informs me that most of the known
breeds are kept in Ceylon. In China, according to Mr. Swinhoe of Amoy, and
Dr. Lockhart of Shangai, carriers, fantails, tumblers, and other varieties are

reared with care, especially by the bonzes or priests. The Chinese fasten a kind
of whistle to the tail-feathers of their pigeons, and as the flock wheels through
the air they produce a sweet sound. In Egypt the late Abbas Pacha was a great
fancier of fantails. Many pigeons are kept at Cairo and Constantinople, and
these have lately been imported by native merchants, as I hear from Sir W.
Elliot, into Southern India, and sold at high prices.
The foregoing statements show in how many countries, and during how long a
period, many men have been passionately devoted to the breeding of pigeons.
Hear how an enthusiastic fancier at the present day writes: "If it were possible
for noblemen and gentlemen to know the amazing amount of solace and
pleasure derived from Almond Tumblers, when they begin to understand their
properties, I should think that scarce any nobleman or gentleman would be
without their aviaries of Almond Tumblers."[350] The pleasure thus taken is of
paramount importance, as it leads amateurs carefully to note and preserve each
slight deviation of structure which strikes their fancy. Pigeons are often closely
confined during their whole lives; they do not partake of their naturally varied
diet; they have often been transported from one climate to another; and all
these changes in their conditions of life would be likely to cause variability.
Pigeons have been domesticated for nearly 5000 years, and have been kept in
many places, so that the numbers reared under domestication must have been
enormous; and this is another circumstance of high importance, for it
obviously favours the chance of rare modifications of structure occasionally
appearing. Slight variations of all kinds would almost certainly be observed,
and, if valued, would, owing to the following circumstances, be preserved and
propagated with unusual facility. Pigeons, differently from any other
domesticated animal, can easily be mated for life, and, though kept with other
pigeons, they rarely prove unfaithful to each other. Even when the male does
break his marriage-vow, he does not permanently desert his mate. I have bred
in the same aviaries many pigeons of different kinds, and never reared a single
bird of an impure strain. Hence a fancier can with the greatest ease select and
match his birds. He will also soon see the good results of his care; for pigeons
breed with extraordinary rapidity. He may freely reject inferior birds, as they
serve at an early age as excellent food. To sum up, pigeons are easily kept,
paired, and selected; vast numbers have been reared; great zeal in breeding
them has been shown by many men in various countries; and this would lead
to their close discrimination, and to a strong desire to exhibit some novelty, or
to surpass other fanciers in the excellence of already established breeds.

History of the principal Races of the Pigeon.[351]

Before discussing the means and steps by which the chief races have
been formed, it will be advisable to give some historical details, for
more is known of the history of the pigeon, little though this be, than
of any other domesticated animal. Some of the cases are interesting
as proving how long domestic varieties may be propagated with
exactly the same or nearly the same characters; and other cases are
still more interesting as showing how slowly but steadily races have
been greatly modified during successive generations. In the last
chapter I stated that Trumpeters and Laughers, both so remarkable
for their voices, seem to have been perfectly characterized in 1735;
and Laughers were apparently known in India before the year 1600.
Spots in 1676, and Nuns in the time of Aldrovandi, before 1600,
were coloured exactly as they now are. Common Tumblers and
Ground Tumblers exhibited in India, before the year 1600, the same
extraordinary peculiarities of flight as at the present day, for they are
well described in the 'Ayeen Akbery.' These breeds may all have
existed for a much longer period; we know only that they were
perfectly characterized at the dates above given. The average length
of life of the domestic pigeon is probably about five or six years; if
so, some of these races have retained their character perfectly for at
least forty or fifty generations.
Pouters.—These birds, as far as a very short description serves for
comparison, appear to have been well characterized in Aldrovandi's
time,[352] before the year 1600. Length of body and length of leg are
at the present time the two chief points of excellence. In 1735 Moore
said (see Mr. J. M. Eaton's edition)—and Moore was a first-rate
fancier—that he once saw a bird with a body 20 inches in length,
"though 17 or 18 inches is reckoned a very good length;" and he has
seen the legs very nearly 7 inches in length, yet a leg 6½ or 6¾ long
"must be allowed to be a very good one." Mr. Bult, the most
successful breeder of Pouters in the world, informs me that at present
(1858) the standard length of the body is not less than 18 inches; but
he has measured one bird 19 inches in length, and has heard of 20
and 22 inches, but doubts the truth of these latter statements. The
standard length of the leg is now 7 inches, but Mr. Bult has recently
measured two of his own birds with legs 7½ long. So that in the 123

years which have elapsed since 1735 there has been hardly any
increase in the standard length of the body; 17 or 18 inches was
formerly reckoned a very good length, and now 18 inches is the
minimum standard; but the length of leg seems to have increased, as
Moore never saw one quite 7 inches long; now the standard is 7, and
two of Mr. Bult's birds measured 7½ inches in length. The extremely
slight improvement in Pouters, except in the length of the leg, during
the last 123 years, may be partly accounted for by the neglect which
they suffered, as I am informed by Mr. Bult, until within the last 20
or 30 years. About 1765[353] there was a change of fashion, stouter
and more feathered legs being preferred to thin and nearly naked
legs.
Fantails.—The first notice of the existence of this breed is in India,
before the year 1600, as given in the 'Ayeen Akbery;'[354] at this date,
judging from Aldrovandi, the breed was unknown in Europe. In 1677
Willughby speaks of a Fantail with 26 tail-feathers; in 1735 Moore
saw one with 36 feathers; and in 1824 MM. Boitard and Corbié
assert that in France birds can easily be found with 42 tail-feathers.
In England, the number of the tail-feathers is not at present so much
regarded as their upward direction and expansion. The general
carriage of the bird is likewise now much regarded. The old
descriptions do not suffice to show whether in these latter respects
there has been much improvement; but if fantails had formerly
existed with their heads and tails touching each other, as at the
present time, the fact would almost certainly have been noticed. The
Fantails which are now found in India probably show the state of the
race, as far as carriage is concerned, at the date of their introduction
into Europe; and some, said to have been brought from Calcutta,
which I kept alive, were in a marked manner inferior to our
exhibition birds. The Java Fantail shows the same difference in
carriage; and although Mr. Swinhoe has counted 18 and 24 tail-
feathers in his birds, a first-rate specimen sent to me had only 14 tail-
feathers.
Jacobins.—This breed existed before 1600, but the hood, judging
from the figure given by Aldrovandi, did not enclose the head nearly
so perfectly as at present: nor was the head then white; nor were the
wings and tail so long, but this last character might have been

overlooked by the rude artist. In Moore's time, in 1735, the Jacobin
was considered the smallest kind of pigeon, and the bill is said to be
very short. Hence either the Jacobin, or the other kinds with which it
was then compared, must have been since considerably modified; for
Moore's description (and it must be remembered that he was a first-
rate judge) is clearly not applicable, as far as size of body and length
of beak are concerned, to our present Jacobins. In 1795, judging
from Bechstein, the breed had assumed its present character.
Turbits.—It has generally been supposed by the older writers on
pigeons, that the Turbit is the Cortbeck of Aldrovandi; but if this be
the case, it is an extraordinary fact that the characteristic frill should
not have been noticed. The beak, moreover, of the Cortbeck is
described as closely resembling that of the Jacobin, which shows a
change in the one or the other race. The Turbit, with its characteristic
frill and bearing its present name, is described by Willughby in 1677;
and the bill is said to be like that of the bullfinch,—a good
comparison, but now more strictly applicable to the beak of the
Barb. The sub-breed called the Owl was well known in Moore's
time, in 1735.
Tumblers.—Common Tumblers, as well as Ground Tumblers, perfect
as far as tumbling is concerned, existed in India before the year
1600; and at this period diversified modes of flight, such as flying at
night, the ascent to a great height, and manner of descent, seem to
have been much attended to, as at the present time, in India.
Belon[355] in 1555 saw in Paphlagonia what he describes as "a very
new thing, viz. pigeons which flew so high in the air that they were
lost to view, but returned to their pigeon-house without separating."
This manner of flight is characteristic of our present Tumblers, but it
is clear that Belon would have mentioned the act of tumbling if the
pigeons described by him had tumbled. Tumblers were not known in
Europe in 1600, as they are not mentioned by Aldrovandi, who
discusses the flight of pigeons. They are briefly alluded to by
Willughby, in 1687, as small pigeons "which show like footballs in
the air." The short-faced race did not exist at this period, as
Willughby could not have overlooked birds so remarkable for their
small size and short beaks. We can even trace some of the steps by
which this race has been produced. Moore in 1735 enumerates

correctly the chief points of excellence, but does not give any
description of the several sub-breeds; and from this fact Mr. Eaton
infers[356] that the short-faced Tumbler had not then come to full
perfection. Moore even speaks of the Jacobin as being the smallest
pigeon. Thirty years afterwards, in 1765, in the Treatise dedicated to
Mayor, short-faced Almond Tumblers are fully described, but the
author, an excellent fancier, expressly states in his Preface (p. xiv.)
that, "from great care and expense in breeding them, they have
arrived to so great perfection and are so different from what they
were 20 or 30 years past, that an old fancier would have condemned
them for no other reason than because they are not like what used to
be thought good when he was in the fancy before." Hence it would
appear that there was a rather sudden change in the character of the
short-faced Tumbler at about this period; and there is reason to
suspect that a dwarfed and half-monstrous bird, the parent-form of
the several short-faced sub-breeds, then appeared. I suspect this
because short-faced Tumblers are born with their beaks (ascertained
by careful measurement) as short, proportionally with the size of
their bodies, as in the adult bird; and in this respect they differ
greatly from all other breeds, which slowly acquire during growth
their various characteristic qualities.
Since the year 1765 there has been some change in one of the chief
characters of the short-faced Tumbler, namely, in the length of the
beak. Fanciers measure the "head and beak" from the tip of the beak
to the front corner of the eyeball. About the year 1765 a "head and
beak" was considered good,[357] which, measured in the usual
manner, was 7/8 of an inch in length; now it ought not to exceed 5/8
of an inch; "it is however possible," as Mr. Eaton candidly confesses,
"for a bird to be considered as pleasant or neat even at 6/8 of an inch,
but exceeding that length it must be looked upon as unworthy of
attention." Mr. Eaton states that he has never seen in the course of
his life more than two or three birds with the "head and beak" not
exceeding half an inch in length; "still I believe in the course of a
few years that the head and beak will be shortened, and that half-
inch birds will not be considered so great a curiosity as at the present
time." That Mr. Eaton's opinion deserves attention cannot be
doubted, considering his success in winning prizes at our
exhibitions. Finally in regard to the Tumbler it may be concluded

from the facts above given that it was originally introduced into
Europe, probably first into England, from the East; and that it then
resembled our common English Tumbler, or more probably the
Persian or Indian Tumbler, with a beak only just perceptibly shorter
than that of the common dovecot-pigeon. With respect to the short-
faced Tumbler, which is not known to exist in the East, there can
hardly be a doubt that the whole wonderful change in the size of the
head, beak, body, and feet, and in general carriage, has been
produced during the last two centuries by continued selection, aided
probably by the birth of a semi-monstrous bird somewhere about the
year 1750.
Runts.—Of their history little can be said. In the time of Pliny the
pigeons of Campania were the largest known; and from this fact
alone some authors assert that they were Runts. In Aldrovandi's time,
in 1600, two sub-breeds existed; but one of them, the short-beaked,
is now extinct in Europe.
Barbs.—Notwithstanding statements to the contrary, it seems to me
impossible to recognise the barb in Aldrovandi's descriptions and
figures; four breeds, however, existed in the year 1600 which were
evidently allied both to Barbs and Carriers. To show how difficult it
is to recognise some of the breeds described by Aldrovandi, I will
give the different opinions in regard to the above four kinds, named
by him C. Indica, Cretensis, Gutturosa, and Persica. Willughby
thought that the Columba Indica was a Turbit, but the eminent
fancier Mr. Brent believes that it was an inferior Barb: C. Cretensis,
with a short beak and a swelling on the upper mandible, cannot be
recognised: C. (falsely called) gutturosa, which from its rostrum,
breve, crassum, et tuberosum seems to me to come nearest to the
Barb, Mr. Brent believes to be a Carrier; and lastly, the C. Persica et
Turcica, Mr. Brent thinks, and I quite concur with him, was a short-
beaked Carrier with very little wattle. In 1687 the Barb was known
in England, and Willughby describes the beak as like that of the
Turbit; but it is not credible that his Barb should have had a beak like
that of our present birds, for so accurate an observer could not have
overlooked its great breadth.
English Carrier.—We may look in vain in Aldrovandi's work for any
bird resembling our prize Carriers; the C. Persica et Turcica of this

author comes the nearest, but is said to have had a short thick beak;
therefore it must have approached in character a Barb, and have
differed greatly from our Carriers. In Willughby's time, in 1677, we
can clearly recognise the Carrier, but he adds, "the bill is not short,
but of a moderate length," a description which no one would apply to
our present Carriers, so conspicuous for the extraordinary length of
their beaks. The old names given in Europe to the Carrier, and the
several names now in use in India, indicate that Carriers originally
came from Persia; and Willughby's description would perfectly apply
to the Bussorah Carrier as it now exists in Madras. In later times we
can partially trace the progress of change in our English Carriers:
Moore in 1735 says "an inch and a half is reckoned a long beak,
though there are very good Carriers that are found not to exceed an
inch and a quarter." These birds must have resembled, or perhaps
been a little superior to, the Carriers, previously described, which are
now found in Persia. In England at the present day "there are," as
Mr. Eaton[358] states, "beaks that would measure (from edge of eye
to tip of beak) one inch and three-quarters, and some few even two
inches in length."

From these historical details we see that nearly all the chief domestic races
existed before the year 1600. Some remarkable only for colour appear to have
been identical with our present breeds, some were nearly the same, some
considerably different, and some have since become extinct. Several breeds,
such as Finnikins and Turners, the swallow-tailed pigeon of Bechstein and the
Carmelite, seem both to have originated and to have disappeared within this
same period. Any one now visiting a well-stocked English aviary would
certainly pick out as the most distinct kinds, the massive Runt, the Carrier with
its wonderfully elongated beak and great wattles, the Barb with its short broad
beak and eye-wattles, the short-faced Tumbler with its small conical beak, the
Pouter with its great crop, long legs and body, the Fantail with its upraised,
widely-expanded, well-feathered tail, the Turbit with its frill and short blunt
beak, and the Jacobin with its hood. Now, if this same person could have
viewed the pigeons kept before 1600 by Akber Khan in India and by
Aldrovandi in Europe, he would have seen the Jacobin with a less perfect
hood; the Turbit apparently without its frill; the Pouter with shorter legs, and in
every way less remarkable—that is, if Aldrovandi's Pouter resembled the old
German kind; the Fantail would have been far less singular in appearance, and
would have had much fewer feathers in its tail; he would have seen excellent

flying Tumblers, but he would in vain have looked for the marvellous short-
faced breeds; he would have seen birds allied to barbs, but it is extremely
doubtful whether he would have met with our actual Barbs; and lastly, he
would have found Carriers with beaks and wattle incomparably less developed
than in our English Carriers. He might have classed most of the breeds in the
same groups as at present; but the differences between the groups were then far
less strongly pronounced than at present. In short, the several breeds had at this
early period not diverged in so great a degree from their aboriginal common
parent, the wild rock-pigeon.

Manner of Formation of the chief Races.
We will now consider more closely the probable steps by which the chief races
have been formed. As long as pigeons are kept semi-domesticated in dovecots
in their native country, without any care in selecting and matching them, they
are liable to little more variation than the wild C. livia, namely, in the wings
becoming chequered with black, in the croup being blue or white, and in the
size of the body. When, however, dovecot-pigeons are transported into
diversified countries, such as Sierra Leone, the Malay archipelago, and
Madeira (where the wild C. livia is not known to exist), they are exposed to
new conditions of life; and apparently in consequence they vary in a somewhat
greater degree. When closely confined, either for the pleasure of watching
them, or to prevent their straying, they must be exposed, even under their
native climate, to considerably different conditions; for they cannot obtain
their natural diversity of food; and, what is probably more important, they are
abundantly fed, whilst debarred from taking much exercise. Under these
circumstances we might expect to find, from the analogy of all other
domesticated animals, a greater amount of individual variability than with the
wild pigeon; and this is the case. The want of exercise apparently tends to
reduce the size of the feet and organs of flight; and then, from the law of
correlation of growth, the beak apparently becomes affected. From what we
now see occasionally taking place in our aviaries, we may conclude that
sudden variations or sports, such as the appearance of a crest of feathers on the
head, of feathered feet, of a new shade of colour, of an additional feather in the
tail or wing, would occur at rare intervals during the many centuries which
have elapsed since the pigeon was first domesticated. At the present day such
"sports" are generally rejected as blemishes; and there is so much mystery in
the breeding of pigeons that, if a valuable sport did occur, its history would
often be concealed. Before the last hundred and fifty years, there is hardly a

chance of the history of any such sport having been recorded. But it by no
means follows from this that such sports in former times, when the pigeon had
undergone much less variation, would have been rejected. We are profoundly
ignorant of the cause of each sudden and apparently spontaneous variation, as
well as of the infinitely numerous shades of difference between the birds of the
same family. But in a future chapter we shall see that all such variations appear
to be the indirect result of changes of some kind in the conditions of life.
Hence, after a long course of domestication, we might expect to see in the
pigeon much individual variability, and occasional sudden variations, as well
as slight modifications from the lessened use of certain parts, together with the
effects of correlation of growth. But without selection all this would produce
only a trifling or no result; for without such aid differences of all kinds would,
from the two following causes, soon disappear. In a healthy and vigorous lot of
pigeons many more young birds are killed for food or die than are reared to
maturity; so that an individual having any peculiar character, if not selected,
would run a good chance of being destroyed; and if not destroyed, the
peculiarity in question would almost certainly be obliterated by free
intercrossing. It might, however, occasionally happen that the same variation
repeatedly occurred, owing to the action of peculiar and uniform conditions of
life, and in this case it would prevail independently of selection. But when
selection is brought into play all is changed; for this is the foundation-stone in
the formation of new races; and with the pigeon, circumstances, as we have
already seen, are eminently favourable for selection. When a bird presenting
some conspicuous variation has been preserved, and its offspring have been
selected, carefully matched, and again propagated, and so onwards during
successive generations, the principle is so obvious that nothing more need be
said about it. This may be called methodical selection, for the breeder has a
distinct object in view, namely, to preserve some character which has actually
appeared; or to create some improvement already pictured in his mind.
Another form of selection has hardly been noticed by those authors who have
discussed this subject, but is even more important. This form may be called
unconscious selection, for the breeder selects his birds unconsciously,
unintentionally, and without method, yet he surely though slowly produces a
great result. I refer to the effects which follow from each fancier at first
procuring and afterwards rearing as good birds as he can, according to his
skill, and according to the standard of excellence at each successive period. He
does not wish permanently to modify the breed; he does not look to the distant

future, or speculate on the final result of the slow accumulation during many
generations of successive slight changes: he is content if he possesses a good
stock, and more than content if he can beat his rivals. The fancier in the time of
Aldrovandi, when in the year 1600 he admired his own jacobins, pouters, or
carriers, never reflected what their descendants in the year 1860 would
become; he would have been astonished could he have seen our jacobins, our
improved English carriers, and our pouters; he would probably have denied
that they were the descendants of his own once admired stock, and he would
perhaps not have valued them, for no other reason, as was written in 1765,
"than because they were not like what used to be thought good when he was in
the fancy." No one will attribute the lengthened beak of the carrier, the
shortened beak of the short-faced tumbler, the lengthened leg of the pouter, the
more perfectly-enclosed hood of the jacobin, &c.,—changes effected since the
time of Aldrovandi, or even since a much later period,—to the direct and
immediate action of the conditions of life. For these several races have been
modified in various and even in directly opposite ways, though kept under the
same climate and treated in all respects in as nearly uniform a manner as
possible. Each slight change in the length or shortness of the beak, in the
length of leg, &c., has no doubt been indirectly and remotely caused by some
change in the conditions to which the bird has been subjected, but we must
attribute the final result, as is manifest in those cases of which we have any
historical record, to the continued selection and accumulation of many slight
successive variations.
The action of unconscious selection, as far as pigeons are concerned, depends
on a universal principle in human nature, namely, on our rivalry, and desire to
outdo our neighbours. We see this in every fleeting fashion, even in our dress,
and it leads the fancier to endeavour to exaggerate every peculiarity in his
breeds. A great authority on pigeons[359] says, "Fanciers do not and will not
admire a medium standard, that is, half and half, which is neither here nor
there, but admire extremes." After remarking that the fancier of short-faced
beard tumblers wishes for a very short beak, and that the fancier of long-faced
beard tumblers wishes for a very long beak, he says, with respect to one of
intermediate length, "Don't deceive yourself. Do you suppose for a moment the
short or the long-faced fancier would accept such a bird as a gift? Certainly
not; the short-faced fancier could see no beauty in it; the long-faced fancier
would swear there was no use in it, &c." In these comical passages, written
seriously, we see the principle which has ever guided fanciers, and has led to

such great modifications in all the domestic races which are valued solely for
their beauty or curiosity.
Fashions in pigeon-breeding endure for long periods; we cannot change the
structure of a bird as quickly as we can the fashion of our dress. In the time of
Aldrovandi, no doubt the more the pouter inflated his crop, the more he was
valued. Nevertheless, fashions do to a certain extent change; first one point of
structure and then another is attended to; or different breeds are admired at
different times and in different countries. As the author just quoted remarks,
"the fancy ebbs and flows; a thorough fancier now-a-days never stoops to
breed toy-birds;" yet these very "toys" are now most carefully bred in
Germany. Breeds which at the present time are highly valued in India are
considered worthless in England. No doubt, when breeds are neglected, they
degenerate; still we may believe that, as long as they are kept under the same
conditions of life, characters once gained will be partially retained for a long
time, and may form, the starting-point for a future course of selection.
Let it not be objected to this view of the action of unconscious selection that
fanciers would not observe or care for extremely slight differences. Those
alone who have associated with fanciers can be thoroughly aware of their
accurate powers of discrimination acquired by long practice, and of the care
and labour which they bestow on their birds. I have known a fancier
deliberately study his birds day after day to settle which to match together and
which to reject. Observe how difficult the subject appears to one of the most
eminent and experienced fanciers. Mr. Eaton, the winner of many prizes, says,
"I would here particularly guard you against keeping too great a variety of
pigeons, otherwise you will know a little about all the kinds, but nothing about
one as it ought to be known." "It is possible there may be a few fanciers that
have a good general knowledge of the several fancy pigeons, but there are
many who labour under the delusion of supposing they know what they do
not." Speaking exclusively of one sub-variety of one race, namely, the short-
faced almond tumbler, and after saying that some fanciers sacrifice every
property to obtain a good head and beak, and that other fanciers sacrifice
everything for plumage, he remarks: "Some young fanciers who are over
covetous go in for all the five properties at once, and they have their reward by
getting nothing." In India, as I hear from Mr. Blyth, pigeons are likewise
selected and matched with the greatest care. But we must not judge of the
slight differences which would have been valued in ancient days, by those
which are now valued after the formation of many races, each with its own

standard of perfection, kept uniform by our numerous Exhibitions. The
ambition of the most energetic fancier may be fully satisfied by the difficulty
of excelling other fanciers in the breeds already established, without trying to
form a new one.

A difficulty with respect to the power of selection will perhaps already have
occurred to the reader, namely, what could have led fanciers first to attempt to
make such singular breeds as pouters, fantails, carriers, &c.? But it is this very
difficulty which the principle of unconscious selection removes. Undoubtedly
no fancier ever did intentionally make such an attempt. All that we need
suppose is that a variation occurred sufficiently marked to catch the
discriminating eye of some ancient fancier, and then unconscious selection
carried on for many generations, that is, the wish of succeeding fanciers to
excel their rivals, would do the rest. In the case of the fantail we may suppose
that the first progenitor of the breed had a tail only slightly erected, as may
now be seen in certain runts,[360] with some increase in the number of the tail-
feathers, as now occasionally occurs with nuns. In the case of the pouter we
may suppose that some bird inflated its crop a little more than other pigeons,
as is now the case in a slight degree with the œsophagus of the turbit. We do
not in the least know the origin of the common tumbler, but we may suppose
that a bird was born with some affection of the brain, leading it to make
somersaults in the air; and the difficulty in this case is lessened, as we know
that, before the year 1600, in India, pigeons remarkable for their diversified
manner of flight were much valued, and by the order of the Emperor Akber
Khan were sedulously trained and carefully matched.

In the foregoing cases we have supposed that a sudden variation,
conspicuous enough to catch a fancier's eye, first appeared; but even this
degree of abruptness in the process of variation is not necessary for the
formation of a new breed. When the same kind of pigeon has been kept
pure, and has been bred during a long period by two or more fanciers, slight
differences in the strain can often be recognised. Thus I have seen first-rate
jacobins in one man's possession which certainly differed slightly in several
characters from those kept by another. I possessed some excellent barbs
descended from a pair which had won a prize, and another lot descended
from a stock formerly kept by that famous fancier Sir John Sebright, and
these plainly differed in the form of the beak; but the differences were so
slight, that they could hardly be described by words. Again, the common
English and Dutch tumbler differ in a somewhat greater degree, both in
length of beak and shape of head. What first caused these slight differences
cannot be explained any more than why one man has a long nose and
another a short one. In the strains long kept distinct by different fanciers,
such differences are so common that they cannot be accounted for by the
accident of the birds first chosen for breeding having been originally as
different as they now are. The explanation no doubt lies in selection of a
slightly different nature having been applied in each case; for no two
fanciers have exactly the same taste, and consequently no two, in choosing
and carefully matching their birds, prefer or select exactly the same. As
each man naturally admires his own birds, he goes on continually
exaggerating by selection whatever slight peculiarities they may possess.
This will more especially happen with fanciers living in different countries,
who do not compare their stocks and aim at a common standard of
perfection. Thus, when a mere strain has once been formed, unconscious
selection steadily tends to augment the amount of difference, and thus
converts the strain into a sub-breed, and this ultimately into a well-marked
breed or race.
The principle of correlation of growth should never be lost sight of. Most
pigeons have small feet, apparently caused by their lessened use, and from
correlation, as it would appear, their beaks have likewise become reduced in
length. The beak is a conspicuous organ, and, as soon as it had thus become
perceptibly shortened, fanciers would almost certainly strive to reduce it
still more by the continued selection of birds with the shortest beaks; whilst

at the same time other fanciers, as we know has actually been the case,
would, in other sub-breeds, strive to increase its length. With the increased
length of the beak, the tongue would become greatly lengthened, as would
the eyelids with the increased development of the eye-wattles; with the
reduced or increased size of the feet the number of the scutellæ would vary;
with the length of the wing the number of the primary wing-feathers would
differ; and with the increased length of the body in the pouter the number of
the sacral vertebræ would be augmented. These important and correlated
differences of structure do not invariably characterise any breed; but if they
had been attended to and selected with as much care as the more
conspicuous external differences, there can hardly be a doubt that they
would have been rendered constant. Fanciers could assuredly have made a
race of tumblers with nine instead of ten primary wing-feathers, seeing how
often the number nine appears without any wish on their part, and indeed in
the case of the white-winged varieties in opposition to their wish. In a
similar manner, if the vertebræ had been visible and had been attended to by
fanciers, assuredly an additional number might easily have been fixed in the
pouter. If these latter characters had once been rendered constant we should
never have suspected that they had at first been highly variable, or that they
had arisen from correlation, in the one case with the shortness of the wings,
and in the other case with the length of the body.
In order to understand how the chief domestic races have become distinctly
separated from each other, it is important to bear in mind, that fanciers
constantly try to breed from the best birds, and consequently that those
which are inferior in the requisite qualities are in each generation neglected;
so that after a time the less improved parent-stocks and many subsequently
formed intermediate grades become extinct. This has occurred in the case of
the pouter, turbit, and trumpeter, for these highly improved breeds are now
left without any links closely connecting them either with each other or
with the aboriginal rock-pigeon. In other countries, indeed, where the same
care has not been applied, or where the same fashion has not prevailed, the
earlier forms may long remain unaltered or altered only in a slight degree,
and we are thus sometimes enabled to recover the connecting links. This is
the case in Persia and India with the tumbler and carrier, which there differ
but slightly from the rock-pigeon in the proportions of their beaks. So again
in Java, the fantail sometimes has only fourteen caudal feathers, and the tail

is much less elevated and expanded than in our improved birds; so that the
Java bird forms a link between a first-rate fantail and the rock-pigeon.
Occasionally a breed may be retained for some particular quality in a nearly
unaltered condition in the same country, together with highly modified
offshoots or sub-breeds, which are valued for some distinct property. We
see this exemplified in England, where the common tumbler, which is
valued only for its flight, does not differ much from its parent-form, the
Eastern tumbler; whereas the short-faced tumbler has been prodigiously
modified, from being valued, not for its flight, but for other qualities. But
the common-flying tumbler of Europe has already begun to branch out into
slightly different sub-breeds, such as the common English tumbler, the
Dutch roller, the Glasgow house-tumbler, and the long-faced beard tumbler,
&c.; and in the course of centuries, unless fashions greatly change, these
sub-breeds will diverge through the slow and insensible process of
unconscious selection, and become modified, in a greater and greater
degree. After a time the perfectly graduated links, which now connect all
these sub-breeds together, will be lost, for there would be no object and
much difficulty in retaining such a host of intermediate sub-varieties.
The principle of divergence, together with the extinction of the many
previously existing intermediate forms, is so important for understanding
the origin of domestic races, as well as of species in a state of nature, that I
will enlarge a little more on this subject. Our third main group includes
carriers, barbs, and runts, which are plainly related to each other, yet
wonderfully distinct in several important characters. According to the view
given in the last chapter, these three races have probably descended from an
unknown race having an intermediate character, and this from the rock-
pigeon. Their characteristic differences are believed to be due to different
breeders having at an early period admired different points of structure; and
then, on the acknowledged principle of admiring extremes, having gone on
breeding, without any thought of the future, as good birds as they could,—
carrier-fanciers preferring long beaks with much wattle,—barb-fanciers
preferring short thick beaks with much eye-wattle,—and runt-fanciers not
caring about the beak or wattle, but only for the size and weight of the body.
This process will have led to the neglect and final extinction of the earlier,
inferior, and intermediate birds; and thus it has come to pass, that in Europe
these three races are now so extraordinarily distinct from each other. But in

the East, whence they were originally brought, the fashion has been
different, and we there see breeds which connect the highly modified
English carrier with the rock-pigeon, and others which to a certain extent
connect carriers and runts. Looking back to the time of Aldrovandi, we find
that there existed in Europe, before the year 1600, four breeds which were
closely allied to carriers and barbs, but which competent authorities cannot
now identify with our present barbs and carriers; nor can Aldrovandi's runts
be identified with our present runts. These four breeds certainly did not
differ from each other nearly so much as do our existing English carriers,
barbs, and runts. All this is exactly what might have been anticipated. If we
could collect all the pigeons which have ever lived, from before the time of
the Romans to the present day, we should be able to group them in several
lines, diverging from the parent rock-pigeon. Each line would consist of
almost insensible steps, occasionally broken by some slightly greater
variation or sport, and each would culminate in one of our present highly
modified forms. Of the many former connecting links, some would be
found to have become absolutely extinct without having left any issue,
whilst others though extinct would be seen to be the progenitors of the
existing races.
I have heard it remarked as a strange circumstance that we occasionally
hear of the local or complete extinction of domestic races, whilst we hear
nothing of their origin. How, it has been asked, can these losses be
compensated, and more than compensated, for we know that with almost all
domesticated animals the races have largely increased in number since the
time of the Romans? But on the view here given, we can understand this
apparent contradiction. The extinction of a race within historical times is an
event likely to be noticed; but its gradual and scarcely sensible modification
through unconscious selection, and its subsequent divergence, either in the
same or more commonly in distant countries, into two or more strains, and
their gradual conversion into sub-breeds, and these into well-marked
breeds, are events which would rarely be noticed. The death of a tree, that
has attained gigantic dimensions, is recorded; the slow growth of smaller
trees and their increase in number excite no attention.
In accordance with the belief of the great power of selection, and of the
little direct power of changed conditions of life, except in causing general
variability or plasticity of organisation, it is not surprising that dovecot-

pigeons have remained unaltered from time immemorial; and that some toy-
pigeons, which differ in little else besides colour from the dovecot-pigeon,
have retained the same character for several centuries. For when one of
these toy-pigeons had once become beautifully and symmetrically coloured,
—when, for instance, a Spot had been produced with the crown of its head,
its tail, and tail-coverts of a uniform colour, the rest of the body being
snow-white,—no alteration or improvement would be desired. On the other
hand, it is not surprising that during this same interval of time our highly-
bred pigeons have undergone an astonishing amount of change; for in
regard to them there is no defined limit to the wish of the fancier, and there
is no known limit to the variability of their characters. What is there to stop
the fancier desiring to give to his carrier a longer and longer beak, or to his
tumbler a shorter and shorter beak? nor has the extreme limit of variability
in the beak, if there be any such limit, as yet been reached. Notwithstanding
the great improvement effected within recent times in the short-faced
almond tumbler, Mr. Eaton remarks, "the field is still as open for fresh
competitors as it was one hundred years ago;" but this is perhaps an
exaggerated assertion, for the young of all highly improved fancy birds are
extremely liable to disease and death.
I have heard it objected that the formation of the several domestic races of
the pigeon throws no light on the origin of the wild species of the
Columbidæ, because their differences are not of the same nature. The
domestic races for instance do not differ, or differ hardly at all, in the
relative lengths and shapes of the primary wing-feathers, in the relative
length of the hind toe, or in habits of life, as in roosting and building in
trees. But the above objection shows how completely the principle of
selection has been misunderstood. It is not likely that characters selected by
the caprice of man should resemble differences preserved under natural
conditions, either from being of direct service to each species, or from
standing in correlation with other modified and serviceable structures. Until
man selects birds differing in the relative length of the wing-feathers or
toes, &c., no sensible change in these parts should be expected. Nor could
man do anything unless these parts happened to vary under domestication: I
do not positively assert that this is the case, although I have seen traces of
such variability in the wing-feathers, and certainly in the tail-feathers. It
would be a strange fact if the relative length of the hind toe should never

vary, seeing how variable the foot is both in size and in the number of the
scutellæ. With respect to the domestic races not roosting or building in
trees, it is obvious that fanciers would never attend to or select such
changes in habits; but we have seen that the pigeons in Egypt, which do not
for some reason like settling on the low mud hovels of the natives, are led,
apparently by compulsion, to perch in crowds on the trees. We may even
affirm that, if our domestic races had become greatly modified in any of the
above specified respects, and it could be shown that fanciers had never
attended to such points, or that they did not stand in correlation with other
selected characters, the fact, on the principles advocated in this chapter,
would have offered a serious difficulty.
Let us briefly sum up the last two chapters on the pigeon. We may conclude
with confidence that all the domestic races, notwithstanding their great
amount of difference, are descended from the Columba livia, including
under this name certain wild races. But the differences between these latter
forms throw no light whatever on the characters which distinguish the
domestic races. In each breed or sub-breed the individual birds are more
variable than birds in a state of nature; and occasionally they vary in a
sudden and strongly-marked manner. This plasticity of organisation
apparently results from changed conditions of life. Disuse has reduced
certain parts of the body. Correlation of growth so ties the organisation
together, that when one part varies other parts vary at the same time. When
several breeds have once been formed, their intercrossing aids the progress
of modification, and has even produced new sub-breeds. But as, in the
construction of a building, mere stones or bricks are of little avail without
the builder's art, so, in the production of new races, selection has been the
presiding power. Fanciers can act by selection on excessively slight
individual differences, as well as on those greater differences which are
called sports. Selection is followed methodically when the fancier tries to
improve and modify a breed according to a prefixed standard of excellence;
or he acts unmethodically and unconsciously, by merely trying to rear as
good birds as he can, without any wish or intention to alter the breed. The
progress of selection almost inevitably leads to the neglect and ultimate
extinction of the earlier and less improved forms, as well as of many
intermediate links in each long line of descent. Thus it has come to pass that

most of our present races are so marvellously distinct from each other, and
from the aboriginal rock-pigeon.

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.

As some naturalists may not be familiar with the chief breeds of the fowl, it
will be advisable to give a condensed description of them.[361] From what I
have read and seen of specimens brought from several quarters of the
world, I believe that most of the chief kinds have been imported into
England, but many sub-breeds are probably still here unknown. The
following discussion on the origin of the various breeds and on their
characteristic differences does not pretend to completeness, but may be of
some interest to the naturalist. The classification of the breeds cannot, as far
as I can see, be made natural. They differ from each other in different
degrees, and do not afford characters in subordination to each other, by
which they can be ranked in group under group. They seem all to have
diverged by independent and different roads from a single type. Each chief
breed includes differently coloured sub-varieties, most of which can be
truly propagated, but it would be superfluous to describe them. I have
classed the various crested fowls as sub-breeds under the Polish fowl; but I
have great doubts whether this is a natural arrangement, showing true
affinity or blood relationship. It is scarcely possible to avoid laying stress
on the commonness of a breed; and if certain foreign sub-breeds had been

largely kept in this country they would perhaps have been raised to the rank
of main-breeds. Several breeds are abnormal in character; that is, they differ
in certain points from all wild Gallinaceous birds. At first I made a division
of the breeds into normal and abnormal, but the result was wholly
unsatisfactory.

Fig. 30.—Spanish Fowl.

1. Game Breed.—This may be considered as the typical breed, as
it deviates only slightly from the wild Gallus bankiva, or, as
perhaps more correctly named, ferrugineus. Beak strong; comb
single and upright. Spurs long and sharp. Feathers closely
adpressed to the body. Tail with the normal number of 14 feathers.
Eggs often pale-buff. Disposition indomitably courageous,
exhibited even in the hens and chickens. An unusual number of
differently coloured varieties exist, such as black and brown-
breasted reds, duckwings, blacks, whites, piles, &c., with their
legs of various colours.
2. Malay Breed.—Body of great size, with head, neck, and legs
elongated; carriage erect; tail small, sloping downwards,
generally formed of 16 feathers; comb and wattle small; ear-lobe

and face red; skin yellowish; feathers closely adpressed to the
body; neck-hackles short, narrow, and hard. Eggs often pale buff.
Chickens feather late. Disposition savage. Of Eastern origin.
3. Cochin, or Shangai Breed.—Size great; wing-feathers short,
arched, much hidden in the soft downy plumage; barely capable
of flight; tail short, generally formed of 16 feathers, developed at
a late period in the young males; legs thick, feathered; spurs short,
thick; nail of middle toe flat and broad; an additional toe not
rarely developed; skin yellowish. Comb and wattle well
developed. Skull with deep medial furrow; occipital foramen,
sub-triangular, vertically elongated. Voice peculiar. Eggs rough,
buff-coloured. Disposition extremely quiet. Of Chinese origin.
4. Dorking Breed.—Size great; body square, compact; feet with
an additional toe; comb well developed, but varies much in form;
wattles well developed; colour of plumage various. Skull
remarkably broad between the orbits. Of English origin.
The white Dorking may be considered as a distinct sub-breed,
being a less massive bird.
5. Spanish Breed.—Tall, with stately carriage; tarsi long; comb
single, deeply serrated, of immense size; wattles largely
developed; the large ear-lobes and sides of face white. Plumage
black glossed with green. Do not incubate. Tender in constitution,
the comb being often injured by frost. Eggs white, smooth, of
large size. Chickens feather late, but the young cocks show their
masculine characters, and crow at an early age. Of Mediterranean
origin.
The Andalusians may be ranked as a sub-breed: they are of a slaty
blue colour, and their chickens are well feathered. A smaller,
short-legged Dutch sub-breed has been described by some authors
as distinct.
6. Hamburgh Breed (fig. 31).—Size moderate; comb flat,
produced backwards, covered with numerous small points; wattle
of moderate dimensions; ear-lobe white; legs blueish, thin. Do not

incubate. Skull, with the tips of the ascending branches of the
premaxillary and with the nasal bones standing a little separate
from each other; anterior margin of the frontal bones less
depressed than usual.
There are two sub-breeds; the spangled Hamburgh, of English
origin, with the tips of the feathers marked with a dark spot; and
the pencilled Hamburgh, of Dutch origin, with dark transverse
lines across each feather, and with the body rather smaller. Both
these sub-breeds include gold and silver varieties, as well as some
other sub-varieties. Black Hamburghs have been produced by a
cross with the Spanish breed.
7. Crested or Polish Breed (fig. 32).—Head with a large,
rounded crest of feathers, supported on a hemispherical
protuberance of the frontal bones, which includes the anterior part
of the brain. The ascending branches of the premaxillary bones
and the inner nasal processes are much shortened. The orifice of
the nostrils raised and crescentic. Beak short. Comb absent, or
small and of crescentic shape; wattles either present or replaced
by a beard-like tuft of feathers. Legs leaden-blue. Sexual
differences appear late in life. Do not incubate. There are several
beautiful varieties which differ in colour and slightly in other
respects.

Fig. 31.—Hamburgh Fowl.

The following sub-breeds agree in having a crest, more or less
developed, with the comb, when present, of crescentic shape. The
skull presents nearly the same remarkable peculiarities of
structure as in the true Polish fowl.
Sub-breed (a) Sultans.—A Turkish breed, resembling white
Polish fowls, with a large crest and beard, with short and well-
feathered legs. The tail is furnished with additional sickle
feathers. Do not incubate.[362]
Sub-breed (b) Ptarmigans.—An inferior breed closely allied to
the last, white, rather small, legs much feathered, with the crest
pointed; comb small, cupped; wattles small.
Sub-breed (c) Ghoondooks.—Another Turkish breed having an
extraordinary appearance; black and tailless; crest and beard
large; legs feathered. The inner processes of the two nasal bones
come into contact with each other, owing to the complete
absorption of the ascending branches of the premaxillaries. I have
seen an allied, white, tailless breed from Turkey.

Fig. 32.—Polish Fowl.

Sub-breed (d) Crève-cœur.—A French breed of large size, barely
capable of flight, with short black legs, head crested, comb

produced into two points or horns, sometimes a little branched
like the horns of a stag; both beard and wattles present. Eggs
large. Disposition quiet.[363]
Sub-breed (e) Horned fowl.—With a small crest; comb produced
into two great points, supported on two bony protuberances.
Sub-breed (f) Houdan.—A French breed; of moderate size, short-
legged with five toes, wings well developed; plumage invariably
mottled with black, white, and straw-yellow; head furnished with
a crest, and a triple comb placed transversely; both wattles and
beard present.[364]
Sub-breed (g) Guelderlands.—No comb, head said to be
surmounted by a longitudinal crest of soft velvety feathers;
nostrils said to be crescentic; wattles well developed; legs
feathered; colour black. From North America. The Breda fowl
seems to be closely allied to the Guelderland.

8. Bantam Breed.—Originally from Japan,[365] characterized by
small size alone; carriage bold and erect. There are several sub-
breeds, such as the Cochin, Game, and Sebright Bantams, some
of which have been recently formed by various crosses. The
Black Bantam has a differently shaped skull, with the occipital
foramen like that of the Cochin fowl.

9. Rump-less Fowls.—These are so variable in character[366] that
they hardly deserve to be called a breed. Any one who will
examine the caudal vertebræ will see how monstrous the breed is.
10. Creepers or Jumpers.—These are characterized by an almost
monstrous shortness of legs, so that they move by jumping rather
than by walking; they are said not to scratch up the ground. I have
examined a Burmese variety, which had a skull of rather unusual
shape.
11. Frizzled or Caffre Fowls.—Not uncommon in India, with
the feathers curling backwards, and with the primary feathers of
the wing and tail imperfect; periosteum of bones black.

12. Silk Fowls.—Feathers silky, with the primary wing and tail-
feathers imperfect; skin and periosteum of bones black; comb and
wattles dark leaden-blue; ear-lappets tinged with blue; legs thin,
often furnished with an additional toe. Size rather small.
13. Sooty Fowls.—An Indian breed, of a white colour stained
with soot, with black skin and periosteum. The hens alone are
thus characterized.

From this synopsis we see that the several breeds differ considerably, and
they would have been nearly as interesting for us as pigeons, if there had
been equally good evidence that all had descended from one parent-species.
Most fanciers believe that they are descended from several primitive stocks.
The Rev. E. S. Dixon[367] argues strongly on this side of the question; and
one fancier even denounces the opposite conclusion by asking, "Do we not
perceive pervading this spirit, the spirit of the Deist?" Most naturalists, with
the exception of a few, such as Temminck, believe that all the breeds have
proceeded from a single species; but authority on such a point goes for
little. Fanciers look to all parts of the world as the possible sources of their
unknown stocks; thus ignoring the laws of geographical distribution. They
know well that the several kinds breed truly even in colour. They assert, but,
as we shall see, on very weak grounds, that most of the breeds are
extremely ancient. They are strongly impressed with the great difference
between the chief kinds, and they ask with force, can differences in climate,
food, or treatment have produced birds so different as the black stately
Spanish, the diminutive elegant Bantam, the heavy Cochin with its many
peculiarities, and the Polish fowl with its great top-knot and protuberant
skull? But fanciers, whilst admitting and even overrating the effects of
crossing the various breeds, do not sufficiently regard the probability of the
occasional birth, during the course of centuries, of birds with abnormal and
hereditary peculiarities; they overlook the effects of correlation of growth—
of the long-continued use and disuse of parts, and of some direct result from
changed food and climate, though on this latter head I have found no
sufficient evidence; and lastly, they all, as far as I know, entirely overlook
the all-important subject of unconscious or unmethodical selection, though
they are well aware that their birds differ individually, and that by selecting
the best birds for a few generations they can improve their stocks.

An amateur writes[368] as follows. "The fact that poultry have until lately
received but little attention at the hands of the fancier, and been entirely
confined to the domains of the producer for the market, would alone
suggest the improbability of that constant and unremitting attention having
been observed in breeding, which is requisite to the consummating, in the
offspring of any two birds, transmittable forms not exhibited by the
parents." This at first sight appears true. But in a future chapter on
Selection, abundant facts will be given showing not only that careful
breeding, but that actual selection was practised during ancient periods, and
by barely civilised races of man. In the case of the fowl I can adduce no
direct facts showing that selection was anciently practised; but the Romans
at the commencement of the Christian era kept six or seven breeds, and
Columella "particularly recommends as the best, those sorts that have five
toes and white ears."[369] In the fifteenth century several breeds were known
and described in Europe; and in China, at nearly the same period, seven
kinds were named. A more striking case is that at present, in one of the
Philippine Islands, the semi-barbarous inhabitants have distinct native
names for no less than nine sub-breeds of the Game Fowl.[370] Azara,[371]
who wrote towards the close of the last century, states that in the interior
parts of South America, where I should not have expected that the least care
would have been taken of poultry, a black-skinned and black-boned breed is
kept, from being considered fertile and its flesh good for sick persons. Now
every one who has kept poultry knows how impossible it is to keep several
breeds distinct unless the utmost care be taken in separating the sexes. Will
it then be pretended that those persons who in ancient times and in semi-
civilized countries took pains to keep the breeds distinct, and who therefore
valued them, would not occasionally have destroyed inferior birds and
occasionally have preserved their best birds? This is all that is required. It is
not pretended that any one in ancient times intended to form a new breed, or
to modify an old breed according to some ideal standard of excellence. He
who cared for poultry would merely wish to obtain, and afterwards to rear,
the best birds which he could; but this occasional preservation of the best
birds would in the course of time modify the breed, as surely, though by no
means as rapidly, as does methodical selection at the present day. If one
person out of a hundred or out of a thousand attended to the breeding of his
birds, this would be sufficient; for the birds thus tended would soon become
superior to others, and would form a new strain; and this strain would, as

explained in the last chapter, slowly have its characteristic differences
augmented, and at last be converted into a new sub-breed or breed. But
breeds would often be for a time neglected and would deteriorate; they
would, however, partially retain their character, and afterwards might again
come into fashion and be raised to a standard of perfection higher than their
former standard; as has actually occurred quite recently with Polish fowls.
If, however, a breed were utterly neglected, it would become extinct, as has
recently happened with one of the Polish sub-breeds. Whenever in the
course of past centuries a bird appeared with some slight abnormal
structure, such as with a lark-like crest on its head, it would probably often
have been preserved from that love of novelty which leads some persons in
England to keep rump-less fowls, and others in India to keep frizzled fowls.
And after a time any such abnormal appearance would be carefully
preserved, from being esteemed a sign of the purity and excellence of the
breed; for on this principle the Romans eighteen centuries ago valued the
fifth toe and the white ear-lobe in their fowls.
Thus from the occasional appearance of abnormal characters, though at first
only slight in degree; from the effects of the use and the disuse of parts;
possibly from the direct effects of changed climate and food; from
correlation of growth; from occasional reversions to old and long-lost
characters; from the crossing of breeds, when more than one had once been
formed; but, above all, from unconscious selection carried on during many
generations, there is no insuperable difficulty, to the best of my judgment,
in believing that all the breeds have descended from some one parent-
source. Can any single species be named from which we may reasonably
suppose that all have descended? The Gallus bankiva apparently fulfils
every requirement. I have already given as fair an account as I could of the
arguments in favour of the multiple origin of the several breeds; and now I
will give those in favour of their common descent from G. bankiva.

But it will be convenient first briefly to describe all the known
species of Gallus. The G. Sonneratii does not range into the
northern parts of India; according to Colonel Sykes,[372] it
presents at different heights on the Ghauts, two strongly marked
varieties, perhaps deserving to be called species. It was at one
time thought to be the primitive stock of all our domestic breeds,

and this shows that it closely approaches the common fowl in
general structure; but its hackles partially consist of highly
peculiar, horny laminæ, transversely banded with three colours;
and I have met with no authentic account of any such character
having been observed in any domestic breed.[373] This species
also differs greatly from the common fowl, in the comb being
finely serrated, and in the loins being destitute of true hackles. Its
voice is utterly different. It crosses readily in India with domestic
hens; and Mr. Blyth [374] raised nearly 100 hybrid chickens; but
they were tender and mostly died whilst young. Those which
were reared were absolutely sterile when crossed inter se, or with
either parent. At the Zoological Gardens, however, some hybrids
of the same parentage were not quite so sterile: Mr. Dixon, as he
informed me, made, with Mr. Yarrell's aid, particular inquiries on
this subject, and was assured that out of 50 eggs only five or six
chickens were reared. Some, however, of these half-bred birds
were crossed with one of their parents, namely, a Bantam, and
produced a few extremely feeble chickens. Mr. Dixon also
procured some of these same birds and crossed them in several
ways, but all were more or less infertile. Nearly similar
experiments have recently been tried on a great scale in the
Zoological Gardens with almost the same result.[375] Out of 500
eggs, raised from various first crosses and hybrids, between G.
Sonneratii, bankiva, and varius, only 12 chickens were reared,
and of these only three were the product of hybrids inter se. From
these facts, and from the above-mentioned strongly-marked
differences in structure between the domestic fowl and G.
Sonneratii, we may reject this latter species as the parent of any
domestic breed.
Ceylon possesses a fowl peculiar to the island, viz. G. Stanleyii;
this species approaches so closely (except in the colouring of the
comb) to the domestic fowl, that Messrs. E. Layard and
Kellaert[376] would have considered it, as they inform me, as one
of the parent-stocks, had it not been for its singularly different
voice. This bird, like the last, crosses readily with tame hens, and
even visits solitary farms and ravishes them. Two hybrids, a male

and female, thus produced, were found by Mr. Mitford to be quite
sterile: both inherited the peculiar voice of G. Stanleyii. This
species, then, may in all probability be rejected as one of the
primitive stocks of the domestic fowl.
Java and the islands eastward as far as Flores are inhabited by G.
varius (or furcatus), which differs in so many characters—green
plumage, unserrated comb, and single median wattle—that no one
supposes it to have been the parent of any one of our breeds; yet,
as I am informed by Mr. Crawfurd,[377] hybrids are commonly
raised between the male G. varius and the common hen, and are
kept for their great beauty, but are invariably sterile; this,
however, was not the case with some bred in the Zoological
Gardens. These hybrids were at one time thought to be
specifically distinct, and were named G. æneus. Mr. Blyth and
others believe that the G. Temminckii[378] (of which the history is
not known) is a similar hybrid. Sir J. Brooke sent me some skins
of domestic fowls from Borneo, and across the tail of one of
these, as Mr. Tegetmeier observed, there were transverse blue
bands like those which he had seen on the tail-feathers of hybrids
from G. varius, reared in the Zoological Gardens. This fact
apparently indicates that some of the fowls of Borneo have been
slightly affected by crosses with G. varius, but the case may
possibly be one of analogous variation. I may just allude to the G.
giganteus, so often referred to in works on poultry as a wild
species; but Marsden,[379] the first describer, speaks of it as a
tame breed; and the specimen in the British Museum evidently
has the aspect of a domestic variety.
The last species to be mentioned, namely, Gallus bankiva, has a
much wider geographical range than the three previous species; it
inhabits Northern India as far west as Sinde, and ascends the
Himalaya to a height of 4000 ft.; it inhabits Burmah, the Malay
peninsula, the Indo-Chinese countries, the Philippine Islands, and
the Malayan archipelago as far eastward as Timor. This species
varies considerably in the wild state. Mr. Blyth informs me that
the specimens, both male and female, brought from near the

Himalaya, are rather paler coloured than those from other parts of
India; whilst those from the Malay peninsula and Java are
brighter coloured than the Indian birds. I have seen specimens
from these countries, and the difference of tint in the hackles was
conspicuous. The Malayan hens were a shade redder on the breast
and neck than the Indian hens. The Malayan males generally had
a red ear-lappet, instead of a white one as in India; but Mr. Blyth
has seen one Indian specimen without the white ear-lappet. The
legs are leaden blue in the Indian, whereas they show some
tendency to be yellowish in the Malayan and Javan specimens. In
the former Mr. Blyth finds the tarsus remarkably variable in
length. According to Temminck[380] the Timor specimens differ as
a local race from that of Java. These several wild varieties have
not as yet been ranked as distinct species; if they should, as is not
unlikely, be hereafter thus ranked, the circumstance would be
quite immaterial as far as the parentage and differences of our
domestic breeds are concerned. The wild G. bankiva agrees most
closely with the black-breasted red Game-breed, in colouring and
in all other respects, except in being smaller, and in the tail being
carried more horizontally. But the manner in which the tail is
carried is highly variable in many of our breeds, for, as Mr. Brent
informs me, the tail slopes much in the Malays, is erect in the
Games and some other breeds, and is more than erect in
Dorkings, Bantams, &c. There is one other difference, namely,
that in G. bankiva, according to Mr. Blyth, the neck-hackles when
first moulted are replaced during two or three months, not by
other hackles, as with our domestic poultry, but by short blackish
feathers.[381] Mr. Brent, however, has remarked that these black
feathers remain in the wild bird after the development of the
lower hackles, and appear in the domestic bird at the same time
with them; so that the only difference is that the lower hackles are
replaced more slowly in the wild than in the tame bird; but as
confinement is known sometimes to affect the masculine
plumage, this slight difference cannot be considered of any
importance. It is a significant fact that the voice of both the male
and female G. bankiva closely resembles, as Mr. Blyth and others
have noted, the voice of both sexes of the common domestic

fowl; but the last note of the crow of the wild bird is rather less
prolonged. Captain Hutton, well known for his researches into the
natural history of India, informs me that he has seen several
crossed fowls from the wild species and the Chinese bantam;
these crossed fowls bred freely with bantams, but unfortunately
were not crossed inter se. Captain Hutton reared chickens from
the eggs of the Gallus bankiva; and these, though at first very
wild, afterwards became so tame that they would crowd round his
feet. He did not succeed in rearing them to maturity; but, as he
remarks, "no wild gallinaceous bird thrives well at first on hard
grain." Mr. Blyth also found much difficulty in keeping G.
bankiva in confinement. In the Philippine Islands, however, the
natives must succeed better, as they keep wild cocks to fight with
their domestic game-birds.[382] Sir Walter Elliot informs me that
the hen of a native domestic breed of Pegu is undistinguishable
from the hen of the wild G. bankiva; and the natives constantly
catch wild cocks by taking tame cocks to fight with them in the
woods.[383] Mr. Crawfurd remarks that from etymology it might
be argued that the fowl was first domesticated by the Malays and
Javanese.[384] It is also a curious fact, of which I have been
assured by Mr. Blyth, that wild specimens of the Gallus bankiva,
brought from the countries east of the Bay of Bengal, are far more
easily tamed than those of India; nor is this an unparalleled fact,
for, as Humboldt long ago remarked, the same species sometimes
evinces a more tameable disposition in one country than in
another. If we suppose that the G. bankiva was first tamed in
Malaya and afterwards imported into India, we can understand an
observation made to me by Mr. Blyth, that the domestic fowls of
India do not resemble the wild G. bankiva more closely than do
those of Europe.

From the extremely close resemblance in colour, general structure, and
especially in voice, between Gallus bankiva and the Game fowl; from their
fertility, as far as this has been ascertained, when crossed; from the
possibility of the wild species being tamed, and from its varying in the wild
state, we may confidently look at it as the parent of the most typical of all
the domestic breeds, namely, the Game-fowl. It is a significant fact, that

almost all the naturalists in India, namely, Sir W. Elliot, Mr. S. N. Ward, Mr.
Layard, Mr. J. C. Jerdon, and Mr. Blyth,[385] who are familiar with G.
bankiva, believe that it is the parent of most or all our domestic breeds. But
even if it be admitted that G. bankiva is the parent of the Game breed, yet it
may be urged that other wild species have been the parents of the other
domestic breeds; and that these species still exist, though unknown, in some
country, or have become extinct. The extinction, however, of several species
of fowls, is an improbable hypothesis, seeing that the four known species
have not become extinct in the most anciently and thickly peopled regions
of the East. There is, in fact, only one kind of domesticated bird, namely,
the Chinese goose or Anser cygnoides, of which the wild parent-form is said
to be still unknown, or extinct. For the discovery of new, or the rediscovery
of old species of Gallus, we must not look, as fanciers often look, to the
whole world. The larger gallinaceous birds, as Mr. Blyth has remarked,[386]
generally have a restricted range: we see this well illustrated in India, where
the genus Gallus inhabits the base of the Himalaya, and is succeeded higher
up by Gallophasis, and still higher up by Phasianus. Australia, with its
islands, is out of the question as the home for unknown species of the
genus. It is, also, as improbable that Gallus should inhabit South
America[387] as that a humming-bird should be found in the Old World.
From the character of the other gallinaceous birds of Africa, it is not
probable that Gallus is an African genus. We need not look to the western
parts of Asia, for Messrs. Blyth and Crawfurd, who have attended to this
subject, doubt whether Gallus ever existed in a wild state even as far west
as Persia. Although the earliest Greek writers speak of the fowl as a Persian
bird, this probably merely indicates its line of importation. For the
discovery of unknown species we must look to India, to the Indo-Chinese
countries, and to the northern parts of the Malay Archipelago. The southern
portion of China is the most likely country; but as Mr. Blyth informs me,
skins have been exported from China during a long period, and living birds
are largely kept there in aviaries, so that any native species of Gallus would
probably have become known. Mr. Birch, of the British Museum, has
translated for me passages from a Chinese Encyclopædia published in 1609,
but compiled from more ancient documents, in which it is said that fowls
are creatures of the West, and were introduced into the East (i.e. China) in a
dynasty 1400 B.C. Whatever may be thought of so ancient a date, we see that
the Indo-Chinese and Indian regions were formerly considered by the