The Project Gutenberg EBook of The Variation of Animals and Plants Under
Domestication, Vol. I., by Charles Darwin

This eBook is for the use of anyone anywhere at no cost and with
almost no restrictions whatsoever.  You may copy it, give it away or
re-use it under the terms of the Project Gutenberg License included
with this eBook or online at www.gutenberg.org


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

Author: Charles Darwin

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

Language: English

Character set encoding: ISO-8859-1

*** START OF THIS PROJECT GUTENBERG EBOOK VARIATION OF ANIMALS AND PLANTS ***




Produced by Steven Gibbs, Keith Edkins and the Online
Distributed Proofreading Team at http://www.pgdp.net





Transcriber's note: A few typographical errors have been corrected: they
are listed at the end of the text.

       *       *       *       *       *


THE VARIATION

OF

ANIMALS AND PLANTS

UNDER DOMESTICATION.

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

IN TWO VOLUMES.--VOL. I.

WITH ILLUSTRATIONS.

LONDON:

JOHN MURRAY, ALBEMARLE STREET.

1868.

_The right of Translation is reserved._

       *       *       *       *       *


BY THE SAME AUTHOR.

       *       *       *       *       *

ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or The PRESERVATION
of FAVOURED RACES in the STRUGGLE for LIFE. Fourth Edition (_Eighth
Thousand_), with Additions and Corrections. 1866. ... MURRAY.

A NATURALIST'S VOYAGE ROUND THE WORLD; or, A JOURNAL OF RESEARCHES into the
NATURAL HISTORY and GEOLOGY of the COUNTRIES visited during the Voyage of
H.M.S. Beagle, under the Command of Capt. FITZ-ROY, R.N. _Tenth Thousand_.
... MURRAY.

ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. ... SMITH, ELDER, & Co.

GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. ... SMITH, ELDER, & Co.

GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. ... SMITH, ELDER, & Co.

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

ON THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND FOREIGN ORCHIDS ARE
FERTILISED BY INSECTS; and on the GOOD EFFECTS of CROSSING. With numerous
Woodcuts. ... MURRAY.

ON THE MOVEMENTS and HABITS of CLIMBING PLANTS. With Woodcuts. ... WILLIAMS
& NORGATE.

       *       *       *       *       *

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

       *       *       *       *       *


{iii}

CONTENTS OF VOLUME I.

INTRODUCTION ... Page 1

CHAPTER I.

DOMESTIC DOGS AND CATS.

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

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

CHAPTER II.

HORSES AND ASSES.

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

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

CHAPTER III.

PIGS--CATTLE--SHEEP--GOATS.

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

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

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

GOATS.--REMARKABLE VARIATIONS OF ... Page 65

CHAPTER IV.

DOMESTIC RABBITS.

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

CHAPTER V.

DOMESTIC PIGEONS.

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

CHAPTER VI.

PIGEONS--_continued_.

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

{v}

CHAPTER VII.

FOWLS.

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

CHAPTER VIII.

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

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

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

PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.

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

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

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

CHAPTER IX.

CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.

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

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

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

{vi}

CHAPTER X.

PLANTS _continued_--FRUITS--ORNAMENTAL TREES--FLOWERS.

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

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

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

CHAPTER XI.

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

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

       *       *       *       *       *


{vii}

LIST OF ILLUSTRATIONS.

  1. 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
  {viii}
  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

       *       *       *       *       *


{1}

THE

VARIATION OF ANIMALS AND PLANTS

UNDER DOMESTICATION.

       *       *       *       *       *

INTRODUCTION.

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

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

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

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

I shall in this volume treat, as fully as my materials permit, the whole
subject of variation under domestication. We may thus hope to obtain some
light, little though it be, on the causes of variability,--on the laws
which govern it, such as the direct action of climate and food, the effects
of use and disuse, and of correlation of growth,--and on the amount of
change to which domesticated organisms are liable. We shall learn something
on the laws of inheritance, on the effects of crossing different breeds,
and on that sterility which often supervenes when organic beings are
removed from their natural conditions of life, and likewise when they are
too closely interbred. During this investigation we shall see that the
principle of Selection is all important. Although man does not cause
variability and cannot even prevent it, he can select, preserve, and
accumulate the variations given to him by the hand of nature in any way
which he chooses; and thus he can certainly produce a great result.
Selection may be followed either methodically and intentionally, or
unconsciously and unintentionally. Man {4} may select and preserve each
successive variation, with the distinct intention of improving and altering
a breed, in accordance with a preconceived idea; and by thus adding up
variations, often so slight as to be imperceptible by an uneducated eye, he
has effected wonderful changes and improvements. It can, also, be clearly
shown that man, without any intention or thought of improving the breed, by
preserving in each successive generation the individuals which he prizes
most, and by destroying the worthless individuals, slowly, though surely,
induces great changes. As the will of man thus comes into play, we can
understand how it is that domesticated breeds show adaptation to his wants
and pleasures. We can further understand how it is that domestic races of
animals and cultivated races of plants often exhibit an abnormal character,
as compared with natural species; for they have been modified not for their
own benefit, but for that of man.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

       *       *       *       *       *


{15}

CHAPTER I.

DOMESTIC DOGS AND CATS.

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

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

The first and chief point of interest in this chapter is, whether the
numerous domesticated varieties of the dog have descended from a single
wild species, or from several. Some authors believe that all have descended
from the wolf, or from the jackal, or from an unknown and extinct species.
Others again believe, and this of late has been the favourite tenet, that
they have descended from several species, extinct and recent, more or less
commingled together. We shall probably never be able to ascertain their
origin with certainty. Palæontology[5] does not throw much light on the
question, owing, on the one hand, to the close similarity of the skulls of
extinct as well as living wolves and jackals, and owing on the other hand
to the great dissimilarity of the skulls of the several breeds of the
domestic dogs. It seems, however, that remains have been found in the {16}
later tertiary deposits more like those of a large dog than of a wolf,
which favours the belief of De Blainville that our dogs are the descendants
of a single extinct species. On the other hand, some authors go so far as
to assert that every chief domestic breed must have had its wild prototype.
This latter view is extremely improbable; it allows nothing for variation;
it passes over the almost monstrous character of some of the breeds; and it
almost necessarily assumes, that a large number of species have become
extinct since man domesticated the dog; whereas we plainly see that the
members of the dog-family are extirpated by human agency with much
difficulty; even so recently as 1710 the wolf existed in so small an island
as Ireland.

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

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

We thus see that, at a period between four and five thousand years ago,
various breeds, viz. pariah dogs, greyhounds, common hounds, mastiffs,
house-dogs, lapdogs, and turnspits, existed, more or less closely
resembling our present breeds. But there is not sufficient evidence that
any of these ancient dogs belonged to the same identical sub-varieties with
our present dogs.[11] As long as man was believed to have existed on this
earth only about 6000 years, this fact of the great diversity of the breeds
at so early a period was an argument of much weight that they had proceeded
from several wild sources, for there would not have been sufficient time
for their divergence and modification. But now that we know, from the
discovery of flint tools embedded with the remains of extinct animals in
districts which have since undergone great geographical changes, that man
has existed for an incomparably longer period, and bearing in mind that the
most barbarous nations possess domestic dogs, the argument from
insufficient time falls away greatly in value.

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

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

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

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

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

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

{23}

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

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

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

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

From this resemblance in several countries of the half-domesticated dogs to
the wild species still living there,--from the facility with which they can
often be crossed together,--from even half-tamed animals being so much
valued by savages,--and from the other circumstances previously remarked on
which favour their domestication, it is highly probable that the domestic
dogs of the world have descended from two good species of wolf (viz. _C.
lupus_ and _C. latrans_), and from two or three other doubtful species of
wolves (namely, the European, Indian, and North African forms); from at
least one or two South American canine species; from several races or
species of the jackal; and perhaps from one or more extinct species. Those
authors who attribute great influence to the action of climate by itself
may thus account for the resemblance of the domesticated dogs and native
animals in the same countries; but I know of no facts supporting the belief
in so powerful an action of climate.

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

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

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

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

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

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

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

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

       *       *       *       *       *

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

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

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

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

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

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

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

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

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

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

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

DOMESTIC CATS.

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

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

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

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

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

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

       *       *       *       *       *


{49}

CHAPTER II.

HORSES AND ASSES.

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

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

The history of the Horse is lost in antiquity. Remains of this animal in a
domesticated condition have been found in the Swiss lake-dwellings,
belonging to the latter part of the Stone period.[100] At the present time
the number of breeds is great, as may be seen by consulting any treatise on
the Horse.[101] Looking only to the native ponies of Great Britain, those
of the Shetland Isles, Wales, the New Forest, and Devonshire are
distinguishable; and so it is with each separate island in the great Malay
archipelago.[102] Some of the breeds present great differences in size,
shape of ears, length of mane, proportions of the body, form of the withers
and hind quarters, and especially in the head. Compare the race-horse,
dray-horse, and a Shetland pony in size, configuration, and disposition;
and see how much greater the difference is than between the six or seven
other living species of the genus Equus.

{50}

Of individual variations not known to characterise particular breeds, and
not great or injurious enough to be called monstrosities, I have not
collected many cases. Mr. G. Brown, of the Cirencester Agricultural
College, who has particularly attended to the dentition of our domestic
animals, writes to me that he has "several times noticed eight permanent
incisors instead of six in the jaw." Male horses alone properly have
canines, but they are occasionally found in the mare, though of small
size.[103] The number of ribs is properly eighteen, but Youatt[104] asserts
that not unfrequently there are nineteen on each side, the additional one
being always the posterior rib. I have seen several notices of variations
in the bones of the leg; thus Mr. Price[105] speaks of an additional bone
in the hock, and of certain abnormal appearances between the tibia and
astragalus, as quite common in Irish horses, and not due to disease. Horses
have often been observed, according to M. Gaudry,[106] to possess a
trapezium and a rudiment of a fifth metacarpal bone, so that "one sees
appearing by monstrosity, in the foot of the horse, structures which
normally exist in the foot of the Hipparion,"--an allied and extinct
animal. In various countries horn-like projections have been observed on
the frontal bones of the horse: in one case described by Mr. Percival they
arose about two inches above the orbital processes, and were "very like
those in a calf from five to six months old," being from half to
three-quarters of an inch in length.[107] Azara has described two cases in
South America in which the projections were between three and four inches
in length: other instances have occurred in Spain.

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

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

With respect to the causes of the modifications which horses have
undergone, the conditions of life seem to produce a considerable direct
effect. Mr. D. Forbes, who has had excellent opportunities of comparing the
horses of Spain with those of South America, informs me that the horses of
Chile, which have lived under nearly the same conditions as their
progenitors in Andalusia, remain unaltered, whilst the Pampas horses and
the Puno ponies are considerably modified. There can be no doubt that
horses become greatly reduced in size and altered in appearance by living
on mountains and islands; and this apparently is due to want of nutritious
or varied food. Every one knows how small and rugged the ponies are on the
Northern islands and on the mountains of Europe. Corsica and Sardinia have
their native ponies; and there were,[116] or still are, on some islands on
the coast of Virginia, ponies like those of the Shetland Islands, which are
believed to have originated through exposure to unfavourable conditions.
The Puno ponies, which inhabit the lofty regions of the Cordillera, are, as
I hear from Mr. D. Forbes, strange little creatures, very unlike their
Spanish progenitors. Further south, in the Falkland Islands, the offspring
of the horses imported in 1764 have already so much deteriorated in
size[117] and strength that they are unfitted for catching wild cattle with
the lasso; so that fresh horses have to be brought for this purpose from La
Plata at a great expense. The reduced size of the horses bred on both
southern and northern islands, and on several mountain-chains, can hardly
have been caused by the cold, as a similar reduction has occurred on the
Virginian and Mediterranean islands. The horse can withstand intense cold,
for wild troops live on the plains of Siberia under lat. 56°,[118] and
aboriginally the horse must {53} have inhabited countries annually covered
with snow, for he long retains the instinct of scraping it away to get at
the herbage beneath. The wild tarpans in the East have this instinct; and,
as I am informed by Admiral Sulivan, this is likewise the case with the
horses which have run wild on the Falkland Islands; now this is the more
remarkable as the progenitors of these horses could not have followed this
instinct during many generations in La Plata: the wild cattle of the
Falklands never scrape away the snow, and perish when the ground is long
covered. In the northern parts of America the horses, descended from those
introduced by the Spanish conquerors of Mexico, have the same habit, as
have the native bisons, but not so the cattle introduced from Europe.[119]

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

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

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

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

[Illustration: Fig. 1.--Dun Devonshire Pony, with shoulder, spinal, and leg
stripes.]

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

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

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

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

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

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

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

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

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

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

{62}

THE ASS.

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

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

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

       *       *       *       *       *


{65}

CHAPTER III.

PIGS--CATTLE--SHEEP--GOATS.

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

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

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

    GOATS.--REMARKABLE VARIATIONS OF.

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

{66}

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

  ----------------+--------+---------+---------+-------------+-----------
                  | English|         |         |             | French
                  |  Long- |         |         |             | Domestic
                  | legged | African | Chinese | Wild Boar,  | Boar, from
                  |  Male. | Female. | Male.   | from Cuvier.| Cuvier.
  ----------------+--------+---------+---------+-------------+-----------
  Dorsal          |        |         |         |             |
  vertebræ        |  15    |    13   |   15    |     14      |    14
                  |        |         |         |             |
  Lumbar          |   6    |     6   |    4    |      5      |     5
                  +--------+---------+---------+-------------+-----------
  Dorsal and      |        |         |         |             |
  lumbar together |  21    |    19   |   19    |     19      |    19
                  |        |         |         |             |
  Sacral          |   5    |     5   |    4    |      4      |     4
                  +--------+---------+---------+-------------+-----------
  Total number    |        |         |         |             |
  of vertebræ     |  26    |    24   |   23    |     23      |    23
  ----------------+--------+---------+---------+-------------+-----------

{75}

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

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

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

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

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

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

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

CATTLE.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

SHEEP.

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

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

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

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

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

  Merinos                            150.3 days.
  Southdowns                         144.2  "
  Half-bred Merinos and Southdowns   146.3  "
  ¾ blood of Southdown             145.5  "
  7/8         "      "               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
foetal development should have been shortened. It is of course possible
that the difference in these two breeds may be due to their descent from
distinct parent-species; but as the early maturity of the Southdowns has
long been carefully attended to by breeders, the difference is more
probably the result of such attention. Lastly, the fecundity of the several
breeds differs much; some generally producing twins or even triplets at a
birth, of which fact the curious Shangai sheep (with their truncated and
rudimentary {98} ears, and great Roman noses), lately exhibited in the
Zoological Gardens, offer a remarkable instance.

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

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

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

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

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

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

GOATS.

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

       *       *       *       *       *


{103}

CHAPTER IV.

DOMESTIC RABBITS.

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

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

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

There are many breeds which transmit their characters more or less truly.
Every one has seen the enormous lop-eared rabbits exhibited at our shows;
various allied sub-breeds are reared on the Continent, such as the
so-called Andalusian, which is said to have a large head with a round
forehead, and to attain a greater size than any other kind; another large
Paris breed is named the Rouennais, and has a square head; the so-called
Patagonian rabbit has remarkably short ears and a large round head.
Although I have not seen all these breeds, I feel some doubt about there
being any marked difference in the {106} shape of their skulls.[254]
English lop-eared rabbits often weigh 8 lbs. or 10 lbs., and one has been
exhibited weighing 18 lbs.; whereas a full-sized wild rabbit weighs only
about 3¼ lbs. The head or skull in all the large lop-eared rabbits examined
by me is much longer relatively to its breadth than in the wild rabbit.
Many of them have loose transverse folds of skin or dewlaps beneath the
throat, which can be pulled out so as to reach nearly to the ends of the
jaws. Their ears are prodigiously developed, and hang down on each side of
their faces. A rabbit has been exhibited with its two ears, measured from
the tip of one to the tip of the other, 22 inches in length, and each ear
was 5-3/8 inches in breadth. In a common wild rabbit I found that the
length of the two ears, from tip to tip, was 7-5/8 inches, and the breadth
only 1-7/8 inch. The great weight of the body in the larger rabbits, and
the immense development of their ears, are the qualities which win prizes,
and have been carefully selected.

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

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

[Illustration: Fig. 5.--Half-lop Rabbit. (Copied from E. S. Delamer's
work.)]

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

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

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

{111}

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

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

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

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

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

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

_Osteological Characters._

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

    [Illustration: Fig. 6.--Skull of Wild Rabbit, of natural size.]

    [Illustration: Fig. 7.--Skull of large Lop-eared Rabbit, of natural
    size.]

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

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

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

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

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

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

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

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

    [Illustration: Fig. 10.--Occipital Foramen, of natural size, in--A.
    Wild Rabbit; B. Large Lop-eared Rabbit.]

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

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

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

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

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

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

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

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

    [Illustration: Fig. 12.--Atlas Vertebræ, of natural size; inferior
    surface viewed obliquely. Upper figure, Wild Rabbit. Lower figure,
    Hare-coloured, large, Lop-eared Rabbit. _a_, supra-median, atlantoid
    process; _b_, infra-median process.]

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

    [Illustration: Fig. 13.--Third Cervical Vertebra, of natural size,
    of--A. Wild Rabbit; B. Hare-coloured, large, Lop-eared Rabbit. _a, a_,
    inferior surface; _b, b_, anterior articular surfaces.]

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

{127}

  ------------------------------------------------+---------+-------------
                                                  | I.      |   II.
                                                  |         |
                                                  |         | Length
          Name of Breed.                          | Length  | of Body from
    WILD AND SEMI-WILD RABBITS.                   | of      | Incisors to
                                                  | Skull.  | Anus.
  ------------------------------------------------+---------+-------------
                                                  | inches. | inches.
   1. Wild rabbit, Kent                           |  3.15   |  17.4
   2.     "        Shetland Islands               |  3.15   |   ..
   3.     "        Ireland                        |  3.15   |   ..
   4. Domestic rabbit, run wild, Sandon           |  3.15   |  18.5
   5. Wild, common variety, small specimen, Kent  |  2.96   |  17.0
   6. Wild, fawn-coloured variety, Scotland       |  3.1    |   ..
   7. Silver-grey, small specimen, Thetford warren|  2.95   |  15.5
   8. Feral rabbit, Porto Santo                   |  2.83   |   ..
   9.       "            "                        |  2.85   |   ..
  10.       "            "                        |  2.95   |   ..
         Average of the three Porto Santo Rabbits |  2.88   |   ..
                                                  |         |
                     Domestic Rabbits             |         |
                                                  |         |
  11. Himalayan                                   |  3.5    |   20.5
  12. Moscow                                      |  3.25   |   17.0
  13. Angora                                      |  3.5    |   19.5
  14. Chinchilla                                  |  3.65   |   22.0
  15. Large lop-eared                             |  4.1    |   24.5
  16.   "         "                               |  4.1    |   25.0
  17.   "         "                               |  4.07   |    ..
  18.   "         "                               |  4.1    |   25.0
  19.   "         "                               |  4.3    |    ..
  20.   "         "                               |  4.25   |    ..
  21. Large hare-coloured                         |  3.86   |   24.0
  22. Average of above seven large lop-eared      |         |
          rabbits                                 |  4.11   |   24.62
  ------------------------------------------------+---------+-------------
  23. Hare (_L. timidus_) English specimen        |  3.61   |
  24.  "         "      German specimen           |  3.82   |
  ------------------------------------------------+---------+-------------

  ----------+-------------+------------+-----------------+-----------
            |   III.      |  IV.       |   V.            |  VI.
            |             |            |                 |
            |             |            | Capacity        |
            |             |            | calculated      | Difference
            |             | Capacity of| according to    | between
            |  Weight     | Skull      | Length of       | actual and
            |  of         | measured by| Skull relatively| calculated
            |  whole Body.| Small Shot.| to that         | capacities
            |             |            | of No. 1.       | of Skulls.
  ----------+-------------+------------+-----------------+-----------
            |  lbs. ozs.  |   grains.  |   grains.       | grains.
   1.       |    3   5    |    972     |     ..          |   ..
   2.       |     ..      |    979     |     ..          |   ..
   3.       |     ..      |    992     |     ..          |   ..
   4.       |     ..      |    977     |                 |
   5.       |    2  14    |    875     |    913          |   38
   6.       |     ..      |    918     |    950          |   32
   7.       |    2  11    |    938     |    910          |   28
   8.       |     ..      |    893     |    873          |   20
   9.       |     ..      |    756     |    879          |  123
  10.       |     ..      |    835     |    910          |   75
            |             |            |                 |
  (Average) |     ..      |    828     |    888          |   60
            |             |            |                 |
  11.       |     ..      |    963     |   1080          |  117
  12.       |    3   8    |    803     |   1002          |  199
  13.       |    3   1    |    697     |   1080          |  383
  14.       |     ..      |    995     |   1126          |  131
  15.       |    7   0    |   1065     |   1265          |  200
  16.       |    7  13    |   1153     |   1265          |  112
  17.       |     ..      |   1037     |   1255          |  218
  18.       |    7   4    |   1208     |   1265          |   57
  19.       |     ..      |   1232     |   1326          |   94
  20.       |     ..      |   1124     |   1311          |  187
  21.       |    6  14    |   1131     |   1191          |   60
  22.       |    7   4    |   1136     |   1268          |  132
  ----------+-------------+------------+-----------------+-----------
  23.       |    7   0    |   1315     |                 |
  24.       |    7   0    |   1455     |                 |
  ----------+-------------+------------+-----------------+-----------

  ----+-----------------------------------
      |  VII.
      |
      | Showing how much per cent.
      | the Brain, by calculation,
      | according to the length of the
      | Skull, is too light or too heavy,
      | relatively to the Brain of the
      | Wild Rabbit No. 1.
  ----+-----------------------------------
   1. |
   2. |
   3. | (  [2 per cent. too heavy in
      | (  comparison with No. 1.]
   4. |
   5. |  4 per cent. too light.
   6. |  3    "         "
   7. |  3    "      too heavy.
   8. |  2    "         "
   9. | 16    "       too light.
  10. |  9    "         "
      |
  Av. |  7    "         "
      |
  11. | 12    "         "
  12. | 24    "         "
  13. | 54    "         "
  14. | 13    "         "
  15. | 18    "         "
  16. |  9    "         "
  17. | 21    "         "
  18. |  4    "         "
  19. |  7    "         "
  20. | 16    "         "
  21. |  5    "         "
  22. | 11    "         "
  ----+-----------------------------------
  23. |
  24. |
  ----+-----------------------------------


{128}

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

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

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

       *       *       *       *       *


{131}

CHAPTER V.

DOMESTIC PIGEONS.

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

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

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

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

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

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

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

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

{136}

COLUMBA LIVIA or ROCK-PIGEON

  GROUP I.               GROUP II.

     1.     SUB-    2.       3.      4.
     .   GROUPS.    .
     .           Kali-Par
     .              .
     .              .
     .              ...Murassa
     .              .
     .              .
     .           Bussorah
     .              .
     .              ....................
     .              .     .      .     .
     .              . Bagadotten .     .
     .              .     .      .     .
     .              . Scanderoon .     .
     .              .     .      .     .
     .              .     .    Tronfo  .
     .              .     .            .
  German P.         .     .            .
     . Lille P.     .     .            .
     .     .     Dragon  Pigeon        .
     .     .        .    Cygne         .
  Dutch P. .        .     .            .
     .     .        .     .            .
     .......        .     .            .
     .              .     .            .
  English       English  Runt.        Barb.
  Pouter.       Carrier.

               GROUP III.                        GROUP IV.

    5.     6.      7.      8.      9.  SUB-      10.     11.
    .      .       .       .       .   GROUPS.   .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
    .      .    Persian    .       .             .  .  .  .  .  .  .
    .      .    Tumbler    .       .             .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
    .      .     Lotan     .       .             .  .  .  .  .  .  .
    .      .    Tumbler    .       .             .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
    .      .    Common     .       .             .  .  .  .  .  .  .
    .      .    Tumbler    .       .             .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
   Java    .       .       .       .             .  .  .  .  .  .  .
  Fantail  .       .       .       .             .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
    .    Turbit    .       .       .             .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
    .      .       .       .       .             .  .  .  .  .  .  .
    .      .       .       .       .    Trumpeter.  .  .  .  .  .  .
    .      .       .       .       .         Laugher.  .  .  .  .  .
    .      .       .       .       . English Frill-back.  .  .  .  .
    .      .       .       .       .                   Nun.  .  .  .
    .      .       .       .       .                     Spot.  .  .
    .      .       .       .       .                     Swallow.  .
    .      .       .       .       .                Dove-cot pigeon.
  Fantail. African Short-  Indian  Jacobin.
           Owl.    faced   Frill-
                   Tumbler. back.

{137}

GROUP I.

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

[Illustration: Fig. 18.--English Pouter.]

RACE I.--POUTER PIGEONS. (Kropf-tauben, German. Grosses-gorges, or boulans,
French.)

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

    _Sub-race I._--The improved English Pouter, when its crop is fully
    inflated, presents a truly astonishing appearance. The habit of
    slightly inflating the crop is common to all domestic pigeons, but is
    carried to an extreme in the Pouter. The crop does not differ, except
    in size, from that of other pigeons; but is less plainly separated by
    an oblique construction from the oesophagus. The diameter of the upper
    part of the oesophagus is immense, even close up to the head. The beak
    in one bird which I possessed was almost completely buried when the
    oesophagus 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 oesophagus assumes a spherical form, as if the pigeon had
    swallowed a large orange, which had stuck close under the beak. This
    inflated ball is represented as rising to a level with the crown of the
    head. The middle toe alone is feathered. A variety of this sub-race,
    called the claquant, is described by MM. Boitard and Corbié; it pouts
    but little, and is characterised {139} by the habit of violently
    hitting its wings together over its back,--a habit which the English
    Pouter has in a slight degree.

    _Sub-race IV. Common German Pouter._--I know this bird only from the
    figures and description given by the accurate Neumeister, one of the
    few writers on pigeons who, as I have found, may be always trusted.
    This sub-race seems considerably different. The upper part of the
    oesophagus 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._

[Illustration: Fig. 19.--English Carrier.]

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

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

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

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

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

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

RACE III.--RUNTS. (Scanderoons: Die Florentiner-Taube and Hinkel-Taube of
Neumeister: Pigeon Bagadais, Pigeon Romain.)

_Beak long, massive; body of great size._

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

    _Sub-race I. Scanderoon of English writers_ (Die Florentiner and
    Hinkel-Taube of Neumeister).--Birds of this sub-race, of which I kept
    one alive {143} and have since seen two others, differ from the
    Bagadotten of Neumeister only in not haying the beak nearly so much
    curved downwards, and in the naked skin round the eyes and over the
    nostrils being hardly at all wattled. Nevertheless I have felt myself
    compelled to place the Bagadotten in Race II., or that of the Carriers,
    and the present bird in Race III., or that of the Runts. The Scanderoon
    has a very short, narrow, and elevated tail; wings extremely short, so
    that the first primary feathers were not longer than those of a small
    tumbler pigeon! Neck long, much bowed; breast-bone prominent. Beak
    long, being 1.15 inch from tip to feathered base; vertically thick;
    slightly curved downwards. The skin over the nostrils swollen, not
    wattled; naked skin round the eyes, broad, slightly carunculated. Legs
    long; feet very large. Skin of neck bright red, often showing a naked
    medial line, with a naked red patch at the distant end of the radius of
    the wing. My bird, as measured from the base of the beak to the root of
    the tail, was fully 2 inches longer than the rock-pigeon; yet the tail
    itself was only 4 inches in length, whereas in the rock-pigeon, which
    is a much smaller bird, the tail is 4-5/8 inches in length.

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

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

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

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

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

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

RACE IV.--BARBS. (Indische-Taube: Pigeons Polonais.)

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

[Illustration: Fig. 20.--English Barb.]

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

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

GROUP III.

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

RACE V.--FANTAILS.

_Sub-race I. European Fantails_ (Pfauen-Taube; Trembleurs). _Tail expanded,
directed upwards, formed of many feathers; oil-gland aborted; body and beak
rather short_.

[Illustration: Fig. 21.--English Fantail.]

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

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

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

RACE VI.--TURBIT AND OWL. (Möven-Taube: Pigeons à cravate.)

_Feathers divergent along the front of the neck and breast; beak very
short, vertically rather thick; oesophagus somewhat enlarged._

[Illustration: 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 oesophagus, which causes a
    movement in the frill. {149} When the oesophagus 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 oesophagus; the Turbit inflates in a much less degree the
    oesophagus alone. The beak of the Turbit is very short, being .28 of an
    inch shorter than that of the rock-pigeon, proportionally with the size
    of their bodies; and in some owls brought by Mr. E. Vernon Harcourt
    from Tunis, it was even shorter. The beak is vertically thicker, and
    perhaps a little broader, in proportion to that of the rock-pigeon.

{150}

RACE VII.--TUMBLERS. (Tümmler, or Burzel-Tauben: Culbutants.)

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

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

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

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

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

[Illustration: Fig. 23.--Short-faced English Tumbler.]

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

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

RACE VIII--INDIAN FRILL-BACK.

_Beak very short; feathers reversed._

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

RACE IX.--JACOBIN. (Zopf or Perücken-Taube: Nonnains.)

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

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

GROUP IV.

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

RACE X.--TRUMPETER. (Trommel-Taube; Pigeon tambour; glougou.)

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

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

RACE XI.--_Scarcely differing in structure from the wild Columba livia._

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

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

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

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

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

       *       *       *       *       *

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

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

_Individual Variability; Variations of a remarkable nature._

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

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

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

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

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

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

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

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

[Illustration: Fig. 24.--Skulls of Pigeons, viewed laterally, of natural
size. A. Wild Rock-pigeon, _Columba livia_. B. Short-faced Tumbler. C.
English Carrier. D. Bagadotten Carrier.]

_Osteological Characters._

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

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

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

[Illustration: Fig. 26.--Skull of Runt, seen from above, of natural size,
showing the reflexed margin of the distal portion of the lower jaw.]

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

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

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

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

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

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

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


  +----------+-------------+--------------+-----------------+-------------+
  |          |  Rock       |Pouter,       |   Tumbler,      |    Bussorah |
  |          |  Pigeon.    |from Mr. Bult.|   Dutch Roller. |    Carrier. |
  +----------+-------------+--------------+-----------------+-------------+
  |Cervical  |             |              |                 |             |
  |Vertebræ  |    12       |     12       |       12        |       12    |
  |          |             |              |                 |  The 12th   |
  |          |             |              |                 |   bore a    |
  |          |             |              |                 |  small rib. |
  |Dorsal    |             |              |                 |             |
  |Vertebræ  |     8       |      8       |        8        |        8    |
  |          |             |              |                 |             |
  |Dorsal    |             |              |                 |             |
  |Ribs      |     8       |      8       |        7        |        7    |
  |          |The 6th pair |The 6th & 7th | The 6th & 7th   |The 6th & 7th|
  |          |    with     |  pair with   |  pair without   |pair without |
  |          | processes,  |  processes.  |  processes.     | processes.  |
  |          |the 7th pair |              |                 |             |
  |          | without a   |              |                 |             |
  |          |  process.   |              |                 |             |
  |          |             |              |                 |             |
  |Sacral    |             |              |                 |             |
  |Vertebræ  |    12       |      14      |        11       |       11    |
  |Caudal    |             |              |                 |             |
  |Vertebræ  |     7       |    8 or 9    |         7       |        7    |
  |          +-------------+--------------+-----------------+-------------+
  |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 {167} tumblers it is straighter, with
    the apex less elongated, than in the rock-pigeon: in the woodcut, fig.
    28, the scapulæ of the rock-pigeon (A), and of a short-faced tumbler
    (B), are given. The processes at the summit of the _coracoid_, which
    receive the extremities of the furcula, form a more perfect cavity in
    some tumblers than in the rock-pigeon: in pouters these processes are
    larger and differently shaped, and the exterior angle of the extremity
    of the coracoid, which is articulated to the sternum, is squarer.

    [Illustration: Fig. 29.--Furculæ, of natural size. A. Short-faced
    Tumbler B and C. Fantails. D. Pouter.]

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

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

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

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

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

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

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

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

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

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

       *       *       *       *       *

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

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

TABLE I.

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

  +-----------------------------------------+--------+-------------------+
  |                                         |        |   Difference      |
  |                                         |        |   between         |
  |                                         |        |   actual and      |
  |                                         |        |   calculated      |
  |                                         |        |   length of       |
  |          Name of Breed.                 | Actual |   feet, in        |
  |                                         | length |   proportion to   |
  |                                         | of     |   length of       |
  |                                         | Feet   |   feet and size   |
  |                                         |        |   of body in the  |
  |                                         |        |   Rock-pigeon     |
  |                                         |        +-------------------+
  | Wild rock-pigeon (mean measurement)     | 2.02   |Too short|Too long |
  |                                         |        |  by     |   by    |
  +-----------------------------------------+--------+---------+---------+
  | Short-faced Tumbler, bald-head          | 1.57   |  0.11   |  ..     |
  |       "        "     almond             | 1.60   |  0.16   |  ..     |
  | Tumbler, red magpie                     | 1.75   |  0.19   |  ..     |
  |    "     red common (by standard        |        |         |         |
  |                      to end of tail)    | 1.85   |  0.07   |  ..     |
  |    "     common bald-head               | 1.85   |  0.18   |  ..     |
  |    "     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 end of tail)    | 1.85   |  0.15   |  ..     |
  |    "           "          "             | 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     |
  +-----------------------------------------+--------+---------+---------+

{173}

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        |
  |          Name of Breed.              |   Actual |    feet, in         |
  |                                      |   length |    proportion to    |
  |                                      |   of     |    length of        |
  |                                      |   Feet   |    feet and size    |
  |                                      |          |    of body in the   |
  |                                      |          |    Rock-pigeon      |
  |                                      |          +----------+----------+
  | Wild rock-pigeon (mean measurement)  |    2.02  | Too short| Too long |
  |                                      |          |   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 cygne             |    2.85  |   ..     |    0.29  |
  | 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 {174}
    greyhound with the head of a bulldog, would be a monstrous production.
    As fancy pigeons are generally kept in small aviaries, and are
    abundantly supplied with food, they must walk about much less than the
    wild rock-pigeon; and it may be admitted as highly probable that the
    reduction in the size of the feet in the twenty-two birds in the first
    table has been caused by disuse,[312] and that this reduction has acted
    by correlation on the beaks of the great majority of the birds in Table
    I. When, on the other hand, the beak has been much elongated by the
    continued selection of successive slight increments of length, the feet
    by correlation have likewise become much elongated in comparison with
    those of the wild rock-pigeon, notwithstanding their lessened use.

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

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

_Length of Sternum._

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

{175}

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

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

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

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

We thus clearly see that the sternum, scapulæ, and furcula are all reduced
in proportional length; but when we turn to the wings we find what at first
appears a wholly different and unexpected result. I may here remark that I
have not picked out specimens, but have used every measurement made by me.
Taking the length from the base of beak to the end of the tail as the
standard of comparison, I find that, out of thirty-five birds of various
breeds, twenty-five have wings of greater, and ten have them of less
proportional length, than in the rock-pigeon. But from the frequently
correlated length of the tail and wing-feathers, it is better to take as
the standard {176} of comparison the length from the base of the beak to
the oil-gland; and by this standard, out of twenty-six of the same birds
which had been thus measured, twenty-one had wings too long, and only five
had them too short. In the twenty-one birds the wings exceeded in length
those of the rock-pigeon, on an average, by 1-1/3 inch; whilst in the five
birds they were less in length by only .8 of an inch. As I was much
surprised that the wings of closely confined birds should thus so
frequently have been increased in length, it occurred to me that it might
be solely due to the greater length of the wing-feathers; for this
certainly is the case with the jacobin, which has wings of unusual length.
As in almost every case I had measured the folded wings, I subtracted the
length of this terminal part from that of the expanded wings, and thus I
obtained, with a moderate degree of accuracy, the length of the wings from
the ends of the two radii, answering from wrist to wrist in our arms. The
wings, thus measured in the same twenty-five birds, now gave a widely
different result; for they were proportionally with those of the
rock-pigeon too short in seventeen birds, and in only eight too long. Of
these eight birds, five were long-beaked,[313] and this fact perhaps
indicates that there is some correlation between the length of the beak and
the length of the bones of the wings, in the same manner as with the feet
and tarsi. The shortening of the humerus and radius in the seventeen birds
may probably be attributed to disuse, as in the case of the scapulæ and
furcula to which the wing-bones are attached;--the lengthening of the
wing-feathers, and consequently the expansion of the wings from tip to tip,
being, on the other hand, as completely independent of use and disuse as is
the growth of the hair or wool on our long-haired dogs or long-woolled
sheep.

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

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

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

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

       *       *       *       *       *


{180}

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 {181} 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 {182}
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, {183} and roosts either on trees or rocks, according to the
    nature of the country. This species belongs to the genus Strictoenas 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 oenas_ 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 {184} 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.

    {185}

    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, {186} 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 {187} 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 {188} 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 {189} 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 {190} 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 {191} 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 {192}
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
{193} 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] {194} 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 {195}
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
{196} 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 {197} 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 {198} 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, {199} 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.

{200}

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 {201} 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 {202} 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. {203}

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 {204} 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 {205}
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 {206} 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 {207} 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 {208} 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 {209} 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." {210} 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 {211} 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 {212} 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 {213}
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 {214} 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 {215} 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 {216} 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 {217} 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 oesophagus 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 {218} 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 {219} 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 {220}
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 {221} 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, {222} 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 {223} 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 of