The Variation of Animals and Plants under Domestication

FOREWORD

Harriet Ritvo

Charles Darwin wrote On the Origin of
Species in a hurry. He had, it was true, been formulating his
ideas and arguments for several decades—since his
round-the-world Beagle voyage of 1831-1836. These ideas and
arguments had been slow to take definitive shape; Darwin had
nurtured and reworked them, amassing evidence for what he projected
to be a weighty magnum opus. Although he had shared his developing
evolutionary speculations with his closest professional colleagues,
Darwin was reluctant to publish them on several grounds. He was
aware that his theory of evolution by natural selection (or descent
with modification) was complex, that it rested on vast but not
incontrovertible evidence, and that the chain of his reasoning was
not uniformly strong. Further, his conclusions challenged not only
the scientific assumptions of many fellow specialists but also the
theological convictions of a much wider circle of fellow
citizens.

In 1859, Darwin did not feel quite ready to
expose his cherished theory to the harsh light of public scrutiny.
In the introduction to the Origin he confessed that although
his work on evolution by natural selection was “nearly finished,”
he would need “two or three more years to complete it.” The
Origin was, he suggested, merely a stopgap, a schematic
“abstract” of a much longer and more fully supported treatise yet
to come. He had been moved to preview his labors in this way, he
explained, because his health was “far from strong” and, perhaps
more importantly, because Alfred Russel Wallace, a younger
naturalist working in isolation in southeast Asia, had sent a paper
to the Linnean Society of London in which he “arrived at almost
exactly the same general conclusions that I have on the origin of
species.” If Darwin had not gone public with his theory at this
point, he would have risked losing credit for the work of many
years.

As its reception showed immediately and has
continued to show, the Origin benefited from the
succinctness imposed by circumstances. Darwin himself may have
appreciated this point; at any rate, he never produced the massive
treatise, although he repeatedly issued revised editions of the
Origin. But he did not abandon his intention to buttress his
initial schematic presentation with additional evidence. In the
course of the next two decades he published several full-length
elaborations of topics summarily discussed in the Origin: The
Variation of Animals and Plants under Domestication; The Descent of
Man, and Selection in Relation to Sex; and The Expression of
the Emotions in Man and Animals. In addition to fleshing out
the Origin, these subsequent studies bolstered its arguments
and responded to questions raised by critical readers, especially
pragmatic questions about the way that descent with modification
actually operated.

In The Variation of Animals and Plants under
Domestication, which appeared first in 1868 and in a revised
edition in 1875, Darwin developed a theme to which he had accorded
great rhetorical and evidentiary significance. He had begun the
Origin with a description of artificial selection as practiced
by farmers, stock breeders, and pet fanciers, thus using a
reassuringly homely example—one recognizable by the general
public as well as by members of the scientific community—to
introduce the most innovative component of his evolutionary theory.
In addition, domesticated animals and plants, because they were
numerous and available for constant observation, provided a readily
available body of evidence.

Reassuring as it was, the analogy between
natural and artificial selection was far from perfect. The point of
Darwin’s analogy was to make the idea of natural selection seem
plausible by characterizing it as a grander version of a well-known
process while emphasizing its efficiency and shaping power. He
noted, for example, that some of the prize birds bred by London
pigeon fanciers diverged so strikingly in size, plumage, beak
shape, flying technique, vocalizations. bone structure, and many
other attributes, that if they had been presented to an
ornithologist as wild specimens, they would unquestionably have
been considered to represent distinct species, perhaps even
distinct genera. Darwin argued that if the relatively brief and
constrained selective efforts of human breeders had produced such
impressive results, it was likely that the more protracted and
thorough-going efforts of nature would work still more
efficaciously.

But as Darwin acknowledged, there were some
fairly obvious reasons why the two processes might diverge. The
superior power of natural selection—“Man can act only on
external and visible characters: nature . . . can act on . . . the
whole machinery of life. Man selects only for his own good; Nature
only for that of the being which she tends” (Origin, chap.
5)—might constitute a difference of kind rather than of
degree, as might the much greater stretches of time available for
natural selection. Further, although the mechanism of the two
processes appeared superficially similar, their outcomes tended to
be rather different. Natural selection produced a constantly
increasing and diversifying variety of forms; it never reversed or
exactly repeated itself. Anyone familiar with artificial selection
would have realized that, although new breeds were constantly being
developed and although neither improved wheat nor improved cattle
showed any tendency to revert to the condition of their aboriginal
wild ancestors, the strains produced by human selection were
neither as prolific nor as durable as those produced by nature.
Indeed, the animals and plants celebrated as the noblest
achievements of the breeder’s art were especially liable to
delicacy and infertility. Highly bred strains, long isolated from
others of their species to preserve their genealogical purity, far
from serving as a springboard for further variation, often had to
be revivified with infusions of less-rarefied blood. Yet any
relaxation of reproductive boundaries threatened subsidence into
the common run of conspecifics.

Darwin firmly connected Variation to the
Origin by devoting its introduction to an overview of his
theory of evolution by natural selection. In particular, the two
volumes of Variation, cumbersomely organized and packed with
zoological and botanical detail, addressed some of the difficulties
inherent in the attractive but paradoxical analogy between natural
selection and artificial selection. For selection of any sort to
operate, diversity already had to exist. With wild populations
living under natural conditions, however, diversity was difficult
to discern. It was widely believed that a heightened propensity to
vary (at least in ways obvious to human observers) was one of the
few general characteristics that differentiated domestic animals as
a group from their wild relatives. This point was conventionally
illustrated with reference to coat color and design. American
bison, for example, were, on the whole, brown, and all Burchell’s
zebras shared similar black and white stripes. A single herd of
either Bos tauras or Equus caballus (domestic cattle
or horses), on the other hand, could display colors ranging from
white through yellow, red, and brown to black, as well as a variety
of spotted and blotched patterns.

In order to demonstrate that such populations
spontaneously produced sufficient variation to support artificial
selection, Darwin devoted most of the first volume of
Variation to a species-by-species survey of domesticated plants
and animals. He began with the dog, the breeds of which differed so
greatly in size, shape, disposition, talents, and every other
characteristic that Darwin attributed its exemplary plasticity to
its derivation from several different species of wild canines.
Domestic cats, on the other hand, differed relatively little from
one another, at least, their variation tended to be individual,
rather than consolidated into breeds. Darwin attributed this to the
minimal influence exerted by cat owners over the mating behavior of
their animals, so that, alone among fully domesticated animals,
cats could not be said to have undergone a genuine process of
artificial selection.

Farmyard ungulates, however, had all proved more
susceptible to human manipulation, whether through the gradual
enhancement of inherent tendencies, such as the relatively early
maturation that distinguished shorthorn cattle, or through the
preservation of spontaneously arising monstrosities, such as the
short, broad foreheads and protruding lower jaws of the niata
cattle of South America, the bulldogs of the bovine world. Among
animals, fancy pigeons, with their short generations, devoted
breeders, and lack of any pragmatic constraints on their
extravagant deformations, provided Darwin with his most abundant
material. He allotted less space to his survey of domesticated
plants, although, with the exception of trees, they tended to he
much shorter lived and more variable even than pigeons. For
example, as Darwin pointed out, a single long-cultivated
species—Brassica oleracea, the ordinary
cabbage—had given rise to strains as distinctive as Brussels
sprouts, cauliflower, broccoli, and kohl-rabi.

Darwin crammed in so much information of this
sort that, in order to confine Variation to two volumes of
manageable size, less crucial evidence was relegated to a smaller
typeface. And so compendious was his survey of domesticates that he
felt constrained to deny that it was intended to he an exhaustive
catalog. After all, many such catalogs, devoted merely to the
accumulation of species- or breed-specific data, existed already;
Darwin cited them generously in his footnotes. The material
included in Variation had been chosen to fulfill a more
focused argumentative purpose. Darwin’s theory of descent with
modification required something further than the simple
demonstration that abundant variation existed among domesticated
animals and plants. The accumulated experience of naturalists and
breeders offered no clear explanation of the causes of variation;
indeed, no consensus existed on this issue. Variation under
domestication was frequently attributed to accidental external
influences, especially climate and food. But environmentally
induced variation was not of much use to Darwin. Instead, he sought
evidence not only that the tendency to vary was inherent in
domesticated animals and plants but also that specific variations
were inherited.

As a result, Darwin’s wealth of detail in
Variation disproportionately featured strong—as well as
puzzling, problematic, or even questionable—versions of
inheritance, in addition to the unsurprising, if still not
completely understood, likelihood that children would resemble
their parents. For example, he devoted an entire chapter to what he
termed “atavism” or “reversion”—that is, the tendency for
offspring to manifest traits apparently derived from their
grandparents, collateral relations, or even remote ancestors,
rather than from their mothers or their fathers. The existence of
this tendency in the lineages of individuals, he argued,
incontrovertibly demonstrated the fact of heritability; and in an
extended or exaggerated version it also demonstrated evolutionary
relations between species. Thus, many breeds of domesticated
chickens revealed their ultimate ancestry by producing occasional
sports with the red and orange plumage of the original Callus
bankiva, or jungle fowl.

Like many other naturalists of his time, Darwin
was receptive to the idea of telegony, also known as “the influence
of the previous sire.” He retailed the famous story of Lord
Morton’s mare, a chestnut of seven-eighths Arabian blood, whose
first foal had been sired by a quagga (a now-extinct relative of
the zebra) her owner was attempting to domesticate. It was not
surprising that the young hybrid faintly echoed his father’s
stripes, but the fact that her next two foals, both sired by a
black Arabian horse, also seemed to resemble the quagga in this
regard, was more remarkable. Darwin pointed out that atavism
offered one possible explanation of this phenomenon—infant
horses and donkeys often showed evanescent striping, which might
indicate the pattern of their ancient shared progenitor—but
he was also drawn to the notion that the first male to impregnate a
female left some permanent, heritable trace of himself behind. He
offered analogous examples from the vegetable kingdom, where the
pollen of related varieties of apples, corn, or orchids, could not
only produce hybrid offspring but occasionally also physically
alter the reproductive tract of the female. Plants also, and more
regularly, demonstrated a kind of variability that could arise
independently of sexual reproduction, such as “bud variation,”
whereby what Darwin called a “monstrosity” might appear on a single
branch or flower and then be transmitted, sexually or asexually, to
future generations.

As he documented the profusion of variation
among domesticated animals and plants, and the tendency of
organisms to transmit these variations down the generations, Darwin
did more than demonstrate that there was ample grist for the mill
of natural selection. He also addressed the most serious weakness
in the argument of the Origin. Despite the incompleteness of
the fossil record, plenty of evidence suggested that evolution had
taken place; indeed the idea of evolution had been current in one
form or another for a century before 1859. Darwin’s explanation of
the way that natural selection should operate was also widely
persuasive. The competitive metaphors with which he characterized
it, especially the “struggle for life” prominently featured in the
Origin’s subtitle, fit well with Victorian understandings
about how things worked in the human arenas of industry, commerce,
and geopolitics. There was, however, a problem that troubled those
inclined to sympathize with Darwin’s reasoning as well as those
inclined to reject it. The efficacy of natural selection, like that
of artificial selection, depended on the inheritance of particular
traits. But before the modern understanding of genetics became
available, no satisfactory mechanism had been adduced to explain
this phenomenon. No consensus yet existed about the way that sexual
reproduction worked, so there was also disagreement about which
characteristics were inherited and which were the result of
environment, and what could he contributed by the male as opposed
to the female parent, let alone why offspring sometimes resembled a
grandparent or some more distant relative rather than their
parents. The special difficulty of accounting for the sudden
emergence of monstrosities, or even less dramatically novel traits,
led Darwin, in later editions of the Origin as well as in
Variation, to become increasingly receptive to the notion
that characteristics acquired by one generation might he inherited
by the next.

In the penultimate chapter of Variation,
Darwin attempted to strengthen the weak link in his chain of
argument by proposing a mechanism for inheritance. He called his
theory “pangenesis,” and he claimed that it explained not only
ordinary inheritance—the influence of parents on their
children—but also reversion, telegony, the regeneration of
amputated limbs in some kinds of animals, the inheritance of
acquired characteristics, and the relationship between sexual and
asexual modes of reproduction and inheritance. The operation of
pangenesis depended on the posited existence of unobservable units
that Darwin called “gemmules,” tiny granules that were thrown off
by individual cells and then circulated through the body. They had,
however, an affinity for each other, which led to their aggregation
in the reproductive organs or in parthenogenetic buds. They could
remain latent for years, until an organism reached a certain stage
of development, or for generations, until they encountered other
gemmules to which they bore some special relationship. In this way
a long-dormant greatgrandparental gemmule might suddenly manifest
itself in a child. Since gemmules could he altered by environmental
influences, they could convert acquired characteristics into the
stuff of heredity. And since they were vulnerable to error, they
could occasionally make mistakes, causing organs, such as limbs or
tails or even heads, to develop in inappropriate numbers or in the
wrong places.

It has doubtless been fortunate for Darwin’s
reputation that his theory of pangenesis is not as well remembered
as his theory of evolution by natural selection. As vague in detail
as it was ambitious and comprehensive in scope, it was unpersuasive
at the time and has since been proven completely wrong. But like
Variation as a whole, which similarly illustrated the
limitations of its author as well as his strengths, pangenesis does
not therefore lack interest or significance. Despite recent
excellent and well-appreciated studies of his entire life and
extended oeuvre (Janet Browne, Charles Darwin:
Voyaging [New York: Knopf, 1995] and Adrian Desmond and James
Moore, Darwin [London: Michael Joseph, 1991], Darwin is
known primarily as the author of the Origin, which is
unrepresentative in its economy of structure, argument, and
evidence, as well as on account of its historical notoriety. Its
enforced streamlining has helped to preserve the Origin’s
accessibility, but its relative paucity of examples was
particularly uncharacteristic of Darwin. Variation, with its
accumulation of evidence about everything from the webbing between
dogs’ toes to the weight of gooseberries, was much more typical; in
addition, it placed Darwin firmly—indeed,
irretrievably—within his time, rather than in an
achronological limbo reserved for intellectual heroes. As a
graduate student from the People’s Republic of China told me
several years ago, after having participated in a seminar that read
excerpts from Variation and The Expression of the
Emotions, if the leaders of his government knew that Darwin had
written such books, he would not be officially admired.

In science as in politics the victors tend to
write the history books. As a result, the record of the past is
edited, intentionally or unintentionally, so that it focuses mainly
on the precursors of contemporary orthodoxy. Such a focus may
accurately represent the genealogy of modem ideas, but it almost
inevitably misrepresents the historical experience of their
progenitors. Viewed without the benefit of hindsight, the
marketplace of Victorian ideas seemed much more competitive than it
does to us. Even the powerful, persuasive, and ultimately
triumphant theory of evolution by natural selection required not
only defense, but repeated buttressing and revision.
Variation showed Darwin hard at work on this rearguard action,
using the materials he had at hand—for the most part, homely
details about the domesticated animals and plants with which his
audience was most familiar. His information was gleaned from the
observations of fanciers, breeders, and amateur naturalists, as
well as from the treatises of those on the cutting edge of zoology
and botany. As hindsight narrows the historical spotlight, it
imposes its own sense of hierarchy on the preoccupations of the
past. But Darwin was interested in all of these topics, valued all
of these sources, and belonged, to a greater or lesser extent, to
all of these communities.

The author of Variation was a Victorian
country gentleman, a lover of dogs and horses, a breeder of pigeons
and peas. He was also, and equally, the author of On the Origin
of Species.

PREFACE TO THE SECOND EDITION

During the seven years which have elapsed since
the publication in 1868 of the first edition of this Work, I have
continued to attend to the same subjects, as far as lay in my
power; and I have thus accumulated a large body of additional
facts, chiefly through the kindness of many correspondents. Of
these facts I have been able here to use only those which seemed to
me the more important. I have omitted some statements, and
corrected some errors, the discovery of which I owe to my
reviewers. Many additional references have been given. The eleventh
chapter, and that on Pangenesis, are those which have been most
altered, parts having been remodelled; but I will give a list of
the more important alterations for the sake of those who may
possess the first edition of this book.

TABLE OF PRINCIPAL ADDITIONS AND CORRECTIONS IN SECOND EDITION

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
fulness.

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

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

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

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

In another work I shall discuss, if time and
health permit, 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 organisation is in some slight degree plastic; granting that
many animals and plants have varied greatly under domestication,
and that man by his power of selection has gone on accumulating
such variations until he has made strongly marked and firmly
inherited races; granting all this, how, it may be asked, have
species arisen in a state of nature? The differences between
natural varieties are slight; whereas the differences are
considerable between the species of the same genus, and great
between the species of distinct genera. How do these lesser
differences become augmented into the greater difference? How do
varieties, or as I have called them incipient species, become
converted into true and well-defined species? How has each new
species been adapted to the surrounding physical conditions, and to
the other forms of life on which it in any way depends? We see on
every side of us innumerable adaptations and contrivances, which
have justly excited the highest admiration of every observer. 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,—has been briefly treated in my ‘Origin of
Species.’ It was there shown that all organic beings, without
exception, tend to increase at so high a ratio, that no district,
no station, not even the whole surface of the land or the whole
ocean, would hold the progeny of a single pair after a certain
number of generations. The inevitable result is an ever-recurrent
Struggle for Existence. It has truly been said that all nature is
at war; the strongest ultimately prevail, the weakest fail; and we
well know that myriads of forms have disappeared from the face of
the earth. If then organic beings in a state of nature vary even in
a slight degree, owing to changes in the surrounding conditions, of
which we have abundant geological evidence, or from any other
cause; if, in the long course of ages, inheritable variations ever
arise in any way advantageous to any being under its excessively
complex and changing relations of life; and it would be a strange
fact if beneficial variations did never arise, seeing how many have
arisen which man has taken advantage of for his own profit or
pleasure; if then these contingencies ever occur, and I do not see
how the probability of their occurrence can be doubted, then the
severe and often-recurrent struggle for existence will determine
that those variations, however slight, which are favourable shall
be preserved or selected, and those which are unfavourable shall be
destroyed.

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

It has been shown from many facts that the
largest amount of life can be supported on each area, by great
diversification or divergence in the structure and constitution of
its inhabitants. We have, also, seen that the continued production
of new forms through natural selection, which implies that each new
variety has some advantage over others, 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 principal 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 end by being 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 comes into competition with 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
organisation 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 organisation
will in some cases be aided by habit, or by the use and disuse of
parts, and they will be governed by the direct action of the
surrounding physical conditions and by correlation of growth.

On the principles here briefly sketched out,
there is no innate or necessary tendency in each being to its own
advancement in the scale of organisation. We are almost compelled
to look at the specialisation 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, organisation 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 organised? Members of a high group might
even become, and this apparently has often occurred, fitted for
simpler conditions of life; and in this case natural selection
would tend to simplify or degrade the organisation, for complicated
mechanism for simple actions would be useless or even
disadvantageous.

The arguments opposed to the theory of Natural
Selection, have been discussed in my ‘Origin of Species,’ as far as
the size of that work permitted, under the following heads: the
difficulty in understanding how very simple organs have been
converted by small and graduated steps into highly perfect and
complex organs; the marvellous facts of Instinct; the whole
question of Hybridity; and, lastly, the absence in our known
geological formations of innumerable links connecting all allied
species. Although some of these difficulties are of great weight,
we shall see that many of them are explicable on the theory of
natural selection, and are otherwise inexplicable.

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

It was the consideration of such large groups of
facts as these 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 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 in their
geological constitution and climate. Still more surprising was the
fact that most of the inhabitants of each separate island in this
small archipelago were specifically different, though most closely
related to each other. The archipelago, with its innumerable
craters and bare streams of lava, appeared to be of recent origin;
and thus I fancied myself brought near to the very act of creation.
I often asked myself how these many peculiar animals and plants had
been produced: the simplest answer seemed to be that the
inhabitants of the several islands had descended from each other,
undergoing modification in the course of their descent; and that
all the inhabitants of the archipelago were 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 realised 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
their conditions of life were incomparably more like parts of South
America, than the different parts of that continent are 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 of South America
would alone explain the wide prevalence of American types
throughout that immense area.

To exhume with one’s own hands the bones of
extinct and gigantic quadrupeds brings the whole question of the
succession of species vividly before one’s mind; and I 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 the 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
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 are
often intermediate in character, like the words of a dead language
with respect to its several offshoots or living tongues. All these
facts seemed to me to point to descent with modification as the
means of production of new 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. These
complex affinities and the rules for classification, receive a
rational explanation on the theory of descent, combined with the
principle of natural selection, which entails divergence of
character and the extinction of intermediate forms. How
inexplicable is the similar pattern of the hand of a man, the foot
of a dog, the wing of a bat, the flipper of a seal, on the doctrine
of independent acts of creation! how simply explained on the
principle of the natural selection of successive slight variations
in the diverging descendants from a single progenitor! So it is
with certain parts or organs in the same individual animal or
plant, for instance, the jaws and legs of a crab, or the petals,
stamens, and pistils of a flower. During the many changes to which
in the course of time 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 useless condition is intelligible on the theory of
descent. It can be shown that modifications of structure are
generally inherited by the offspring at the same age at which each
successive variation appeared in the parents; it can further be
shown that variations do not commonly supervene at a very early
period of embryonic growth, and on these two principles we can
understand that most wonderful fact in the whole circuit of natural
history, namely, the close similarity of the embryos within the
same great class—for instance, those of mammals, birds,
reptiles, and fish.

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 Creation; they cannot be grouped together under one
point of view, but each has to be considered as an ultimate fact.
As the first origin of life on this earth, as well as the continued
life of each individual, is at present quite beyond the scope of
science, I do not wish to lay much stress on the greater simplicity
of the view of a few forms or of only one form having been
originally created, instead of innumerable miraculous creations
having been necessary at innumerable periods; though this more
simple view accords well with Maupertuis’s philosophical axiom of
“least action.”

In considering how far the theory of natural
selection may be extended, —that is, in determining from how
many progenitors the inhabitants of the world have
descended,—we may conclude that at least all the members of
the same class have descended from a single ancestor. A number of
organic beings are included in the same class, because they
present, independently of their habits of life, the same
fundamental type of structure, and because they graduate into each
other. Moreover, members of the same class can in most cases be
shown to be closely alike at an early embryonic age. These facts
can be explained on the belief of their descent from a common form;
therefore it may be safely admitted that all the members of the
same class are 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 would lead us one step
further, and to infer as probable that all living creatures are
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. 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,
etc.—on which we are confessedly ignorant; nor do we know how
ignorant we are. If we are much more ignorant than is generally
supposed, most of these difficulties wholly disappear. Let the
reader reflect on the difficulty of looking at whole classes of
facts from a new point of view. Let him observe how slowly, but
surely, the noble views of Lyell on the gradual changes now in
progress on the earth’s surface have been accepted as sufficient to
account for all that we see in its past history. The present action
of natural selection may seem more or less probable; but I believe
in the truth of the theory, because it collects, under one point of
view, and gives a rational explanation of, many apparently
independent classes of facts.^[4]

REFERENCES

[1]
To any one who has attentively read my ‘Origin of Species’
this Introduction will be superfluous. As I stated in that work
that I should soon publish the facts on which the conclusions
given in it were founded, I here beg permission to remark that
the great delay in publishing this first work has been caused by
continued ill-health.

[2]
M. Pouchet has recently (‘Plurality of Races,’ Eng.
Translat., 1864, p. 83, etc.) insisted that variation under
domestication throws no light on the natural modification of
species. I cannot perceive the force of his arguments, or, to
speak more accurately, of his assertions to this effect.

[3]
Léon Dufour in ‘Annales des Science. Nat.’ (3rd series,
Zoolog.), tom. v. p. 6.

[4]
In treating the several subjects included in the present and
my other works I have continually been led to ask for information
from many zoologists, botanists, geologists, breeders of animals,
and horticulturists, and I have invariably received from them the
most generous assistance. Without such aid I could have effected
little. I have repeatedly applied for information and specimens
to foreigners, and to British merchants and officers of the
Government residing in distant lands, and, with the rarest
exceptions, I have received prompt, open-handed, and valuable
assistance. I cannot express too strongly my obligations to the
many persons who have assisted me, and who, I am convinced, would
be equally willing to assist others in any scientific
investigation.

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^[1] does not throw much light on the question,
owing, on the one hand, to the close similarity of the skulls of extinct as
well as living wolves and jackals, and owing, on the other hand, to the
great dissimilarity of the skulls of the several breeds of the domestic
dogs. It seems, however, that remains have been found in the later tertiary
deposits more like those of a large dog than of a wolf, which favours the
belief of De Blainville that our dogs are the descendants of a single
extinct species. On the other hand, some authors go so far as to assert
that every chief domestic breed must have had its wild prototype. This
latter view is extremely improbable: it allows nothing for variation; it
passes over the almost monstrous character of some of the breeds; and it
almost necessarily assumes that a large number of species have become
extinct since man domesticated the dog; whereas we plainly see that wild
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.^[2] 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.^[3] At this latter period various breeds,
namely hounds, house-dogs, lapdogs, etc, 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^[4] 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 Egyptian
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^[5] 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^[6] states that the Arab boar-hound is “an
eccentric hieroglyphic animal, such as Cheops once hunted with, somewhat
resembling the rough Scotch deer-hound; their tails are curled tight round
on their backs, and their ears stick out at right angles.” With this most
ancient variety a pariah-like dog coexisted.

We thus see that, at a period between four and five thousand years ago,
various breeds, viz. pariah dogs, greyhounds, common hounds, mastiffs,
house-dogs, lapdogs, and turnspits, existed, more or less closely
resembling our present breeds. But there is not sufficient evidence that
any of these ancient dogs belonged to the same identical sub-varieties with
our present dogs.^[7] 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 embedded, 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.^[8] This ancient dog was succeeded in Denmark
during the Bronze period by a larger kind, presenting certain differences,
and this again during the Iron period, by a still larger kind. In
Switzerland, we hear from Prof. Rütimeyer,^[9] 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;^[10] 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.^[11]

The main argument in favour of the several breeds of the dog being the
descendants of distinct wild stocks, is their resemblance in various
countries to distinct species still existing there. It must, however, be
admitted that the comparison between the wild and domesticated animal has
been made but in few cases with sufficient exactness. Before entering on
details, it will be well to show that there is no a priori difficulty in
the belief that several canine species have been domesticated. 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^[12] 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 a 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^[13] how slowly the native birds of several
islands have acquired and inherited a salutary dread of man: at the
Galapagos Archipelago I pushed with the muzzle of my gun hawks from a
branch, and held out a pitcher of water for other birds to alight on and
drink. Quadrupeds and birds which have seldom been disturbed by man, dread
him no more than do our English birds, the cows, or horses grazing in the
fields.

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

We will now enter into details. The accurate and sagacious Richardson says,
“The resemblance between the Northern American wolves (Canis lupus,
var. occidentalis) and the domestic dogs of the Indians is so great
that the size and strength of the wolf seems to be the only difference. I
have more than once mistaken a band of wolves for the dogs of a party of
Indians; and the howl of the animals of both species is prolonged so
exactly in the same key that even the practised ear of the Indian fails at
times to discriminate them.” He adds that the more northern Esquimaux dogs
are not only extremely like the grey wolves of the Arctic circle in form
and colour, but also nearly equal them in size. Dr. Kane has often seen in
his teams of sledge-dogs the oblique eye (a character on which some
naturalists lay great stress), the drooping tail, and scared look of the
wolf. In disposition the Esquimaux dogs differ little from wolves, and,
according to Dr. Hayes, they are capable of no attachment to man, and are
so savage that when hungry they will attack even their masters. According
to Kane they readily become feral. Their affinity is so close with wolves
that they frequently cross with them, and the Indians take the whelps of
wolves “to improve the breed of their dogs.” The half-bred wolves sometimes
(Lamare-Picquot) cannot be tamed, “though this case is rare;” but they do
not become thoroughly well broken in till the second or third generation.
These facts show that there can be but little, if any, sterility between
the Esquimaux dog and the wolf, for otherwise they would not be used to
improve the breed. As Dr. Hayes says of these dogs, “reclaimed wolves they
doubtless are.”^[14]

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.^[15]

Turning to the southern parts of the new world, Columbus found two kinds of
dogs in the West Indies; and Fernandez^[16] 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,^[17] 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^[18]
states that they suffer from cold in the Cordillera. This naked dog is,
however quite distinct from that found preserved in the ancient Peruvian
burial-places, and described by Tschudi, under the name of Canis
ingæ, as withstanding cold well and as barking. It is not known whether
these two distinct kinds of dog are the descendants of native species, and
it might be argued that when man first migrated into America he brought
with him from the Asiatic continent dogs which had not learned to bark; but
this view does not seem probable, as the natives along the line of their
march from the north reclaimed, as we have seen, at least two N. American
species of Canidæ.

Turning to the Old World, some European dogs closely resemble the wolf;
thus the shepherd dog of the plains of Hungary is white or reddish-brown,
has a sharp nose, short, erect ears, shaggy coat, and bushy tail, and so
much resembles a wolf that Mr. p.t, 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.^[19] 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.^[20]

With respect to Jackals, Isidore Geoffroy Saint-Hilaire^[21] says that not one
constant difference can be pointed out between their structure and that of
the smaller races of dogs. They agree closely in habits: jackals, when
tamed and called by their master, wag their tails, lick his hands, crouch,
and throw themselves on their backs; they smell at the tails of other dogs,
and void their urine sideways; they roll on carrion or on animals which
they have killed; and, lastly, when in high spirits, they run round in
circles or in a figure of eight, with their tails between their legs.^[22] 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^[23]
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.^[24] 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.^[25] On the east coast of Africa, between
latitude 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^[26] says that the dogs of the Bosjemans
present a striking resemblance even in colour (excepting the black stripe
down the back) with the C. mesomelas of South Africa. Mr. E. Layard
informs me that he has seen a Caffre dog which closely resembled an
Esquimaux dog. In Australia the Dingo is both domesticated and wild; though
this animal may have been introduced aboriginally by man, yet it must be
considered as almost an endemic form, for its remains have been found in a
similar state of preservation and associated with extinct mammals, so that
its introduction must have been ancient.^[27]

From this resemblance of the half-domesticated dogs in several countries 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 are descended from two
well-defined species of wolf (viz. C. lupus and C. latrans),
and from two or three other doubtful species (namely, the European, Indian,
and North African wolves); from at least one or two South American canine
species; from several races or species of jackal; and perhaps from one or
more extinct species. Although it is possible or even probable that
domesticated dogs, introduced into any country and bred there for many
generations, might acquire some of the characters proper to the aboriginal
Canidæ of the country, we can hardly thus account for introduced dogs
having given rise to two breeds in the same country, resembling two of its
aboriginal species, as in the above-given cases of Guiana and of North
America.^[28]

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,^[29] and became as
sensible of 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 further see, in habits between the domestic dogs of the North
American Indians and the wolves of that country, or between the Eastern
pariah dogs and jackals, or between the dogs which have run wild in various
countries and the several natural species of the family. The habit of
barking, however, which is almost universal with domesticated dogs, forms
an exception, as it does not characterise a single natural species of the
family, though I am assured that the Canis latrans of North America
utters a noise which closely approaches a bark. But this habit is soon lost
by dogs when they become feral and is soon reacquired when they are again
domesticated. 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^[30] 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^[31] “made his voice sound as loudly as any
other dog of the same age and size.” According to Professor Nillson,^[32] 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.^[33] An interesting
account has been given by Mr. G. Clarke^[34] 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 singly or in packs, and burrow holes for their
young.^[35] 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.^[36] These feral dogs have not become
uniform in colour on Juan Fernandez, Juan de Nova, or La Plata.^[37] In
Cuba the feral dogs are described by Poeppig as nearly all mouse-coloured,
with short ears and light-blue eyes. In St. Domingo, Col. Ham. Smith says^[38] 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^[39]
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.” This dog, or another exactly the
same colour, ran at the Scottish National Club on the 21st of March, 1865;
and I hear from Mr. C. M. Browne, that “there was no reason either on the
sire or dam side for the appearance of this unusual colour.” 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^[40]
figures the magnificent black mastiff of Thibet with a tan-coloured stripe
over the eyes, feet, and chaps; and what is more singular, he figures the
Alco, or native domestic dog of Mexico, as black and white, with narrow
tan-coloured rings round the eyes; at the Exhibition of dogs in London, May
1863, a so-called forest dog from North-West Mexico was shown, which had
pale tan-coloured spots over the eyes. The occurrence of these tan-coloured
spots in dogs of such extremely different breeds, living in various parts
of the world, makes the fact highly remarkable.

We shall hereafter see, especially in the chapter on Pigeons, that coloured
marks are strongly inherited, and that they often aid us in discovering the
primitive forms of our domestic races. Hence, if any wild canine species
had distinctly exhibited the tan-coloured spots over the eyes, it might
have been argued that this was the parent-form of nearly all our domestic
races. But after looking at many coloured plates, and through the whole
collection of skins in the British Museum, I can find no species thus
marked. It is no doubt possible that some extinct species was thus
coloured. On the other hand, in looking at the various species, there seems
to be a tolerably plain correlation between tan-coloured legs and face; and
less frequently between black legs and a black face; and this general rule
of colouring explains to a certain extent the above-given cases of
correlation between the eye-spots and the colour of the feet. Moreover,
some jackals and foxes have a trace of a white ring round their eyes, as in
C. mesomelas, C. aureus, and (judging from Colonel H. 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 à rappeler la tache terminale de même
couleur, qui caractérise la plupart des Canidés sauvages.”^[41] This
rule, however, as I am assured by Mr. Jesse, does not invariably hold good.

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.^[42] Tessier, who has closely attended to
this subject, allows a difference of four days in the gestation of the dog.
The Rev. W. D. Fox has given me three carefully recorded cases of
retrievers, in which the bitch was put only once to the dog; and not
counting this day, but counting that of parturition, the periods were
fifty-nine, sixty-two, and sixty-seven days. The average period is
sixty-three days; but Bellingeri states that this applies only to 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;^[43] 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^[44] gave to a dog the same odour as that
from a jackal by feeding it on raw flesh.

The belief that our dogs are descended from wolves, jackals, South American
Canidæ, and other species, suggests a far more important difficulty. These
animals in their undomesticated state, judging from a widely-spread
analogy, would have been in some degree sterile if intercrossed; and such
sterility will be admitted as almost certain by all those who believe that
the lessened fertility of crossed forms is an infallible criterion of
specific distinctness. Anyhow these animals keep distinct in the countries
which they inhabit in common. On the other hand, all domestic dogs, which
are here supposed to be descended from several distinct species, are, as
far as is known, mutually fertile together. But, as Broca has well
remarked,^[45] 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^[46]
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^[47] 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 inclined to admit the truth of this hypothesis.

There is another and closely allied difficulty consequent on the doctrine
of the descent of our domestic dogs from several wild species, namely, that
they do not seem to be perfectly fertile with their supposed parents. But
the experiment has not been quite fairly tried; the Hungarian dog, for
instance, which in external appearance so closely resembles the European
wolf, ought to be crossed with this wolf: and the pariah dogs of India with
Indian wolves and jackals; and so in other cases. That the sterility is
very slight between certain dogs and wolves and other Canidæ is shown by
savages taking the trouble to cross them. Buffon got four successive
generations from the wolf and dog, and the mongrels were perfectly fertile
together.^[48] 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.^[49] 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.^[50] Some hounds from Central Africa,
brought home by Major Denham, never bred in the Town of London;^[51] 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; and this
circumstance would most certainly increase the tendency to sterility.
Several years ago I saw confined in the Zoological Gardens of London a
female hybrid from an English dog and jackal, which even in this the first
generation was so sterile that, as I was assured by her keeper, she did not
fully exhibit her proper periods; but this case was certainly exceptional,
as numerous instances have occurred of fertile hybrids from these two
animals. 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,^[52] with
its elongated muzzle; that of the larger dogs from the larger wolves, and
the smaller and slighter dogs from the 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^[53] a
large amount of variation. The intercrossing of the several aboriginal wild
stocks, and of the subsequently formed races, has probably increased the
total number of breeds, and, as we shall presently see, has greatly
modified some of them. But we cannot explain by crossing the origin of such
extreme forms as thoroughbred greyhounds, bloodhounds, bulldogs, Blenheim
spaniels, terriers, pugs, etc., 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 civilisation.

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^[54] 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 fossae; the position of the
occiput—all vary considerably.^[55] The difference in size between the
brains of dogs belonging to large and small breeds “is something
prodigious.” “Some dogs’ brains are high and rounded, while others are low,
long, and narrow in front.” In the latter, “the olfactory lobes are visible
for about half their extent, when the brain is seen from above, but they
are wholly concealed by the hemispheres in other breeds.”^[56] 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;^[57] and Professor Gervais says that there
are dogs “qui ont sept paires de dents supérieures et huit inférieures.” De
Blainville^[58] 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,^[59] the molar teeth stand
obliquely, whilst in long-muzzled races they are placed longitudinally,
with open spaces between them. The naked, so-called Egyptian or Turkish dog
is extremely deficient in its teeth,^[60] —sometimes having none except one
molar on each side; but this, though characteristic of the breed, must be
considered as a monstrosity. M. Girard,^[61] 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. On the other hand
small dogs are mature, and the females have arrived at the best age for
breeding, when one year old, whereas large dogs “are still in their
puppyhood at this time, and take fully twice as long to develop their
proportions.”^[62]

With respect to minor differences little need be said. Isidore Geoffroy has
shown^[63] 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.^[64] 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 vertebrae,
according to F. Cuvier, on whose authority the two last statements rest,
vary in number; and the tail in English cattle and some shepherd dogs is
almost absent. The mammae vary from seven to ten in number; Daubenton,
having examined twenty-one dogs, found eight with five mammae on each side;
eight with four on each side; and the others with an unequal number on the
two sides.^[65] 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^[66] 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^[67] “varies materially according to the
breed, as well as to the size of the animal.” Different breeds of dogs are
subject in different degrees to various diseases. They certainly become
adapted to different climates under which they have long existed. It is
notorious that most of our best European breeds deteriorate in India.^[68] The
Rev R. Everest^[69] believes that no one has succeeded in
keeping the Newfoundland dog long alive in India; so it is, according to
Lichtenstein,^[70] even at the Cape of Good Hope. The
Thibet mastiff degenerates on the plains of India, and can live only on the
mountains.^[71] Lloyd^[72] 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,^[73] admits “si l’on prenait sans contrôle
les alterations 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 mammae; the additional digit is generally
present with mastiffs, and some of the more important differences in the
skull and lower jaw are more or less characteristic of various breeds. But
we must not forget that the predominant power of selection has not been
applied in any of these cases; we have variability in important parts, but
the differences have not been fixed by selection. Man cares for the form
and fleetness of his greyhounds, for the size of his mastiffs, and formerly
for the strength of the jaw in his bulldogs, etc.; but he cares nothing
about the number of their molar teeth or mammae 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^[74] 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,^[75] who carefully attended to this subject,
states that “hounds are the most rapid in their decline;” “greyhounds and
pointers, also, rapidly decline.” But spaniels, after eight or nine
generations, and without a cross from Europe, are as good as their
ancestors. Dr. Falconer informs me that bulldogs, which have been known,
when first brought into the country, to pin down even an elephant by its
trunk, not only fall off after two or 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. So
again on the coast of Guinea, dogs, according to Bosman, “alter strangely;
their ears grow long and stiff like those of foxes, to which colour they
also incline, so that in three or four years, they degenerate into very
ugly creatures; and in three or four broods their barking turns into a
howl.”^[76] This remarkable tendency to rapid
deterioration in European dogs subjected to the climate of India and
Africa, may be largely accounted for by reversion to a primordial condition
which many animals exhibit, as we shall hereafter see, when their
constitutions are in any way disturbed.

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

It has already been remarked that dogs differ in the degree to which their
feet are webbed. In dogs of the Newfoundland breed, which are eminently
aquatic in their habits, the skin, according to Isidore Geoffroy,^[78]
extends to the third phalanges whilst in ordinary dogs it extends only to
the second. In two Newfoundland dogs which I examined, when the toes were
stretched apart and viewed on the under side, the skin extended in a nearly
straight line between the outer margins of the balls of the toes; whereas,
in two terriers of distinct sub-breeds, the skin viewed in the same manner
was deeply scooped out. In Canada there is a dog which is peculiar to the
country and common there, and this has “half-webbed feet and is fond of the
water.”^[79] English otter-hounds are said to have
webbed feet: a friend examined for me the feet of two, in comparison with
the feet of some harriers and bloodhounds; he found the skin variable in
extent in all, but more developed in the otter-hounds than in the others.^[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. The effects of
use from the frequent stretching apart of the toes will likewise aid in the
result. 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 soft; 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 necessary habits of life.

As changes in domestic breeds which take place so slowly are not to be
noticed at any one period, whether due to the selection of individual
variations or of differences resulting from crosses, are most important in
understanding the origin of our domestic productions, and likewise in
throwing indirect light on the changes effected under nature, I will give
in detail such cases as I have been able to collect. Lawrence,^[82] who
paid particular attention to the history of the foxhound, writing in 1829,
says that between eighty and ninety years before “an entirely new foxhound
was raised through the breeder’s art,” the ears of the old southern hound
being reduced, the bone and bulk lightened, the waist increased in length,
and the stature somewhat added to. It is believed that this was effected by
a cross with a 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 erroneously supposed to be 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. The bulldog is an English breed, and as I hear
from Mr. G. R. Jesse,^[85] seems to have originated from the
mastiff since the time of Shakspeare; but certainly existed in 1631, as
shown by Prestwick Eaton’s letters. 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 present names, Don, Ponto, Carlos, etc., 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 alter 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 desired owing to 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 Irish
wolf-dog, the old English bulldog, and several other breeds, such
as the alaunt, as I am informed by Mr. Jesse. 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 some difficulty, apparently from 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
individuals, modified in many cases by crosses with other breeds;
and as we shall hereafter see that the breeding of dogs was
attended to in ancient times, as it still is by savages, we may
conclude that we have in selection, even if only occasionally
practised, a potent means of modification.

DOMESTIC CATS.

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

Whether domestic cats have descended from
several distinct species, or have only been modified by occasional
crosses, their fertility, as far as is known, is unimpaired. The
large Angora or Persian cat is the most distinct in structure and
habits of all the domestic breeds; and is believed by Pallas, but
on no distinct evidence, to be descended from the F. manul
of middle Asia; and I am assured by Mr. Blyth that the Angora 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 one another.

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

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

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

REFERENCES

[1]
Owen ‘British Fossil Mammals,’ pp. 123 to
133. Pictet’s ‘Traité de Pal.,’ 1853, tom. i. p. 202. De
Blainville in his ‘Ostéographie, Canidæ,’ p. 142, has
largely discussed the whole subject, and concludes that the
extinct parent of all domesticated dogs came nearest to the wolf
in organisation, and to the jackal in habits. See also
Boyd Dawkins, ‘Cave Hunting,’ 1874, p. 131, etc., and his other
publications. Jeitteles has discussed in great detail the
character of the breeds of pre-historic dogs: ‘Die
vorgeschichtlichen Alterthümer der Stadt Olmütz,’ II.
Theil, 1872, p. 44 to end.

[2]
Pallas, I believe, originated this doctrine
in ‘Act. Acad. St. Petersburgh,’ 1780, Part ii. Ehrenberg has
advocated it, as may be seen in De Blainville’s
‘Ostéographie,’ p. 79. It has been carried to an extreme
extent by Col. Hamilton Smith in the ‘Naturalist Library,’ vols
ix and x. Mr. W. C. Martin adopts it in his excellent ‘History of
the Dog,’ 1845; as does Dr. Morton, as well as Nott and Gliddon,
in the United States. Prof. Low, in his ‘Domesticated Animals,’
1845, p. 666, comes to this same conclusion. No one has argued on
this side with more clearness and force than the late James
Wilson, of Edinburgh, in various papers read before the Highland
Agricultural and Wernerian Societies. Isidore Geoffroy
Saint-Hilaire (‘Hist. Nat. Gén.,’ 1860, tom. iii. p. 107),
though he believes that most dogs have descended from the jackal,
yet inclines to the belief that some are descended from the wolf.
Prof. Gervais (‘Hist. Nat. Mamm.’ 1855, tom. ii. p. 69, referring
to the view that all the domestic races are the modified
descendants of a single species, after a long discussion, says,
“Cette opinion est, suivant nous du moins, la moins
probable.”

[3]
Berjeau, ‘The Varieties of the Dog; in old Sculptures and Pictures,’ 1863. ‘Der
Hund,’ von Dr. F. L. Walther, Giessen, 1817, s. 48: this author seems carefully
to have studied all classical works on the subject. See also Volz,
‘Beiträge zur Kulturgeschichte,’ Leipzig, 1852, s. 115, ‘Youatt on the Dog,’
1845, p. 6. A very full history is given by De Blainville in his ‘Ostéographie,
Canidæ.’

[4]
I have seen drawings of this dog from the tomb of the son of Esar Haddon, and
clay models in the British Museum. Nott and Gliddon, in their ‘Types of
Mankind,’ 1854, p. 393, give a copy of these drawings. This dog has been called
a Thibetan mastiff, but Mr. H. A. Oldfield, who is familiar with the so-called
Thibet mastiff, and has examined the drawings in the British Museum, informs me
that he considers them different.

[5]
‘Proc. Zoolog. Soc.,’ July 12th, 1831.

[6]
‘Sporting in Algeria,’ p. 51.

[7]
Berjeau gives facsimiles of the Egyptian drawings. Mr. C. L. Martin in his
‘History of the Dog,’ 1845, copies several figures from the Egyptian monuments,
and speaks with much confidence with respect to their identity with still
living dogs. Messrs. Nott and Gliddon (‘Types of Mankind,’ 1854, p. 388) give
still more numerous figures. Mr. Gliddon asserts that a curl-tailed greyhound,
like that represented on the most ancient monuments, is common in Borneo; but
the Rajah, Sir J. Brooke, informs me that no such dog exists there.

[8]
These, and the following facts on the Danish remains, are taken from M.
Morlot’s most interesting memoir in ‘Soc. Vaudoise des Sc. Nat.’ tom. vi.,
1860, pp. 281, 299, 320.

[9]
‘Die Fauna der Pfahlbauten,’ 1861, s. 117, 162.

[10]
De Blainville ‘Ostéographie, Canidæ.’

[11]
Sir R. Schomburgk has given me information on this head. See also
‘Journal of R. Geographical Soc.’ vol. xiii. 1843, p. 65.

[12]
‘Domestication of Animals:’ Ethnological Soc., Dec. 22nd, 1863.

[13]
‘Journal of Researches,’ etc., 1845, p. 393. With respect to Canis
antarcticus, see p. 193. For the case of the antelope, see ‘Journal
Royal Geograph. Soc.,’ vol. xxiii. p. 94.

[14]
The authorities for the foregoing statements are as follow:—Richardson in
‘Fauna Boreali-Americana,’ 1829, pp. 64, 75; Dr. Kane ‘Arctic Explorations,’
1856, vol. i. pp. 398, 455; Dr. Hayes ‘Arctic Boat Journey,’ 1860, p. 167.
Franklin’s ‘Narrative,’ vol. i. p. 269, gives the case of three whelps of a
black wolf being carried away by the Indians. Parry, Richardson, and others,
give accounts of wolves and dogs naturally crossing in the eastern parts of
North America. Seeman in his ‘Voyage of H.M.S. Herald,’ 1853, vol. ii.
p. 26, says the wolf is often caught by the Esquimaux for the purpose of
crossing with their dogs, and thus adding to their size and strength. M.
Lamare-Picquot in ‘Bull. de la Soc. d’Acclimat,’ tom. vii., 1860, p. 148, gives
a good account of the half-bred Esquimaux dogs.

[15]
‘Fauna Boreali-Americana,’ 1829, pp. 73, 78, 80. Nott and Gliddon, ‘Types of
Mankind,’ p. 383. The naturalist and traveller Bartram is quoted by Hamilton
Smith, in ‘Naturalist Lib.,’ vol. x. p. 156. A Mexican domestic dog seems also
to resemble a wild dog of the same country; but this may be the prairie-wolf.
Another capable judge, Mr. J. K. Lord (‘The Naturalist in Vancouver Island,’
1866, vol. ii. p. 218), says that the Indian dog of the Spokans, near the Rocky
Mountains, “is beyond all question nothing more than a tamed Cayote or
prairie-wolf,” or Canis latrans.)

[16]
I quote this from Mr. R. Hill’s excellent account of the Alco or domestic dog
of Mexico, in Gosse’s ‘Naturalist’s Sojourn in Jamaica,’ 1851, p. 329.

[17]
‘Naturgeschichte der Säugethiere von
Paraguay,’ 1830, s. 151.

[18]
Quoted in Humboldt’s ‘Aspects of Nature’ (Eng. trans.), vol. i. p. 108.

[19]
p.t’s ‘Travels in Hungary and Transylvania,’ vol. i. p. 501. Jeitteles ‘Fauna
Hungariæ Superioris,’ 1862, s. 13. See Pliny ‘Hist. of the World’ (Eng.
trans.), 8th book, ch. xl., about the Gauls crossing their dogs. See
also Aristotle ‘Hist. Animal.’lib. viii. c. 28. For good evidence about
wolves and dogs naturally crossing near the Pyrenees, see M. Mauduyt ‘Du
Loup et de ses Races,’ Poitiers, 1851; also Pallas in ‘Acta Acad. St.
Petersburgh,’ 1780, part ii. p. 94.

[20]
I give this on excellent authority, namely Mr. Blyth (under the signature of
Zoophilus), in the ‘Indian Sporting Review,’ Oct. 1856, p. 134. Mr. Blyth
states that he was struck with the resemblance between a brush-tailed race of
pariah-dogs, north-west of Cawnpore, and the Indian wolf. He gives
corroborative evidence with respect to the dogs of the valley of the Nerbudda.

[21]
For numerous and interesting details on the resemblance of dogs and jackals
see Isid. Geoffroy St.-Hilaire ‘Hist. Nat. Gén.,’ 1860, tom. iii. p.
101. See also ‘Hist. Nat. des Mammifères,’ par Prof. Gervais, 1855,
tom. ii. p. 60.

[22]
Also Güldenstädt ‘Nov. Comment. Acad. Petrop.,’ tom. xx., pro anno 1775, p.
449. Also Salvin in ‘Land and Water,’ Oct. 1869.

[23]
Quoted by De Blainville in his ‘Ostéographie, Canidæ,’ pp. 79, 98.

[24]
See Pallas in ‘Act. Acad. St. Petersburgh,’ 1780, part ii. p. 91. For
Algeria, see Isid. Geoffroy St.-Hilaire ‘Hist. Nat. Gén.,’ tom. iii. p.
177. In both countries it is the male jackal which pairs with female domestic
dogs.

[25]
John Barbut’s ‘Description of the Coast of Guinea in 1746.’

[26]
‘Travels in South Africa,’ vol. ii. p. 272.

[27]
Selwyn, Geology of Victoria; ‘Journal of Geolog. Soc.,’ vol. xiv., 1858, p.
536, and vol. xvi., 1860, p. 148; and Prof. M’Coy, in ‘Annals and Mag. of Nat.
Hist.’ (3rd series) vol. ix., 1862, p. 147. The Dingo differs from the dogs of
the central Polynesian islands. Dieffenbach remarks (‘Travels,’ vol. ii. p. 45)
that the native New Zealand dog also differs from the Dingo.

[28]
These latter remarks afford, I think, a sufficient answer to some criticisms by
Mr. Wallace, on the multiple origin of dogs, given in Lyell’s ‘Principles of
Geology,’ 1872, vol. ii. p. 295.

[29]
‘Proceedings Zoolog. Soc.,’ 1833, p. 112. See also, on the taming of
the common wolf, L. Lloyd, ‘Scandinavian Adventures,’ 1854, vol. i. p. 460.
With respect to the jackal, see Prof. Gervais ‘Hist. Nat. Mamm.’ tom.
ii. p. 61. With respect to the aguara of Paraguay see Rengger’s work.

[30]
Roulin, in ‘Mém. présent. par divers Savans,’ tom. vi. p. 341.

[31]
Martin, ‘History of the Dog,’ p. 14.

[32]
Quoted by L. Lloyd in ‘Field Sports of North of Europe,’ vol. i. p. 387.

[33]
Quatrefages, ‘Soc. d’Acclimat.,’ May 11th, 1863, p. 7.

[34]
‘Annals and Mag. of Nat. Hist.’ vol. xv., 1845, p. 140.

[35]
Azara, ‘Voyages dans l’Amér. Mérid.’ tom. i. p. 381; his account is fully
confirmed by Rengger. Quatrefages gives an account of a bitch brought from
Jerusalem to France which burrowed a hole and littered in it. See
‘Discours, Exposition des Races Canines,’ 1865, p. 3.

[36]
With respect to wolves burrowing holes see Richardson, ‘Fauna
Boreali-Americana,’ p. 64; and Bechstein ‘Naturgeschichte Deutschlands,’ B. i.
s. 617.

[37]
See Poeppig, ‘Reise in Chile,’ B. i. s. 290; Mr. G. Clarke, as above;
and Rengger, s. 155.

[38]
Dogs, ‘Nat. Library,’ vol. x. p. 121; an endemic South American dog seems also
to have become feral in this island. See Gosse’s ‘Jamaica,’ p. 340.

[39]
Low ‘Domesticated Animals,’ p. 650.

[40]
‘The Naturalist Library,’ Dogs, vol. x. pp. 4, 19.

[41]
Quoted by Prof. Gervais, ‘Hist. Nat. Mamm.,’ tom. ii. p. 66.

[42]
J. Hunter shows that the long period of seventy-three days given by Buffon is
easily explained by the bitch having received the dog many times during a
period of sixteen days (‘Phil. Transact.,’ 1787, p. 353). Hunter found that the
gestation of a mongrel from wolf and dog (‘Phil. Transact.,’ 1789, p. 160)
apparently was sixty-three days, for she received the dog more than once. The
period of a mongrel dog and jackal was fifty-nine days. Fred. Cuvier found the
period of gestation of the wolf to be (‘Dict. Class. d’Hist. Nat.’ tom. iv. p.
8) two months and a few days, which agrees with the dog. Isid G. St.-Hilaire,
who has discussed the whole subject, and from whom I quote Bellingeri, states
(‘Hist. Nat. Gén.,’ tom. iii. p. 112) that in the Jardin des Plantes the period
of the jackal has been found to be from sixty to sixty-three days, exactly as
with the dog.

[43]
See Isid. Geoffroy St.-Hilaire ‘Hist. Nat. Gén.,’ tom. iii. p. 112, on
the odour of jackals. Col. Ham. Smith in ‘Nat. Lib.,’ vol. x. p. 289.

[44]
Quoted by Quatrefages in ‘Bull. Soc. d’Acclimat.,’ May 11th, 1863.

[45]
‘Journal de la Physiologie,’ tom. ii. p. 385.

[46]
See Mr. R. Hill’s excellent account of this breed in Gosse’s ‘Jamaica,’
p. 338; Rengger ‘Säugethiere von Paraguay,’ s. 153. With respect to Spitz dogs,
see Bechstein’s ‘Naturgesch. Deutschlands,’ 1801, B. i. s. 638. With
respect to Dr. Hodgkin’s statement made before Brit. Assoc. see ‘The
Zoologist,’ vol. iv. for 1845-46 p. 1097.

[47]
‘Acta Acad. St. Petersburgh,’ 1780, part ii. pp. 84, 100.

[48]
M. Broca has shown (‘Journal de Physiologie,’ tom. ii. p. 353) that Buffon’s
experiments have been often misrepresented. Broca has collected (pp. 390-395)
many facts on the fertility of crossed dogs, wolves, and jackals.

[49]
‘De la Longévité Humaine,’ par M. Flourens, 1855, p. 143. Mr. Blyth says
(‘Indian Sporting Review,’ vol. 2 p. 137) that he has seen in India several
hybrids from the pariah-dog and jackal; and between one of these hybrids and a
terrier. The experiments of Hunter on the jackal are well-known. See
also Isid. Geoffroy St.-Hilaire, ‘Hist. Nat. Gén.,’ tom. iii. p. 217, who
speaks of the hybrid offspring of the jackal as perfectly fertile for three
generations.

[50]
On authority of F. Cuvier quoted in Bronn’s ‘Geschichte der Natur,’ B ii. s.
164.

[51]
W. C. L. Martin ‘History of the Dog,’ 1845, p. 203. Mr. Philip P. King, after
ample opportunities of observation, informs me that the Dingo and European dogs
often cross in Australia.

[52]
Rüppel ‘Neue Wirbelthiere von Abyssinien,’ 1835-40 ‘Mammif.,’ s. 39 pl. xiv.
There is a specimen of this fine animal in the British Museum.

[53]
Even Pallas admits this; see ‘Act. Acad. St. Petersburgh,’ 1780, p. 93.

[54]
Quoted by I. Geoffroy, ‘Hist. Nat. Gén.,’ tom. iii. p. 453.

[55]
F. Cuvier in ‘Annales du Muséum,’ tom. xviii. p. 337; Godron ‘De l’Espèce,’
tom. i. p. 342; and Col. H. Smith in ‘Nat. Library,’ vol. ix. p. 101. See
also some observations on the degeneracy of the skull in certain breeds, by
Prof. Bianconi, ‘La Theorie Darwinienne,’ 1874, p. 279.

[56]
Dr. Burt Wilder, ‘American Assoc. Advancement of Science,’ 1873, pp. 236, 239.

[57]
Isid. Geoffroy Saint-Hilaire ‘Hist. des Anomalies,’ 1832, tom. i. p. 660,
Gervais ‘Hist. Nat. des Mammifères,’ tom. ii., 1855, p. 66. De Blainville
(‘Ostéographie, Canidæ,’ p. 137) has also seen an extra molar on both sides.

[58]
‘Ostéographie, Canidæ,’ p. 137.

[59]
Würzburger ‘Medecin. Zeitschrift,’ 1860, B. i. s. 265.

[60]
Mr. Yarrell in ‘Proc. Zoolog. Soc.,’ Oct. 8th, 1833. Mr. Waterhouse showed me a
skull of one of these dogs, which had only a single molar on each side and some
imperfect incisors.

[61]
Quoted in ‘The Veterinary,’ London, vol. viii. p. 415.

[62]
This is quoted from Stonehenge, a great authority, ‘The Dog,’ 1867, p. 187.

[63]
‘Hist. Nat. Général,’ tom. iii. p. 448.

[64]
W. Scrope ‘Art of Deer-Stalking,’ p. 354.

[65]
Quoted by Col. Ham. Smith in ‘Nat. Lib.,’ vol. x. p. 79.

[66]
De Blainville ‘Ostéographie, Canidæ,’ p. 134. F. Cuvier ‘Annales du Muséum,’
tom. xviii. p. 342. In regard to mastiffs, see Col. H. Smith ‘Nat. Lib.’
vol. x. p. 218. For the Thibet mastiff, see Mr. Hodgson in ‘Journal of
As. Soc. of Bengal,’ vol. i., 1832, p. 342.

[67]
‘The Dog,’ 1845, p. 186. With respect to diseases Youatt asserts (p. 167) that
the Italian greyhound is “strongly subject” to polypi in the matrix or vagina.
The spaniel and pug (p. 182) are most liable to bronchocele. The liability to
distemper (p. 232) is extremely different in different breeds. On the
distemper, see also Col. Hutchinson on ‘Dog Breaking,’ 1850, p. 279.

[68]
See Youatt on the Dog, p. 15; ‘The Veterinary,’ London, vol. xi. p. 235.

[69]
‘Journal of As. Soc. of Bengal,’ vol. iii. p. 19.

[70]
‘Travels,’ vol. ii. p. 15.

[71]
Hodgson in ‘Journal of As. Soc. of Bengal,’ vol. i. p. 342.

[72]
‘Field Sports of the North of Europe,’ vol. ii. p. 165.

[73]
‘Hist. Nat. des Mammif.,’ 1855, tom. ii. pp. 66, 67.

[74]
‘History of Quadrupeds,’ 1793, vol. i. p. 238.

[75]
‘Oriental Field Sports,’ quoted by Youatt, ‘The Dog,’ p. 15.

[76]
A. Murray gives this passage in his ‘Geographical Distribution of Mammals,’
4to, 1866, p. 8.

[77]
Quoted by Mr. Galton, ‘Domestication of Animals,’ p. 13.

[78]
‘Hist. Nat. Gén.,’ tom. iii. p. 450.

[79]
Mr. Greenhow on the Canadian Dog in Loudon’s ‘Mag. of Nat. Hist.,’ vol. vi.,
1833, p. 511.

[80]
See Mr. C. O. Groom-Napier on the webbing of the hind feet of
Otterhounds in ‘Land and Water,’ Oct. 13, 1866, p. 270.

[81]
‘Fauna Boreali-Americana,’ 1829, p. 62.

[82]
‘The Horse in all his Varieties,’ etc., 1829, pp. 230, 234.

[83]
‘The Dog,’ 1845, pp. 31, 35; with respect to King Charles’s spaniel, p. 45; for
the setter, p. 90.

[84]
In the ‘Encyclop. of Rural Sports,’ p. 557.

[85]
Author of ‘Researches into the History of the British Dog.’

[86]
See Col. Hamilton Smith on the antiquity of the Pointer, in ‘Nat. Lib.’
vol. x. p. 196.

[87]
The Newfoundland dog is believed to have originated from a cross between the
Esquimaux dog and a large French hound. See Dr. Hodgkin ‘British
Assoc.,’ 1844; Bechstein ‘Naturgesch. Deutschland,’ B. i. s. 574; ‘Nat. Lib.,’
vol. x. p. 132; also Mr. Jukes’ ‘Excursion in and about Newfoundland.’

[88]
De Blainville ‘Ostéographie, Felis,’ p. 65, on the character of F.
caligulata; pp. 85, 89, 90, 175, on the other mummied species. He quotes
Ehrenberg on F. maniculata being mummied.

[89]
Asiatic Soc. of Calcutta; Curator’s Report, Aug. 1856. The passage from Sir W.
Jardine is quoted from this Report. Mr. Blyth, who has especially attended to
the wild and domestic cats of India, has given in this Report a very
interesting discussion on their origin.

[90]
‘Fauna Hungariæ Sup.,’ 1862, s. 12.

[91]
Isid. Geoffroy Saint-Hilaire, ‘Hist. Nat. Gén.,’ tom. iii. p. 177.

[92]
‘Proc. Zoolog. Soc.,’ 1863, p. 184.

[93]
‘Säugethiere von Paraguay,’ 1830, s. 212.

[94]
‘Mem. présentés par divers Savans: Acad. Roy. des Sciences,’ tom. vi. p. 346.
Gomara first noticed this fact in 1554.

[95]
‘Narrative of Voyages,’ vol. ii. p. 180.

[96]
J. Crawfurd ‘Descript. Dict. of the Indian Islands,’ p. 255. The Madagascar cat
is said to have a twisted tail; see Desmarest in ‘Encyclop. Nat. Mamm.,’
1820, p. 233, for some of the other breeds.

[97]
Admiral Lutké’s Voyage, vol. iii. p. 308.

[98]
‘Zoology of the Voyage of the Beagle, Mammalia,’ p. 20. Dieffenbach ‘Travels in
New Zealand,’ vol. ii. p. 185. Ch. St. John ‘Wild Sports of the Highlands,’
1846, p. 40.

[99]
Quoted by Isid. Geoffroy ‘Hist. Nat. Gén.,’ tom. iii. p. 427.

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 Neolithic period.^[1] At the present time the number of breeds
is great, as may be seen by consulting any treatise on the Horse.^[2] 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,
amongst other instances, with each separate island in the great Malay
archipelago.^[3] 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 seven or eight other living species of the genus Equus.

Of individual variations not known to characterise particular breeds, and
not great or injurious enough to be called monstrosities, I have not
collected many cases. Mr. G. Brown, of the Cirencester Agricultural
College, who has particularly attended to the dentition of our domestic
animals, writes to me that he has “several times noticed eight permanent
incisors instead of six in the jaw.” Male horses only should have canines,
but they are occasionally found in the mare, though a small size.^[4] The
number of ribs on each side is properly eighteen, but Youatt^[5] asserts
that not unfrequently there are nineteen, the additional one being always
the posterior rib. It is a remarkable fact that the ancient Indian horse is
said in the Rig-Vêda to have only seventeen ribs; and M. Piétrement,^[6] who has
called attention to this subject, gives various reasons for placing full
trust in this statement, more especially as during former times the Hindoos
carefully counted the bones of animals. I have seen several notices of
variations in the bones of the leg; thus Mr. Price^[7] 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,^[8] 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.^[9] Azara has described two cases in South
America in which the projections were between three and four inches in
length: other instances have occurred in Spain.

That there has been much inherited variation in the horse cannot be
doubted, when we reflect on the number of the breeds existing throughout
the world or even within the same country, and when we know that they have
largely increased in number since the earliest known records.^[10] Even
in so fleeting a character as colour, Hofacker^[11] found that, out of
216 cases in which horses of the same colour were paired, only eleven pairs
produced foals of a quite different colour. As Professor Low^[12] 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
201,505 pounds sterling, and begot 497 winners; “Eclipse” begot 334
winners.

Whether the whole amount of difference between the various breeds has
arisen under domestication is doubtful. From the fertility of the most
distinct breeds^[13] 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.^[14] But as several species and varieties of
the horse existed^[15] during the later tertiary periods, and
as Rutimeyer found differences in the size and form of the skull in the
earliest known domesticated horses,^[16] we ought not to feel sure that all our
breeds are descended from a single species. The savages of North and South
America easily reclaim the feral horses, so that there is no improbability
in savages in various quarters of the world having domesticated more than
one native species or natural race. M. Sanson^[17] thinks that he has
proved that two distinct species have been domesticated, one in the East,
and one in North Africa; and that these differed in the number of their
lumbar vertebra and in various other parts; but M. Sanson seems to believe
that osteological characters are subject to very little variation, which is
certainly a mistake. At present no aboriginal or truly wild horse is
positively known to exist; for it is commonly believed that the wild horses
of the East are escaped domestic animals.^[18] If therefore our
domestic breeds are descended from several species or natural races, all
have become extinct in the wild state.

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 horses 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,^[19] 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^[20] 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°,^[21] and aboriginally the
horses must have inhabited countries annually covered with snow, for he
long retains the instinct of scraping it away to get at the herbage
beneath. The wild tarpans in the East have this instinct; and so it is, as
I am informed by Admiral Sulivan, with the horses recently and formerly
introduced into the Falkland Islands from La Plata, some of which have run
wild; this latter fact is remarkable, as the progenitors of these horses
could not have followed this instinct during many generations in La Plata.
On the other hand, 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.^[22]

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 circumstance may perhaps partly
account for the singular fact that to the eastward of the Bay of Bengal,^[23] 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.^[24]

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^[25] the case of a mare which produced
successively three foals without tails; so that a tailless race might have
been formed like the tailless races of dogs and cats. A Russian breed of
horses is said to have curled hair, and Azara^[26] 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 be derived from the
commingled blood of Arabs, Turks, and Barbs; but selection, which was
carried on during very early times in England,^[27] together with
training, have 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 pounds weight; and when both parents are of these countries a
discount of 36 pounds.^[28] 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 Bible we
hear of studs carefully kept for breeding, and of horses imported at high
prices from various countries.^[29] 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 been lost for a
long time. 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^[30] 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,”^[31] 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,^[32] in the same manner as is so conspicuous
with grey horses. This fact does not throw any clear light on the colouring
of the aboriginal horse, but is a case of analogous variation, for even
asses are sometimes dappled, and I have seen, in the British Museum, a
hybrid from the ass and zebra dappled on its hinder quarters. By the
expression analogous variation (and it is one that I shall often have
occasion to use) I mean a variation occurring in a species or variety which
resembles a normal character in another and distinct species or variety.
Analogous variations may arise, as will be explained in a future chapter,
from two or more forms with a similar constitution having been exposed to
similar conditions,—or from one of two forms having reacquired
through reversion a character inherited by the other form from their common
progenitor,—or from both forms having reverted to the same ancestral
character. We shall immediately see that horses occasionally exhibit a
tendency to become striped over a large part of their bodies; and as we
know that in the varieties of the domestic cat and in several feline
species stripes readily pass into spots and cloudy marks—even the
cubs of the uniformly-coloured lion being spotted with dark marks on a
lighter ground—we may suspect that the dappling of the horse, which
has been noticed by some authors with surprise, is a modification or
vestige of a tendency to become striped.

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

This tendency in the horse to become striped is in several respects an
interesting fact. 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.^[33] Occasionally horses are transversely
barred on the legs, chiefly on the under side; and more rarely they have a
distinct stripe on the shoulder, like that on the shoulder of the ass, or a
broad dark patch representing a stripe. Before entering on any details I
must premise that the term dun-coloured is vague, and includes three groups
of colours, viz., that between cream-colour and reddish-brown, which
graduates into light-bay or light-chestnut—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 (Figure 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 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 chestnut-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 chestnut 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 back
ill-defined stripe, running obliquely half-way down each shoulder; neither
had leg-stripes.

The most interesting case which I have met with occurred in a colt of my
own breeding. A bay mare (descended from a dark-brown Flemish mare by a
light grey Turcoman horse) was put to Hercules, a thoroughbred dark bay,
whose sire (Kingston) and dam were both bays. The colt ultimately turned
out brown; but when only a fortnight old it was a dirty bay, shaded with
mouse-grey, and in parts with a yellowish tint: it had only a trace of the
spinal stripe, with a few obscure transverse bars on the legs; but almost
the whole body was marked with very narrow dark stripes, in most parts so
obscure as to be visible only in certain lights, like the stripes which may
be seen on black kittens. These stripes were distinct on the hind-quarters,
where they diverged from the spine, and pointed a little forwards; many of
them as they diverged 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.^[34] My son estimated that
about a third of the ponies which he saw there 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 H. Smith^[35] alludes to dun-horses with the spinal
stripe in the Sierras of Spain; and the horses originally derived from
Spain, in some parts of South America, are now duns. Sir W. Elliot informs
me that he inspected a herd of 300 South American horses imported into
Madras, and many of these had transverse stripes on the legs and short
shoulder-stripes; the most strongly marked individual, of which a coloured
drawing was sent me, was a mouse-dun, with the shoulder-stripes slightly
forked.

In the North-Western parts of India striped horses of more than one breed
are apparently commoner than in any other part of the world; and I have
received information respecting them from several officers, especially from
Colonel Poole, Colonel Curtis, Major Campbell, Brigadier St. John, and
others. The Kattywar horses are often fifteen or sixteen hands in height,
and are well but lightly built. They are of all colours, but the several
kinds of duns prevail; and these are so generally striped, that a horse
without stripes is not considered pure. Colonel Poole believes that all the
duns have the spinal stripe, the leg-stripes are generally present, and he
thinks that about half the horses have the shoulder-stripe; this stripe is
sometimes double or treble on both shoulders. Colonel Poole has often seen
stripes on the cheeks and sides of the nose. He has seen stripes on the
grey and bay Kattywars when first foaled, but they soon faded away. I have
received other accounts of cream-coloured, bay, brown, and grey Kattywar
horses being striped. Eastward of India, the Shan (north of Burmah) ponies,
as I am informed by Mr. Blyth, have spinal, leg, and shoulder stripes. Sir
W. Elliot informs me that he saw two bay Pegu ponies with leg-stripes.
Burmese and Javanese ponies are frequently dun-coloured, and have the three
kinds of stripes, “in the same degree as in England.”^[36] Mr. Swinhoe informs
me that he examined two light-dun ponies of two Chinese breeds, viz., those
of Shanghai 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 dusty black), and rarely when almost white tinged
with yellow, grey, bay, and chestnut, 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.^[37]

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 dun; and it is generally
asserted that, when this is the case, the dun-colour and the stripes are
strongly inherited.^[38] One case, however, 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^[39] gives two instances of mouse-duns
(Mausrapp) being produced from two parents of different colours and neither
duns.

The stripes of all kinds are generally plainer in the foal than in the
adult horse, being commonly lost at the first shedding of the hair.^[40]
Colonel Poole believes that “the stripes in the Kattywar breed 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. 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 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. With natural species, the young often exhibit
characters which disappear at maturity.

The stripes are variable in colour, but are
always darker than the rest of the body. They do not by any means
always coexist on the different parts of the body: the legs may be
striped without any shoulder-stripe, or the converse case, which is
rarer, may occur; but I have never heard of either shoulder or
leg-stripes without the spinal stripe. The latter is by far the
commonest of all the stripes, as might have been expected, as it
characterises the other seven or eight species of the genus. It is
remarkable that so trifling a character as the shoulder-stripe
being double or triple should occur in such different breeds as
Welch and Devonshire ponies, the Shan pony, heavy cart-horses,
light South American horses, and the lanky Kattywar breed. Colonel
Hamilton Smith believes that one of his five supposed primitive
stocks was dun-coloured and striped; and that the stripes in all
the other breeds result from ancient crosses with this one
primitive dun; but it is extremely improbable that different breeds
living in such distant quarters of the world should all have been
crossed with any one aboriginally distinct stock. Nor have we any
reason to believe that the effects of a cross at a very remote
period would 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^[41] 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 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,” that is brown; not more than one in 2000 being black. In North
America the feral horses show a strong tendency to become roans of various
shades; but in certain parts, as I hear from Dr. Canfield, they are mostly
duns and striped.^[42]

In the following chapters on the Pigeon we shall
see that a blue bird is occasionally produced by pure breeds of
various colours and that when this occurs 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 horse certainly wild is known as a standard of
comparison; because the stripes when they appear are variable in
character; because there is far from sufficient evidence that the
crossing of distinct breeds produces stripes, and lastly, because
all the species of the genus Equus have the spinal stripe, and
several species 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 occasionally
revert.

THE ASS.

Four species of Asses, besides three zebras, have been described by
naturalists. There is now little doubt that our domesticated animal is
descended from the Equus tæniopus of Abyssinia.^[43] 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;^[44] 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 the South of France also there are
several breeds, and one of extraordinary size, some individuals being as
tall as full-sized horses. Although the ass in England is by no means
uniform in appearance, distinct breeds have not been formed. 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.”^[45]

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 more plainly than those of dun-coloured horses.
Thirteen or fourteen transverse stripes have been counted on both the fore
and hind legs. With the horse the occasional appearance of leg-stripes was
accounted for by reversion to a supposed parent-form, and in the case of
the ass we may confidently believe in this explanation, as E.
tæniopus is known to be barred, though only in a slight degree, and not
quite invariably. The stripes are believed to occur most frequently and to
be plainest on the legs of the domestic ass during early youth,^[46] as
likewise occurs 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 one 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;^[47] 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. If therefore grey and
reddish-brown asses had been steadily selected and bred from, the shoulder
stripe would probably have been lost almost as generally and completely 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.^[48] 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 in the above dun
Devonshire pony. I have seen three cases of the terminal portion abruptly
and angularly bent; and have seen and heard of four cases of a distinct
though slight forking of the stripe. In Syria, Dr. Hooker and his party
observed for me no less than five similar instances of the shoulder-stripe
plainly bifurcating over the fore leg. In the common mule it likewise
sometimes bifurcates. When I first noticed the forking and angular bending
of the shoulder-stripe, I had seen enough of the stripes in the various
equine species to feel convinced that even a character so unimportant as
this had a distinct meaning, and was thus led to attend to the subject. I
now find that in the E. 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 bent angularly backwards. The bifurcation
and angular bending of the stripes on the shoulders apparently are
connected with the nearly upright stripes on the sides of the body and neck
changing their direction and becoming transverse 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 downwards,—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 the genus. We shall
hereafter return to this subject, and discuss it more fully.

REFERENCES

[1]
Rütimeyer ‘Fauna der Pfahlbauten,’ 1861, s. 122.

[2]
See ‘Youatt on the Horse’: J. Lawrence on the Horse, 1829; W. C.
L. Martin, ‘History of the Horse,’ 1845: Col. H. Smith, in ‘Nat.
Library, Horses,’ 1841, vol. xii.: Prof. Veith, ‘Die naturgesch.
Haussäugethiere,’ 1856.

[3]
Crawfurd, ‘Descript. Dict. of Indian Islands,’ 1856, p. 153. “There are
many different breeds, every island having at least one peculiar to
it.” Thus in Sumatra there are at least two breeds; in Achin and
Batubara one; in Java several breeds; one in Bali, Lomboc, Sumbawa (one
of the best breeds), Tambora, Bima, Gunung-api, Celebes, Sumba, and
Philippines. Other breeds are specified by Zollinger in the ‘Journal of
the Indian Archipelago,’ vol. v, p. 343, etc.

[4]
‘The Horse,’ etc. by John Lawrence, 1829, p. 14.

[5]
‘The Veterinary,’ London, vol. v, p. 543.

[6]
‘Mémoire sur les chevaux à trente-quatre côtes,’ 1871.

[7]
Proc. Veterinary Assoc., in ‘The Veterinary,’ vol. xiii. p. 42.

[8]
‘Bulletin de la Soc. Géolog.,’ tom. xxii., 1866, p. 22.

[9]
Mr. Percival of the Enniskillen Dragoons, in ‘The Veterinary,’ vol. i.
p. 224: see Azara, ‘Des Quadrupèdes du Paraguay,’ tom. ii. p.
313. The French translator of Azara refers to other cases mentioned by
Huzard as having occurred in Spain.

[10]
Godron, ‘De l’Espèce’ tom. i. p. 378.

[11]
‘Ueber die Eigenschaften,’ etc., 1828, s. 10.

[12]
‘Domesticated Animals of the British Islands,’ pp. 527, 532. In all the
veterinary treatises and papers which I have read, the writers insist
in the strongest terms on the inheritance by the horse of all good and
bad tendencies and qualities. Perhaps the principle of inheritance is
not really stronger in the horse than in any other animal; but, from
its value, the tendency has been more carefully observed.

[13]
Andrew Knight crossed breeds so different in size as a dray-horse and
Norwegian pony: see A. Walker on ‘Intermarriage,’ 1838, p. 205.

[14]
‘Nat. Library, Horses,’ vol. xii. p. 208.

[15]
Gervais, ‘Hist. Nat. Mamm.,’ tom. ii. p. 143. Owen, ‘British Fossil Mammals,’
p. 383.

[16]
‘Kenntniss der fossilen Pferde,’ 1863, s. 131.

[17]
‘Comptes rendus,’ 1866, p. 485, and ‘Journal de l’Anat. et de la
Phys.,’ Mai 1868.

[18]
Mr. W. C. L. Martin, (‘The Horse,’ 1845, p. 34), in arguing against the
belief that the wild Eastern horses are merely feral, has remarked on
the improbability of man in ancient times having extirpated a species
in a region where it can now exist in numbers.

[19]
‘Transact. Maryland Academy,’ vol. i. part i. p. 28.

[20]
Mr. Mackinnon ‘The Falkland Islands,’ p. 25. The average height of the
Falkland horses is said to be 14 hands 2 inches. See also my
‘Journal of Researches.’

[21]
Pallas, ‘Act. Acad. St. Petersburgh,’ 1777, part ii. p. 265. With
respect to the tarpans scraping away the snow see Col. Hamilton
Smith in ‘Nat. Lib.,’ vol. xii. p. 165.

[22]
Franklin’s ‘Narrative,’ vol. i. p. 87; note by Sir J. Richardson.

[23]
Mr. J. H. Moor, ‘Notices of the Indian Archipelago;’ Singapore, 1837,
p. 189. A pony from Java was sent (‘Athenæum,’ 1842, p. 718) to the
Queen only 28 inches in height. For the Loo Choo Islands, see
Beechey’s ‘Voyage,’ 4th. edit., vol. i. p. 499.

[24]
J. Crawford, ‘History of the Horse;’ ‘Journal of Royal United Service
Institution,’ vol. iv.

[25]
‘Essays on Natural History,’ 2nd series, p. 161.

[26]
‘Quadrupédes du Paraguay,’ tom. ii. p. 333. Dr. Canfield informs me
that a breed with curly hair was formed by selection at Los Angeles in
North America.

[27]
See the evidence on this head in ‘Land and Water,’ May 2nd, 1868.

[28]
Prof. Low, ‘Domesticated Animals,’ p. 546. With respect to the writer
in India see ‘India Sporting Review,’ vol. ii. p. 181. As
Lawrence has remarked (‘The Horse,’ p. 9), “perhaps no instance has
ever occurred of a three-part bred horse (i.e. a horse, one of
whose grandparents was of impure blood) saving his distance in running
two miles with thoroughbred racers.” Some few instances are on record
of seven-eights racers having been successful.

[29]
Prof. Gervais (in his ‘Hist. Nat. Mamm.,’ tom. ii. p. 144) has
collected many facts on this head. For instance Solomon (Kings, B. i.
ch. x. v. 28) bought horses in Egypt at a high price.

[30]
‘The Field,’ July 13th, 1861, p. 42.

[31]
E. Vernon Harcourt, ‘Sporting in Algeria,’ p. 26.

[32]
I state this from my own observations made during several years on the
colours of horses. I have seen cream-coloured, light-dun and mouse-dun
horses dappled, which I mention because it has been stated (Martin,
‘History of the Horse,’ p. 134) that duns are never dappled. Martin (p.
205) refers to dappled asses. In the ‘Farrier’ (London, 1828, pp. 453,
455) there are some good remarks on the dappling of horses; and
likewise in Col. Hamilton Smith on ‘The Horse.’

[33]
Some details are given in ‘The Farrier,’ 1828, pp. 452, 455. One of the
smallest ponies I ever saw, of the colour of a mouse, had a conspicuous
spinal stripe. A small Indian chestnut pony had the same stripe, as had
a remarkably heavy chestnut cart-horse. Race-horses often have the
spinal stripe.

[34]
I have received information, through the kindness of the
Consul-General, Mr. J. R. Crowe, from Prof. Boeck, Rasck, and Esmarck,
on the colours of the Norwegian ponies. See also ‘The Field,’
1861, p. 431.

[35]
Col. Hamilton Smith, ‘Nat. Lib.,’ vol. xii. p. 275.

[36]
Mr. G. Clark, in ‘Annal and Mag. of Nat. History,’ 2nd series, vol. ii.
1848, p. 363. Mr. Wallace informs me that he saw in Java a dun and
clay-coloured horse with spinal and leg stripes.

[37]
See also on this point, ‘The Field,’ July 27th, 1861, p. 91.

[38]
‘The Field,’ 1861, pp. 431, 493, 545.

[39]
‘Ueber die Eigenschaften,’ etc., 1828, s. 13, 14.

[40]
Von Nathusius, ‘Vorträge über Viehzucht,’ 1872, 135.

[41]
‘Nat. Library,’ vol. xii. (1841), pp. 109, 156 to 163, 280, 281.
Cream-colour, passing into Isabella (i.e. the colour of the
dirty linen of Queen Isabella), seems to have been common in ancient
times. See also Pallas’s account of the wild horses of the East,
who speaks of dun and brown as the prevalent colours. In the Icelandic
sagas, which were committed to writing in the twelfth century,
dun-coloured horses with a black spinal stripe are mentioned;
see Dasent’s translation, vol. i. p. 169.

[42]
Azara, ‘Quadrupèdes du Paraguay,’ tom. ii. p. 307. In North America,
Catlin (vol. ii. p. 57) describes the wild horses, believed to have
descended from the Spanish horses of Mexico, as of all colours, black,
grey, roan, and roan pied with sorrel. F. Michaux (‘Travels in North
America,’ Eng. translat., p. 235) describes two wild horses from Mexico
as roan. In the Falkland Islands, where the horse has been feral only
between 60 and 70 years, I was told that roans and iron-greys were the
prevalent colours. These several facts show that horses do not soon
revert to any uniform colour.

[43]
Dr. Sclater, in ‘Proc. Zoolog. Soc.,’ 1862, p. 164. Dr. Hartmann says
(‘Annalen der Landw.’ B. xliv. p. 222) that this animal in its wild
state is not always striped across the legs.

[44]
W. C. Martin, ‘History of the Horse,’ 1845, p. 207.

[45]
Col. Sykes’ Cat. of Mammalia, ‘Proc. Zoolog. Soc.’ July 12th, 1831.
Williamson ‘Oriental Field Sports,’ vol. ii., quoted by Martin, p. 206.

[46]
Blyth, in ‘Charlesworth’s Mag. of Nat. Hist.,’ vol. iv., 1840, p. 83. I
have also been assured by a breeder that this is the case.

[47]
(One case is given by Martin, ‘The Horse,’ p. 205.

[48]
‘Journal As. Soc. of Bengal,’ vol. xxviii. 1860, p. 231. Martin on the
Horse, p. 205.

CHAPTER III.
PIGS—CATTLE—SHEEP—GOATS

PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND
INDICUS—TORFSCHWEIN—JAPAN PIGS—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.^[1] Nathusius
has shown that all the known breeds may be divided into 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 indicus, 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.

First for 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,^[2] 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. indicus 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;^[3] and in Hindostan an accurate observer,^[4] Sir
Walter Elliot, after describing the differences between wild Indian and
wild German boars, remarks that “the same differences are perceptible in
the domesticated individuals of the two countries.” We may therefore
conclude that the breeds of the Sus scrofa type are descended from,
or have been modified by crossing with, forms which may be ranked as
geographical races, but which, according to some naturalists, ought to be
ranked as distinct species.

Pigs of the Sus indicus type are best
known to Englishmen under the form of the Chinese breed. The skull
of S. indicus, 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. indicus 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.
indicus 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.
indicus; 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. indicus,
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 was found in the
buried city of Herculaneum.

Rütimeyer has made the remarkable discovery that there lived
contemporaneously in Switzerland, during the Neolithic period, two
domesticated forms, the S. scrofa, and the S. scrofa
palustris or Torfschwein. Rütimeyer perceived that the latter
approached the Eastern breeds, and, according to Nathusius, it certainly
belongs to the S. indicus 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.^[5] 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 the above conclusion; and
Rütimeyer himself seems now to feel some doubt. Other naturalists have also
argued strongly on the same side as Nathusius.^[6]

Several breeds, differing in the proportions of the body, in the length of
the ears, in the nature of the hair, in colour, etc., come under the S.
indicus 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^[7] 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.^[8] Hence
these pigs, as Nathusius has remarked,^[9] display in an eminent degree the
characters of a highly-cultivated race, and hence, no doubt, their 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. indicus 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. indicus, such as
the shortness of the lachrymal bones, etc., being common to several species
of the genus; for in crosses characters which are common to many species
apparently tend to be prepotent over those appertaining to only a few
species.

Fig. 2.—Head of Japan or Masked Pig.

The Japan pig (S. pliciceps of Gray), which was formerly exhibited
in the Zoological Gardens, has an extraordinary appearance from its short
head, broad forehead and nose, great fleshy ears, and deeply furrowed skin.
Figure 2 is copied from that given by Mr. Bartlett.^[10] Not only is the face
furrowed, but thick folds of skin, which are harder than the other parts,
almost like the plates on the Indian rhinoceros, hang about the shoulders
and rump. It is coloured black, with white feet, and breeds true. That it
has long been domesticated there can be little doubt; and this might have
been inferred even from the fact that its young are not longitudinally
striped; for this is a character common to all the species included within
the genus Sus and the allied genera whilst in their natural state.^[11] Dr.
Gray^[12] 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. indicus type. Hence Nathusius considers the
Japan pig as only a domesticated variety of S. indicus: if this
really be the case, it is a wonderful instance of the amount of
modification which can be effected under domestication.

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^[13] 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 of European and
Chinese pigs into these islands, 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.”^[14]

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

Fig. 3—Head of Wild Boar, and of “Golden
Days,” a pig of the Yorkshire Large Breed

The modification of the skull in the most highly cultivated races is
wonderful. To appreciate the amount of change, Nathusius’ work, with its
excellent figures, should be studied. The whole of the exterior 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 beyond 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-racen” the
proportion is as 1 to 9, and even recently as 1 to 11.^[15] The following
woodcut^[16] of the head of a wild boar and of a sow
from a photograph of the Yorkshire Large Breed, may aid in showing how
greatly the head in a highly cultivated race has been modified and
shortened.

Nathusius has well discussed the causes of the remarkable changes in the
skull and shape of the body which the highly cultivated races have
undergone. These modifications occur chiefly in the pure and crossed races
of the S. indicus type; but their commencement may be clearly
detected in the slightly improved breeds of the S. scrofa type.^[17]
Nathusius states positively (s. 99, 103), as the result of common
experience and of his experiments, that rich and abundant food, given
during youth, tends by some direct action to make the head broader and
shorter; and that poor food works a contrary result. He lays much stress on
the fact that all wild and semi-domesticated pigs, in ploughing up the
ground with their muzzles, have, whilst young, to exert the powerful
muscles fixed to the hinder part of the head. In highly cultivated races
this habit is no longer followed, and consequently the back of the skull
becomes modified in shape, entailing other changes in other parts. There
can hardly be a doubt that so great a change in habits would affect the
skull; but it seems rather doubtful how far this will account for the
greatly reduced length of the skull and for its concave front. It is well
known (Nathusius himself advancing many cases, s. 104) that there is a
strong tendency in many domestic animals—in bull- and pug-dogs, in
the niata cattle, in sheep, in Polish fowls, short-faced tumbler pigeons,
and in one variety of the carp—for the bones of the face to become
greatly shortened. In the case of the dog, as H. Müller has shown, this
seems caused by an abnormal state of the primordial cartilage. We may,
however, readily admit that abundant and rich food supplied during many
generations would give an inherited tendency to increased size of body, and
that, from disuse, the limbs would become finer and shorter.^[18] We
shall in a future chapter see also 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.^[19]

The nature of the food supplied during many generations has apparently
affected the length of the intestines; for, according to Cuvier,^[20] their
length to that of the body in the wild boar is as 9 to 1,—in the
common domestic boar as 13·5 to 1,—and in the Siam breed as 16
to 1. In this latter breed the greater length may be due either to descent
from a distinct species or to more ancient domestication. The number of
mammæ vary, as does the period of gestation. The latest authority says^[21] 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 the course of their 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,^[22] than in the case of common swine. 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,^[23] 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
this paper, cross-bred animals from the African and English races were
found by Lord Hill to be perfectly fertile.

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^[24] 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.

Old Irish Pig, with jaw-appendages.

Another curious anomaly is offered by the appendages, described by M.
Eudes-Deslongchamps as often characterizing the Normandy pigs. These
appendages are always attached to the same spot, to the corners of the jaw;
they are cylindrical, about three inches in length, covered with bristles,
and with a pencil of bristles rising out of a sinus on one side: they have
a cartilaginous centre, with two small longitudinal muscles they occur
either symmetrically on both sides of the face or on one side alone.
Richardson figures them on the gaunt old “Irish Greyhound pig;” and
Nathusius states that they occasionally appear in all the long eared races,
but are not strictly inherited, for they occur or fail in animals of the
same litter.^[25] 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 a somewhat complex, though apparently useless, structure may be
suddenly developed without the aid of selection.

It is a remarkable fact that the boars of all
domesticated breeds have much shorter tusks than wild boars. Many
facts show that with many 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,^[26] 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;”^[27] whether these latter
pigs belong to the same curly-haired race as 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.^[28]

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, namely, S.
scrofa and indicus, have not been distinguished. The young, as
we have just seen, reacquire their longitudinal stripes, and the boars
invariably reassume their tusks. They revert also in the general shape of
their bodies, and in the length of their legs and muzzles, to the state of
the wild animal, as might have been expected from the amount of exercise
which they are compelled to take in search of food. In Jamaica the feral
pigs do not acquire the full size of the European wild boar, “never
attaining a greater height than 20 inches at the shoulder.” In various
countries they reassume their original bristly covering, but in different
degrees, dependent on the climate; thus, according to Roulin, the
semi-feral pigs in the hot valleys of New Granada are very scantily
clothed; whereas, on the Paramos, at the height of 7000 to 8000 feet, they
acquire a thick covering of wool lying under the bristles, like that on the
truly wild pigs of France. These pigs on the Paramos are small and stunted.
The wild boar of India is said to have the bristles at the end of its tail
arranged like the plumes of an arrow, whilst the European boar has a simple
tuft; and it is a curious fact that many, but not all, of the feral pigs in
Jamaica, derived from a Spanish stock, have a plumed tail.^[29] 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.^[30] 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,^[31] the Berkshire breed of 1780 is quite
different from that of 1810; and, since this latter period, at least two
distinct forms have borne the same name.

CATTLE.

Domestic cattle are 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,^[32] than do the fossil and prehistoric
European species, namely, Bos primigenius and longifrons,
from each other. They differ, also, as Mr. Blyth,^[33] who has particularly
attended to this subject, remarks, in general configuration, in the shape
of their ears, in the point where the dewlap commences, in the typical
curvature of their horns, in their manner of carrying their heads when at
rest, in their ordinary variations of colour, especially in the frequent
presence of “nilgau-like markings on their feet,” and “in the one being
born with teeth protruding through the jaws, and the other not so.” They
have different habits, and their voice is entirely different. The humped
cattle in India “seldom seek shade, and never go into the water and there
stand knee-deep, like the cattle of Europe.” They have run wild in parts of
Oude and Rohilcund, and can maintain themselves in a region infested by
tigers. They have given rise to many races differing greatly in size, in
the presence of one or two humps, in length of horns, and other respects.
Mr. Blyth sums up emphatically that the humped and humpless cattle must be
considered as distinct species. When we consider the number of points in
external structure and habits, independently of 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;^[34] 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
enormous horns sometimes measuring above five feet from tip to tip:^[35] the
Podolian cattle also are remarkable from the height of their fore-quarters.
In the most recent work on Cattle,^[36] 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-civilised 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,^[37] and more especially from Rütimeyer’s
works and those of Boyd Dawkins. Two or three species or forms of Bos,
closely allied to still living domestic races, have been found in the more
recent tertiary deposits or amongst prehistoric remains in 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, etc., 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.^[38]

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.^[39]

Bos longifrons (or brachyceros) of Owen.—This very
distinct species was of small size, and had a short body with fine legs.
According to Boyd Dawkins^[40] it was introduced as a domesticated
animal into Britain at a very early period, and supplied food to the Roman
legionaries.^[41] Some remains have been found in Ireland
in certain crannoges, of which the dates are believed to be from 843-933
A.D.^[42] It was also the commonest form in a
domesticated condition in Switzerland during the earliest part of the
Neolithic period. Professor Owen^[43] 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, and, according to the high authority of Mr. Boyd Dawkins,
is identical with it, but in the opinion of some judges is distinct. Both
co-existed in Scania during the same late geological period,^[44] and
both have been found in the Irish crannoges.^[45] Nilsson believes that
his B. frontosus may be the parent of the mountain cattle of Norway,
which have a high protuberance on the skull between the base of the horns.
As Professor Owen and others believe that the Scotch Highland cattle are
descended from his B. longifrons, it is worth notice that a capable
judge^[46] has remarked that he saw no cattle in
Norway like the Highland breed, but that they more nearly resembled the
Devonshire breed.

On the whole we may conclude, more especially from the researches of Boyd
Dawkins, that European cattle are descended from two species; and there is
no improbability in this fact, for the genus Bos readily yields to
domestication. Besides these two species and the zebu, the yak, the gayal,
and the arni^[47] (not to mention the buffalo or genus
Bubalus) have been domesticated; making altogether six species of Bos. The
zebu and the two European species are now extinct in a wild state. Although
certain races of cattle were domesticated at a very ancient period in
Europe, it does not follow that they were first domesticated here. Those
who place much reliance on philology argue that they were imported from the
East.^[48] It is probable that they 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, 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.”^[49]

Bos primigenius and longifrons have been ranked by nearly all
palæontologists as distinct species; and it would not be reasonable to take
a different view simply because their domesticated descendants now
intercross with the utmost freedom. All the European breeds have so often
been crossed both intentionally and unintentionally, that, if any sterility
had 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^[50] 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 have been in all probability
in some degree sterile, become perfectly fertile after a long course of
domestication. In a future chapter we shall see that this doctrine throws
some light on the difficult subject of Hybridism.

I have alluded to the cattle in Chillingham Park, which, according to
Rütimeyer, have been very little changed from the Bos primigenius
type. This park is so ancient that it is referred to in a record of the
year 1220. The cattle in their instincts and habits are truly wild. They
are white, with the inside of the ears reddish-brown, eyes rimmed with
black, muzzles brown, hoofs black, and horns white tipped with black.
Within a period of thirty-three years about a dozen calves were born with
“brown and blue spots upon the cheeks or necks; but these, together with
any defective animals, were always destroyed.” According to Bewick, about
the year 1770 some calves appeared with black ears; but these were also
destroyed by the keeper, and black ears have not since reappeared. The wild
white cattle in the Duke of Hamilton’s park, where I have heard of the
birth of a black calf, are said by Lord Tankerville to be inferior to those
at Chillingham. The cattle kept until the year 1780 by the Duke of
Queensberry, but now extinct, had their ears, muzzle, and orbits of the
eyes black. Those which have existed from time immemorial at Chartley,
closely resemble the cattle at Chillingham, but are larger, “with some
small difference in the colour of the ears.” “They frequently tend to
become entirely black; and a singular superstition prevails in the vicinity
that, when a black calf is born, some calamity impends over the noble house
of Ferrers. All the black calves are destroyed.” The cattle at Burton
Constable in Yorkshire, now extinct, had ears, muzzle, and the tip of the
tail black. Those at Gisburne, also in Yorkshire, are said by Bewick to
have been sometimes without dark muzzles, with the inside alone of the ears
brown; and they are elsewhere said to have been low in stature and
hornless.^[51]

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

Boyd Dawkins believes that the park-cattle are descended from anciently
domesticated, and not truly wild animals; and from the occasional
appearance of dark-coloured calves, it is improbable that the aboriginal
Bos primigenius was white. It is curious what a strong, though not
invariable, tendency there is in wild or escaped cattle to become white
with coloured ears, under widely different conditions of life. If the old
writers Boethius and Leslie^[52] 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. In Wales,^[53] 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, 150 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 shorthorn breed are white, the extremities of
their ears are more or less tinged with red.

The cattle which have run wild on the Pampas, in Texas, and in two parts of
Africa, have become of a nearly uniform dark brownish-red.^[54] 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.”^[55] 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,^[56] 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 Usborn,
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 a 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 Shorthorns,
Longhorns (now rarely seen), Herefords, Highland cattle, Alderneys,
etc., must be familiar to every one. A part of this difference may
be attributed to descent from primordially distinct species; but we
may feel sure that there has been a considerable amount of
variation. Even during the Neolithic period, the domestic cattle
were to a certain extent variable. 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 Shorthorns, eleven bulls reached an
average of 214 pounds, and lately Shorthorn 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 Shorthorns 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,^[57] 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.”^[58] With respect to the period of
conception, it seems certain that Alderney and Zetland cows often become
pregnant earlier than other breeds.^[59] Lastly, as four fully developed mammæ
is a generic character in the genus Bos,^[60] 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-civilised countries, it
may be well to show that in some countries inhabited by barbarous races,
who are frequently at war with each other, and therefore have little free
communication, several distinct breeds of cattle now exist or formerly
existed. At the Cape of Good Hope Leguat observed, in the year 1720, three
kinds.^[61] 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^[62] 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 also 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. 8-1/4 in. long, as measured in a straight line
from tip to tip, and no less than 13 ft. 5 in. 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^[63] describes two peculiar breeds, namely,
pelones, with extremely thin and fine hair, and calongos,
absolutely naked. According to Castelnau there are two races in Brazil, one
like European cattle, the other different, with remarkable horns. In
Paraguay, Azara describes a breed which certainly originated in S. America,
called chivos, “because they have straight vertical horns, conical,
and very large at the base.” He likewise describes a dwarf race in
Corrientes, with short legs and a body larger than usual. Cattle without
horns, and others with reversed hair, have also originated in Paraguay.

Another monstrous breed, called niatas or natas, of which I saw two small
herds on the northern bank of the Plata, is so remarkable as to deserve a
fuller description. This breed bears the same relation to other breeds, as
bull or pug dogs do to other dogs, or as improved pigs, according to H. von
Nathusius, do to common pigs.^[64] Rütimeyer believes that these cattle
belong to the primigenius type.^[65] 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 confirmation characterizes, as I am 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:^[66] “It is remarkable from the stunted
development of the nasals, premaxillaries, and fore-part of the lower jaw,
which is unusually curved upwards to come into contact with the
premaxillaries. The nasal bones are about one-third the ordinary length,
but retain almost their normal breadth. The triangular vacuity is left
between them, the frontal and lachrymal, which latter bone articulates with
the premaxillary, and thus excludes the maxillary from any junction with
the nasal.” So that even the connexion of some of the bones is changed.
Other differences might be added: thus the plane of the condyles is
somewhat modified, and the terminal edge of the premaxillaries forms an
arch. In fact, on comparison with the skull of a common ox, scarcely a
single bone presents the same exact shape, and the whole skull has a
wonderfully different appearance.

The first brief published notice of this race
was by Azara, between the years 1783-96; but Don F. Muniz, of
Luxan, who has kindly collected information for me, states that
about 1760 these cattle were kept as curiosities near Buenos Ayres.
Their origin is not positively known, but they must have originated
subsequently to the year 1552, when cattle were first introduced.
Senor 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 civilised 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 Senor 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 to keep alive by browsing with their lips on
the twigs of trees and on reeds: this the niatas cannot so well do,
as their lips do not join, and hence they are found to perish
before the common cattle. This strikes me as a good illustration of
how little we are able to judge from the ordinary habits of an
animal, on what circumstances, occurring only at long intervals of
time, its rarity or extinction may depend. It shows us, also, how
natural selection would have determined the rejection of the niata
modification had it arisen in a state of nature.

Having described the semi-monstrous niata breed, I may allude to a white
bull, said to have been brought from Africa, which was exhibited in London
in 1829, and which has been well figured by Mr. Harvey.^[67] 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 cause is far from
sufficient. 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.^[68] That climate directly affects the
thickness of the skin and the hair is likewise certain: thus Roulin
asserts^[69] 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^[70] that the cattle which inhabit the more
humid parts of Britain have longer hair and thicker skins than other
British cattle. 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, etc., have appeared suddenly owing to what we may call in
our ignorance spontaneous variation; but even in this case a rude kind of
selection is necessary, and the animals thus characterised must be at least
partially separated from others. This degree of care, however, has
sometimes been taken even in little-civilised 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 Longhorn Bull, Shakespeare, though of the pure Canley stock,
scarcely inherited a single point of the long-horned breed, his horns
excepted;^[71] yet in the hands of Mr. Fowler, this
bull greatly improved his race. We have also reason to believe that
selection, carried on so far unconsciously that there was at no one time
any distinct intention to improve or change the breed, has in the course of
time modified most of our cattle; for by this process, aided by more
abundant food, all the lowland British breeds have increased greatly in
size and in early maturity since the reign of Henry VII.^[72] 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, etc., 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 were 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. Mr. Blyth, who has carefully
attended to the subject, believes that fourteen wild species now exist, but
“that not one of them can be identified as the progenitor of any one of the
interminable domestic races.” M. Gervais thinks that there are six species
of Ovis,^[73] but that our domestic sheep form a
distinct genus, now completely extinct. A German naturalist^[74]
believes that our sheep descend from ten aboriginally distinct species, of
which only one is still living in a wild state! Another ingenious
observer,^[75] 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^[76] found
in the Swiss lake-dwellings the remains of a small breed, with thin tall
legs, and horns like those of a goat, thus differing somewhat from any kind
now known. Almost every country has its own peculiar breed; and many
countries have several 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 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, bear
in their drooping ears the stamp of long domestication. This is likewise
the case with those sheep which have two great masses of fat on the rump,
with the tail in a rudimentary condition. The Angola variety of the
long-tailed race has curious masses of fat on the back of the head and
beneath the jaws.^[77] Mr. Hodgson in an admirable paper^[78] 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 not rarely absent, especially in the
female sex, 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.”^[79] This correlation, however, is far from
being general; for instance, 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 Càgias (a
sub-Himalayan domestic race) possessed of four teats.”^[80] 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^[81] has
shown that these pits or pouches are absent in some breeds.

In sheep there is a strong tendency for characters, which have apparently
been acquired under domestication, to become attached either exclusively to
the male sex, or to be more highly developed in this than in the other sex.
Thus in many breeds the horns are deficient in the ewe, though this
likewise occurs occasionally with the female of the wild musmon. In the
rams of the Wallachian breed, “the horns spring almost perpendicularly from
the frontal bone, and then take a beautiful spiral form; in the ewes they
protrude nearly at right angles from the head, and then become twisted in a
singular manner.”^[82] 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.^[83] 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^[84] 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.”^[85] Marshall relates^[86] that a flock of heavy
Lincolnshire and light Norfolk sheep which had been bred together in a
large sheep-walk, part of which was low, rich, and moist, and another part
high and dry, with benty grass, when turned out, regularly separated from
each other; the heavy sheep drawing off to the rich soil, and the lighter
sheep to their own soil; so that “whilst there was plenty of grass the two
breeds kept themselves as distinct as rooks and pigeons.” Numerous sheep
from various parts of the world have been brought during a long course of
years to the Zoological Gardens of London; but as Youatt, who attended the
animals as a veterinary surgeon, remarks, “few or none die of the rot, but
they are phthisical; not one of them from a torrid climate lasts out the
second year, and when they die their lungs are tuberculated.”^[87] There
is very good evidence that English breeds of sheep will not succeed in
France.^[88] 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^[89] that the owner could not keep them; the
coarser-skinned sheep never being affected.

The period of gestation was formerly thought to be of so unalterable a
character, that a supposed difference of this kind between the wolf and the
dog 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,^[90] that
Merino and Southdown sheep, when both have long been kept under exactly the
same conditions, differ in their average period of gestation, as is seen in
the following Table:—

In this graduated difference in 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 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.^[91] 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.^[92] On
the other hand, many wool-bearing sheep live on the hot plains of India.
Roulin asserts that in the lower and heated valleys of the Cordillera, if
the lambs are sheared as soon as the wool has grown to a certain thickness,
all goes on afterwards as usual; but if not sheared, the wool detaches
itself in flakes, and short shining hair like that on a goat is produced
ever afterwards. This curious result seems merely to be an exaggerated
tendency natural to the Merino breed, for as a great authority, namely,
Lord Somerville, remarks, “the wool of our Merino sheep after shear-time is
hard and coarse to such a degree as to render it almost impossible to
suppose that the same animal could bear wool so opposite in quality,
compared to that which has been clipped from it: as the cold weather
advances, the fleeces recover their soft quality.” As in sheep of all
breeds the fleece naturally consists of longer and coarser hair covering
shorter and softer wool, the change which it often undergoes in hot
climates is probably merely a case of unequal development; for even with
those sheep which like goats are covered with hair, a small quantity of
underlying wool may always be found.^[93] In the wild mountain-sheep (0vis
montana) of North America there is an analogous annual 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, etc.,
which shed their winter coat in the spring.”^[94]

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

That methodical selection has effected great changes in several breeds of
sheep no one who knows anything on the subject, entertains a doubt. The
case of the Southdowns, as improved by Ellman, offers perhaps the most
striking instance. Unconscious or occasional selection has likewise slowly
produced a great effect, as we shall see in the chapters on Selection. That
crossing has largely modified some breeds, no one who will study what has
been written on this subject—for instance, Mr. Spooner’s
paper—will dispute; but to produce uniformity in a crossed breed,
careful selection and “rigorous weeding,” as this author expresses it, are
indispensable.^[95]

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. The sheep are remarkable
from transmitting their character so truly that Colonel Humphreys^[96] 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; even one of twins has resembled one parent and the
second the other. Lastly, “the ancons have been observed to keep together,
separating themselves from the rest of the flock when put into enclosures
with other sheep.”

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

GOATS.

From the recent researches of M. Brandt, most naturalists now believe that
all our goats are descended from the Capra ægagrus of the mountains
of Asia, possibly mingled with the allied Indian species C.
falconeri of India.^[97] In Switzerland, during the neolithic
period, the domestic goat was commoner than the sheep; and this very
ancient race differed in no respect from that now common in Switzerland.^[98] 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,^[99] they
are all quite fertile when crossed. So numerous are the breeds, that Mr. G.
Clark^[100] 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 4-3/4 inches in breadth. As with cattle, the mammæ of
those breeds which are regularly milked become greatly developed; and, as
Mr. Clark remarks, “it is not rare to see their teats touching the ground.”
The following cases are worth notice as presenting unusual points of
variation. According to Godron,^[101] 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;^[102] and in some breeds
the females are destitute of horns.^[103] M. Ramu of Nancy informs me that many
of the goats there bear on the upper part of the throat a pair of hairy
appendages, 70 mm. in length and about 10 mm. in diameter, which in
external appearance resemble those above described on the jaws of pigs. The
presence of inter-digital 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.^[104] 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!

REFERENCES

[1]
Hermann von Nathusius ‘Die Racen des Schweines,’ Berlin, 1860; and ‘Vorstudien
für Geschichte,’ etc., ‘Schweineschädel,’ Berlin, 1864. Rütimeyer, ‘Die Fauna
der Pfahlbauten,’ Basel, 1861.

[2]
Nathusius, ‘Die Racen des Schweines,’ Berlin, 1860. An excellent appendix is
given with references to published and trustworthy drawings of the breeds of
each country.

[3]
For Europe see Bechstein, ‘Naturgesch. Deutschlands,’ 1801, B. i., s.
505. Several accounts have been published on the fertility of the offspring
from wild and tame swine. See Burdach’s ‘Physiology,’ and Godron ‘De
l’Espèce,’ tom. i. p. 370. For Africa, ‘Bull. de la Soc. d’Acclimat.’ tom. iv.
p. 389. For India, see Nathusius, ‘Schweineschädel,’ s. 148.

[4]
Sir W. Elliot, Catalogue of Mammalia, ‘Madras Journal of Lit. and Science,’
vol. x. p. 219.

[5]
‘Pfahlbauten,’ s. 163 et passim.

[6]
See J. W. Schütz’ interesting essay, ‘Zur Kenntniss des Torfschweins,’
1868. This author believes that the Torfschwein is descended from a distinct
species, the S. sennariensis of Central Africa.

[7]
Stan. Julien quoted by de Blainville, ‘Ostéographie,’ p. 163.

[8]
Richardson, ‘Pigs, their Origin,’ etc., p. 26.

[9]
‘Die Racen des Schweines’ s. 47, 64.

[10]
‘Proc. Zoolog. Soc.,’ 1861, p. 263.

[11]
Sclater, in ‘Proc. Zoolog. Soc.,’ Feb. 26, 1861.

[12]
‘Proc. Zoolog. Soc.,’ 1862, p. 13. The skull has since been described much more
fully by Professor Lucae in a very interesting essay, ‘Der Schädel des
Maskenschweines,’ 1870. He confirms the conclusion of von Nathusius on the
relationship of this kind of pig.

[13]
‘Journal of Voyages and Travels from 1821 to 1829,’ vol. i. p. 300.

[14]
Rev. G. Low ‘Fauna Orcadensis,’ p. 10. See also Dr. Hibbert’s account of
the pig of the Shetland Islands.

[15]
‘Die Racen des Schweines’ s. 70.

[16]
These woodcuts are copied from engravings given in Mr. S. Sidney’s excellent
edition of ‘The Pig,’ by Youatt, 1860. See pp. 1, 16, 19.

[17]
‘Schweineschädel’ s. 74, 135.

[18]
Nathusius, ‘Die Racen des Schweines,’ s. 71.

[19]
‘Die Racen des Schweines,’ s. 47. ‘Schweineschädel’ s. 104. Compare, also, the
figures of the old Irish and the improved Irish breeds in Richardson on ‘The
Pig,’ 1847.

[20]
Quoted by Isid. Geoffroy, ‘Hist. Nat. Gén.,’ tom. iii. p. 441.

[21]
S. Sidney, ‘The Pig,’ p. 61.

[22]
‘Schweineschädel,’ s. 2, 20.

[23]
‘Proc. Zoolog. Soc.,’ 1837, p. 23. I have not given the caudal vertebræ, as Mr.
Eyton says some might possibly have been lost. I have added together the dorsal
and lumbar vertebræ, owing to Prof. Owen’s remarks (‘Journal Linn. Soc.,’ vol.
ii. p. 28) on the difference between dorsal and lumbar vertebræ depending only
on the development of the ribs. Nevertheless the difference in the number of
the ribs in pigs deserves notice. M. Sanson gives the number of lumbar vertebræ
in various pigs; ‘Comptes Rendus,’ lxiii. p. 843.

[24]
‘Edinburgh New Philosoph. Journal,’ April, 1863. See also De
Blainville’s ‘Ostéographie,’ p. 128, for various authorities on this subject.

[25]
Eudes-Deslongchamps, ‘Mémoires de la Soc. Linn. de Normandie,’ vol. vii., 1842,
p. 41. Richardson, ‘Pigs, their Origin, etc.,’ 1847, p. 30. Nathusius, ‘Die
Racen des Schweines,’ 1863, s. 54.

[26]
D. Johnson’s ‘Sketches of Indian Field Sports,’ p. 272. Mr. Crawfurd informs me
that the same fact holds good with the wild pigs of the Malay peninsula.

[27]
For Turkish pigs see Desmarest, ‘Mammalogie,’ 1820, p. 391. For those of
Westphalia see Richardson’s ‘Pigs, their Origin, etc.,’ 1847, p. 41.

[28]
With respect to the several foregoing and following statements on feral pigs,
see Roulin, in ‘Mém. présentés par divers Savans a l’Acad.,’ etc.,
Paris, tom. vi. 1835, p. 326. It should be observed that his account does not
apply to truly feral pigs; but to pigs long introduced into the country and
living in a half-wild state. For the truly feral pigs of Jamaica, see
Gosse’s ‘Sojourn in Jamaica,’ 1851, p. 386; and Col. Hamilton Smith, in ‘Nat.
Library,’ vol. ix. p. 93. With respect to Africa see Livingstone’s
‘Expedition to the Zambesi,’ 1865, p. 153. The most precise statement with
respect to the tusks of the West Indian feral boars is by P. Labat (quoted by
Roulin); but this author attributes the state of these pigs to descent from a
domestic stock which he saw in Spain. Admiral Sulivan, R.N., had ample
opportunities of observing the wild pigs on Eagle Islet in the Falklands; and
he informs me that they resembled wild boars with bristly ridged backs and
large tusks. The pigs which have run wild in the province of Buenos Ayres
(Rengger ‘Säugethiere,’ s. 331) have not reverted to the wild type. De
Blainville (‘Ostéographie,’ p. 132) refers to two skulls of domestic pigs sent
from Patagonia by Al. d’Orbigny, and he states that they have the occipital
elevation of the wild European boar, but that the head altogether is “plus
courte et plus ramassée.” He refers, also, to the skin of a feral pig from
North America, and says “il ressemble tout à fait à un petit sanglier, mais il
est presque tout noir, et peut-être un peu plus ramassé dans ses formes.”

[29]
Gosse’s ‘Jamaica,’ p. 386, with a quotation from Williamson’s ‘Oriental Field
Sports.’ Also Col. Hamilton Smith, in ‘Naturalist Library,’ vol. ix. p. 94.

[30]
S. Sidney’s edition of ‘Youatt on the Pig,’ 1860, pp. 7, 26, 27, 29, 30.

[31]
‘Schweineschädel’ s. 140.

[32]
‘Die Fauna der Pfahlbauten,’ 1861, s. 109, 149, 222. See also Geoffroy
Saint-Hilaire in ‘Mém. du Mus. d’Hist. Nat.,’ tom. x. p. 172; and his son
Isidore in ‘Hist. Nat. Gen.’ tom. iii. p. 69. Vasey, in his ‘Delineations of
the Ox Tribe,’ 1851, p. 127, says the zebu has four, and common ox five, sacral
vertebræ. Mr. Hodgson found the ribs either thirteen or fourteen in number;
see a note in ‘Indian Field,’ 1858, p. 62.

[33]
‘The Indian Field,’ 1858, p. 74, where Mr. Blyth gives his authorities with
respect to the feral humped cattle. Pickering, also, in his ‘Races of Man,’
1850, p. 274, notices the peculiar grunt-like character of the voice of the
humped cattle.

[34]
Mr. H. E. Marquand, in ‘The Times,’ June 23rd, 1856.

[35]
Vasey, ‘Delineations of the Ox-Tribe,’ p. 124. Brace’s ‘Hungary,’ 1851, p. 94.
The Hungarian cattle descend, according to Rütimeyer ‘Zahmen Europ. Rindes,’
1866, s. 13 from Bos primigenius.

[36]
Moll and Gayot, ‘La Connaissance Gén. du Bœuf,’ Paris, 1860. Fig. 82 is that of
the Podolian breed.

[37]
A translation appeared in three parts in the ‘Annals and Mag. of Nat. Hist.,’
2nd series, vol. iv., 1849.

[38]
See also Rütimeyer’s ‘Beiträge pal. Gesch. der Wiederkäuer Basel,’ 1865,
s. 54.

[39]
Pictet ‘Paléontologie,’ tom. i. p. 365 (2nd edit.). With respect to B.
trochoceros, see Rütimeyer ‘Zahmen Europ. Rindes,’ 1866, s. 26.

[40]
W. Boyd Dawkins on the British Fossil Oxen, ‘Journal of the Geolog. Soc.,’ Aug.
1867, p. 182. Also ‘Proc. Phil. Soc. of Manchester,’ Nov. 14th, 1871, and ‘Cave
Hunting,’ 1875, p. 27, 138.

[41]
‘British Pleistocene Mammalia,’ by W. B. Dawkins and W. A. Sandford, 1866, p.
15.

[42]
W. R. Wilde, ‘An Essay on the Animal Remains, etc. Royal Irish Academy,’ 1860,
p. 29. Also ‘Proc. of R. Irish Academy,’ 1858, p. 48.

[43]
‘Lecture: Royal Institution of G. Britain,’ May 2nd, 1856, p. 4. ‘British
Fossil Mammals,’ p. 513.

[44]
Nilsson, in ‘Annals and Mag. of Nat. Hist.,’ 1849, vol. iv. p. 354.

[45]
See W. R. Wilde, ut supra; and Mr. Blyth, in ‘Proc. Irish Academy,’
March 5th, 1864.

[46]
Laing’s ‘Tour in Norway,’ p. 110.

[47]
Isid. Geoffroy Saint-Hilaire, ‘Hist. Nat. Gén.,’ tom. iii. 96.

[48]
Idem, tom. iii. pp. 82, 91.

[49]
‘Quadrupèdes du Paraguay,’ tom. ii. p. 360.

[50]
Walther ‘Das Rindvieh,’ 1817, s. 30.

[51]
I am much indebted to the present Earl of Tankerville for information about his
wild cattle; and for the skull which was sent to Prof. Rütimeyer. The fullest
account of the Chillingham cattle is given by Mr. Hindmarsh, together with a
letter by the late Lord Tankerville, in ‘Annals and Mag. of Nat. Hist.,’ vol.
ii., 1839, p. 274. See Bewick, ‘Quadrupeds,’ 2nd edit., 1791, p. 35,
note. With respect to those of the Duke of Queensberry, see Pennant’s
‘Tour in Scotland,’ p. 109. For those of Chartley, see Low’s
‘Domesticated Animals of Britain,’ 1845, p. 238. For those of Gisburne,
see Bewick ‘Quadrupeds,’ and ‘Encyclop. of Rural Sports,’ p. 101.

[52]
Boethius was born in 1470; ‘Annals and Mag. of Nat. Hist.,’ vol. ii., 1839, p.
281; and vol. iv., 1849, p. 424.

[53]
n’Youatt on Cattle,’ 1834, p. 48: See also p. 242, on short-horn cattle.
Bell, in his ‘British Quadrupeds,’ p. 423, states that, after long attending to
the subject, he has found that white cattle invariably have coloured ears.ote

[54]
Azara, ‘Quadrupèdes du Paraguay,’ tom. ii. p. 361. Azara quotes Buffon for the
feral cattle of Africa. For Texas see ‘Times,’ Feb. 18th, 1846.

[55]
Anson’s Voyage. See Kerr and Porter’s ‘Collection,’ vol. xii. p. 103.

[56]
See also Mr. Mackinnon’s pamphlet on the Falkland Islands, p. 24.

[57]
‘The Age of the Ox, Sheep, Pig,’ etc., by Prof. James Simonds, published by
order of the Royal Agricult. Soc.

[58]
‘Ann. Agricult. France,’ April, 1837, as quoted in ‘The Veterinary,’ vol. xii.
p. 725. I quote Tessier’s observations from ‘Youatt on Cattle,’ p. 527.

[59]
‘The Veterinary,’ vol. viii. p. 681 and vol. x. p. 268. Low’s ‘Domest. Animals,
etc.’ p. 297.

[60]
Mr. Ogleby in ‘Proc. Zoolog. Soc.,’ 1836, p. 138, and 1840, p. 4. Quatrefages
quotes Philippi (‘Revue des Cours Scientifiques,’ Feb. 12, 1688, p. 657), that
the cattle of Piacentino have thirteen dorsal vertebræ and ribs in the place of
the ordinary number of twelve.

[61]
Leguat’s Voyage, quoted by Vasey in his ‘Delineations of the Ox-tribe,’ p. 132.

[62]
‘Travels in South Africa,’ pp. 317, 336.

[63]
‘Mem. de l’Institut présent. par divers Savans,’ tom. vi., 1835, p. 333. For
Brazil, see ‘Comptes Rendus,’ June 15, 1846. See Azara
‘Quadrupèdes du Paraguay,’ tom. ii. pp. 359, 361.

[64]
‘Schweineschädel,’ 1864, s. 104. Nathusius states that the form of skull
characteristic in the niata cattle occasionally appears in European cattle; but
he is mistaken, as we shall hereafter see, in supposing that these cattle do
not form a distinct race. Prof. Wyman, of Cambridge, United States, informs me
that the common cod-fish presents a similar monstrosity, called by the
fishermen “bull-dog cod.” Prof. Wyman also concluded, after making numerous
inquiries in La Plata, that the niata cattle transmit their peculiarities or
form a race.

[65]
‘Ueber Art des zahmen Europ. Rindes,’ 1866, s. 28.

[66]
‘Descriptive Cat. of Ost. Collect. of College of Surgeons,’ 1853, p. 624. Vasey
in his ‘Delineations of the Ox-tribe’ has given a figure of this skull; and I
sent a photograph of it to Prof. Rütimeyer.

[67]
Loudon’s ‘Magazine of Nat. Hist.,’ vol. i. 1829, p. 113. Separate figures are
given of the animal, its hoofs, eye, and dewlap.

[68]
Low, ‘Domesticated Animals of the British Isles,’ p. 264.

[69]
‘Mém. de l’Institut présent. Par divers Savans,’ tom. vi., 1835, p. 332.

[70]
Idem, pp. 304, 368, etc.

[71]
‘Youatt on Cattle,’ p. 193. A full account of this bull is taken from Marshall.

[72]
‘Youatt on Cattle,’ p. 116. Lord Spencer has written on this same subject.

[73]
Blyth, on the genus Ovis, in ‘Annals and Mag. of Nat. History,’ vol. vii.,
1841, p. 261. With respect to the parentage of the breeds see Mr.
Blyth’s excellent articles in ‘Land and Water,’ 1867, pp. 134, 156. Gervais,
‘Hist. Nat. des Mammifères,’ 1855, tom. ii. p. 191.

[74]
Dr. L. Fitzinger, ‘Ueber die Racen des Zahmen Schafes,’ 1860, s. 86.

[75]
J. Anderson, ‘Recreations in Agriculture and Natural History,’ vol. ii. p. 264.

[76]
‘Pfahlbauten’ s. 127, 193.

[77]
‘Youatt on Sheep,’ p. 120.

[78]
‘Journal of the Asiatic Soc. of Bengal,’ vol.xvi. pp. 1007, 1016.

[79]
‘Youatt on Sheep,’ pp. 142-169.

[80]
‘Journal Asiat. Soc. of Bengal,’ vol. xvi., 1847, p. 1015.

[81]
‘Hist. Nat. Gén.,’ tom. iii. p. 435.

[82]
‘Youatt on Sheep,’ p. 138.

[83]
‘Journal Asiat. Soc. of Bengal,’ vol. xvi., 1847, pp. 1015, 1016.

[84]
‘Racen des Zahmen Schafes,’ s. 77.

[85]
‘Rural Economy of Norfolk,’ vol. ii. p. 136.

[86]
‘Youatt on Sheep,’ p. 312. On same subject, see excellent remarks in
‘Gardener’s Chronicle,’ 1858, p. 868. For experiments in crossing Cheviot sheep
with Leicesters see Youatt, p. 325.

[87]
‘Youatt on Sheep,’ note, p. 491.

[88]
M. Malingié-Nouel, ‘Journal R. Agricult. Soc.,’ vol. xiv. 1853, p. 214.
Translated and therefore approved by a great authority, Mr. Pusey.

[89]
‘The Veterinary,’ vol. x. p. 217.

[90]
A translation of his paper is given in ‘Bull. Soc. Imp. d’Acclimat.,’ tom. ix.,
1862, p. 723.

[91]
Erman’s ‘Travels in Siberia,’ (Eng. trans.) vol. i. p. 228. For Pallas on the
fat-tailed sheep I quote from Anderson’s account of the ‘Sheep of Russia,’
1794, p. 34. With respect to the Crimean sheep see Pallas’ ‘Travels’
(Eng. trans.) vol. ii. p. 454. For the Karakool sheep see Burnes’
‘Travels in Bokhara,’ vol. iii. p. 151.

[92]
See Report of the Directors of the Sierra Leone Company, as quoted in
White’s ‘Gradation of Man,’ p. 95. With respect to the change which sheep
undergo in the West Indies see also Dr. Davy, in ‘Edin. New. Phil.
Journal,’ Jan. 1852. For the statement made by Roulin, see ‘Mém. de
l’Institut présent. par divers Savans,’ tom. vi., 1835, p. 347.

[93]
‘Youatt on Sheep,’ p. 69, where Lord Somerville is quoted. See p. 117 on
the presence of wool under the hair. With respect to the fleeces of Australian
sheep, p. 185. On selection counteracting any tendency to change, see
pp. 70, 117, 120, 168.

[94]
Audubon and Bachman, ‘The Quadrupeds of North America,’ 1846, vol. v. p. 365.

[95]
‘Journal of R. Agricult. Soc. of England,’ vol. xx., part ii., W. C. Spooner on
cross-Breeding.

[96]
‘Philosoph. Transactions,’ London, 1813, p. 88.

[97]
Isidore Geoffroy St. Hilaire, ‘Hist. Nat. Générale,’ tom. iii. p. 87. Mr.
Blyth, (‘Land and Water,’ 1867, p. 37) has arrived at a similar conclusion, but
he thinks that certain Eastern races may perhaps be in part descended from the
Asiatic markhor.

[98]
Rütimeyer ‘Pfahlbauten,’ s. 127.

[99]
Godron ‘De l’Espèce,’ tom. i. p. 402.

[100]
‘Annals and Mag. of Nat. History,’ vol ii. (2nd series), 1848, p. 363.

[101]
‘De l’Espèce,’ tom. i. p. 406. Mr. Clark also refers to differences in the
shape of the mammæ. Godron states that in the Nubian race the scrotum is
divided into two lobes; and Mr. Clark gives a ludicrous proof of this fact, for
he saw in the Mauritius a male goat of the Muscat breed purchased at a high
price for a female in full milk. These differences in the scrotum are probably
not due to descent from distinct species: for Mr. Clark states that this part
varies much in form.

[102]
Mr. Clark, ‘Annals and Mag. of Nat. Hist.,’ vol. ii. (2nd series), 1848, p.
361.

[103]
Desmarest, ‘Encyclop. Méthod. Mammalogie,’ p. 480.

[104]
‘Journal of Asiatic Soc. of Bengal,’ vol. xvi., 1847, pp. 1020, 1025.

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^[1] states “that the true wild rabbit is
smaller than the domestic; its proportions are not absolutely the same; its
tail is smaller; its ears are shorter and more thickly clothed with hair;
and these characters, without speaking of colour, are so many indications
opposed to the opinion which unites these animals under the same specific
denomination.” Few naturalists will agree with this author that such slight
differences are sufficient to separate as distinct species the wild and
domestic rabbit. How extraordinary it would be, if close confinement,
perfect tameness, unnatural food, and careful breeding, all prolonged
during many generations, had not produced at least some effect! The tame
rabbit has been domesticated from an ancient period. Confucius ranges
rabbits among animals worthy to be sacrificed to the gods, and, as he
prescribes their multiplication, they were probably at this early period
domesticated in China. They are mentioned by several of the classical
writers. In 1631 Gervaise Markham writes, “You shall not, as in other
cattell, looke to their shape, but to their richnesse, onely elect your
buckes, the largest and goodliest conies you can get; and for the richnesse
of the skin, that is accounted the richest which hath the equallest mixture
of blacke and white haire together, yet the blacke rather shadowing the
white; the furre should be thicke, deepe, smooth, and shining; … they are
of body much fatter and larger, and, when another skin is worth two or
three pence, they are worth two shillings.” From this full description we
see that silver-grey rabbits existed in England at this period; and what is
far more important, we see that the breeding or selection of rabbits was
then carefully attended to. Aldrovandi, in 1637, describes, on the
authority of several old writers (as Scaliger, in 1557), rabbits of various
colours, some “like a hare,” and he adds that P. Valerianus (who died a
very old man in 1558) saw at Verona rabbits four times bigger than ours.^[2]

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.^[3] 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.^[4] Wild rabbits, if taken
young, can be domesticated, though the process is generally very
troublesome.^[5] The various domestic races are often
crossed, and are believed to be quite 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.^[6] 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, these, with the exception of 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 marvellous success in France in rearing hybrids between the hare and
rabbit,^[7]
it is possible, though not probable, from the great difficulty in making
the first cross, that some of the larger races, which are coloured like the
hare, may have been modified by crosses with this animal. Nevertheless, the
chief differences in the skeletons of the several domestic breeds cannot,
as we shall presently see, have been derived from a cross with the hare.

There are many breeds which transmit their characters more or less truly.
Every one has seen the enormous lop-eared rabbits exhibited at our shows;
various allied sub-breeds are reared on the Continent, such as the
so-called Andalusian, which is said to have a large head with a round
forehead, and to attain a greater size than any other kind; another large
Paris breed is named the Rouennais, and has a square head; the so-called
Patagonian rabbit has remarkably short ears and a large round head.
Although I have not seen all these breeds, I feel some doubt about there
being any marked difference in the shape of their skulls.^[8] English lop-eared
rabbits often weigh 8 pounds or 10 pounds, and one has been exhibited
weighing 18 pounds; whereas a full-sized wild rabbit weighs only about
3-1/4 pounds. 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 was exhibited in 1867 with its two ears, measured
from the tip of one to the tip of the other, 22 inches in length, and each
ear 5-3/8 inches in breadth. In 1869 one was exhibited with ears, measured
in the same manner, 23-1/8 in length and 5-1/2 in breadth; “thus exceeding
any rabbit ever exhibited at a prize show.” In a common wild rabbit I found
that the length of two ears, from tip to tip, was 7-5/8 inches, and the
breadth only 1-7/8 inch. The weight of body in the larger rabbits, and the
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.^[9]
Two live rabbits were brought to me from Moscow, of about the size of the
wild species, but with long soft fur, different from that of the Angora.
These Moscow rabbits had pink eyes and were snow-white, excepting the ears,
two spots near the nose, the upper and under surface of the tail, and the
hinder tarsi, which were blackish-brown. In short, they were coloured
nearly like the so-called Himalayan rabbits, presently to be described, and
differed from them only in the character of their fur. There are two other
breeds which come true to colour, but differ in no other respect, namely
silver-greys and chinchillas. Lastly, the Nicard or Dutch rabbit may be
mentioned, which varies in colour, and is remarkable from its small size,
some specimens weighing only 1-1/4 pounds; rabbits of this breed make
excellent nurses for other and more delicate kinds.^[10]

Fig. 5—Half-lop Rabbit.

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,^[11] “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 down flat 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 characterised. 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;^[12] 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 results from the great length and weight of the ear, favoured
no doubt by the weakness of the muscles consequent on disuse. Anderson^[13]
mentions a breed having only a single ear; and Professor Gervais another
breed destitute of ears.

We come now to the Himalayan breed, which is sometimes called Chinese,
Polish, or Russian. These pretty rabbits are white, or occasionally yellow,
excepting their ears, nose, feet, and the upper side of the 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.^[14] Some
good observers thought that they could detect a difference in their habits,
and stoutly maintained that they formed a new species. The origin of this
breed is so curious, both in itself and as throwing some light on the
complex laws of inheritance that it is worth giving in detail. But it is
first necessary briefly to describe two other breeds: silver-greys or
silver-sprigs generally have black heads and legs, and their fine grey fur
is interspersed with numerous black and white long hairs. They breed
perfectly true, and have long been kept in warrens. When they escape and
cross with common rabbits, the product, as I hear from Mr. Wyrley Birch, of
Wretham Hall, is not a mixture of the two colours, but about half take
after the one parent, and the other half after the other parent. Secondly,
chinchillas or tame silver-greys (I will use the former name) have short,
paler, mouse or slate-coloured fur, interspersed with long, blackish,
slate-coloured, and white hairs.^[15] These rabbits breed perfectly true. A
writer stated in 1857^[16] that he had produced Himalayan rabbits
in the following manner. He 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^[17] 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.
But I have recently been assured the pure silver-greys of any sub-breed
occasionally produce Himalayans.

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 with the Himalayans in their colour whilst quite young,
for they are born perfectly black, but soon assume their characteristic
grey or silver tints. The same thing occurs with grey horses, which, as
long as they are foals, are generally of a nearly black colour, but soon
become grey, and get whiter and whiter as they grow older. Hence the usual
rule is that Himalayans are born white and afterwards become in certain
parts of their bodies dark-coloured; whilst silver-greys are born black and
afterwards become sprinkled with white. Exceptions, however, and of a
directly opposite nature, occasionally occur in both cases. For young
silver-greys are sometimes born in warrens, as I hear from Mr. W. Birch, of
a cream-colour, but these young animals ultimately become black. The
Himalayans, on the other hand, sometimes produce, as is stated by an
experienced amateur,^[18] a single black young one in a litter;
and this, before two months elapse, becomes perfectly white.

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

It is, also, remarkable that Himalayans, though produced so suddenly; breed
true. But as, whilst young, they are albinoes, the case falls under a very
general rule; albinism being well known to be strongly inherited, for
instance with white mice and many other quadrupeds, and even 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 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^[19] 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 the several coloured marks on
the Himalayan rabbits, as they grow old, are rendered intelligible. I may
add a nearly analogous case: fancy rabbits very often have a white star on
their foreheads; and the common English hare, whilst young, generally has,
as I have myself observed, a similar white star on its forehead.

When variously coloured rabbits are set free in Europe, and are thus placed
under their natural conditions, they generally revert to the aboriginal
grey colour; this may be in part due to the tendency in all crossed
animals, as lately observed, to revert to their primordial state. But this
tendency does not always prevail; thus silver-grey rabbits are kept in
warrens, and remain true though living almost in a state of nature; but a
warren must not be stocked with both silver-greys and common rabbits;
otherwise “in a few years there will be none but common greys surviving.”^[20] When
rabbits run wild in foreign countries under new conditions of life, they by
no means always revert to their aboriginal colour. In Jamaica the feral
rabbits are described as having been “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.”^[21] But in this tropical
island the conditions were not favourable to their increase, and they never
spread widely, and are now extinct, as I hear from Mr. R. Hill, owing to a
great fire which occurred in the woods. 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.^[22] 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 of 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^[23]
happened to have a female rabbit on board which had produced young during
the voyage, and he turned them all out on the island. These animals soon
increased so rapidly, that they became a nuisance, and actually caused the
abandonment of the settlement. Thirty-seven years subsequently, Cada Mosto
describes them as innumerable; nor is this surprising, as the island was
not inhabited by any beast of prey or by any terrestrial mammal. We do not
know the character of the mother-rabbit; but it was probably 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; and 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 17-3/4 inches in
length; whilst two of the Porto Santo rabbits were only 14-1/2 and 15
inches in length. But the decrease in size is best shown by weight; four
wild English rabbits averaged 3 pounds 5 ounces, whilst one of the Porto
Santo rabbits, which had lived for four years in the Zoological Gardens,
but had become thin, weighed only 1 pound 9 ounces. A fairer test is
afforded by the comparison of the well-cleaned limb-bones of a Porto 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.^[24] The head has not
decreased in length proportionally with the body; and the capacity of the
brain case is, as we shall hereafter see, singularly variable. I prepared
four skulls, and these resembled each other more closely than do generally
the skulls of wild English rabbits; but the only difference in structure
which they presented was that the supra-orbital processes of the frontal
bones were narrower.

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

The two little Porto Santo rabbits, whilst alive
in the Zoological Gardens, had a remarkably different appearance
from the common kind. They were extraordinarily wild and active, so
that many persons exclaimed on seeing them that they were more like
large rats than rabbits. They were nocturnal to an unusual degree
in their habits, and their wildness was never in the least subdued;
so that the superintendent, Mr. Bartlett, assured me that he had
never had a wilder animal under his charge. This is a singular
fact, considering that they are descended from a domesticated
breed. I was so much surprised at it, that I requested Mr. Haywood
to make inquiries on the spot, whether they were much hunted by the
inhabitants, or persecuted by hawks, or cats, or other animals; but
this is not the case, and no cause can be assigned for their
wildness. They live both on the central, higher rocky land and near
the sea-cliffs, and, from 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 colour, reddish above and grey beneath, their tails and
ears not 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, as certainly originated since the year 1420. Finally, from
the three cases of the rabbits which have run wild in Porto Santo,
Jamaica, and the Falkland Islands, we see that these animals do
not, under new conditions of life, revert to or retain their
aboriginal character, as is so generally asserted to be the case by
most authors.

Osteological Characters.

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

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

Skull.—I have
carefully examined skulls of ten large lop-eared 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 4·3; 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 Porto
Santo rabbit, on the other hand, the head relatively to the length
of body is about a quarter of an inch too long.

This elongation of the skull
relatively to its breadth, I find a universal character, not only
with the large lop-eared rabbits, but in all the artificial breeds;
as is well seen in the skull of the Angora. I was at first much
surprised at the fact, and could not imagine why domestication
could 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 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.

Fig. 6—Skull of Wild Rabbit. Fig. 7—Skull of
large Lop-eared Rabbit.

Fig. 8—Part of Zygomatic Arch.

In all the skulls of the large lop-eared rabbits, the supra-orbital plates
or processes of the frontal bones are 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, that is more
extended in the line of the longitudinal axis of the skull, than in the
wild rabbit. The posterior margin of “the square raised platform”^[25] 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.

Fig. 9—Posterior end of skull of Rabbits.

Fig. 10—Occipital Foramen of Rabbits.

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

In all the skulls of the large
lop-eared rabbits, the bony auditory meatus is conspicuously larger
than in the wild rabbit. In a skull 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 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 (fig. 11) of
the skull of a half-lop; and it may be observed that the suture
between the parietal and frontal bones does not run strictly at
right angles to the longitudinal axis of the skull; the left
frontal bone projects beyond the right one; both the posterior and
anterior margins of the left zygomatic arch on the side of the
lopping ear stand a little in advance of the corresponding bones on
the opposite side. Even the lower jaw is affected, and the condyles
are not quite symmetrical, that on the left standing a little in
advance of that on the right. This seems to me a remarkable case of
correlation of growth. Who would have surmised that by keeping an
animal during many generations under confinement, and so leading to
the disuse of the muscles of the ears, and by continually selecting
individuals with the longest and largest ears, he would thus
indirectly have affected almost every suture in the skull and the
form of the lower jaw!

Fig. 11—Skull of Half-lop Rabbit.

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 proportionately a little longer. The molar teeth
have increased in size proportionately with the increased width of
the skull, measured across the zygomatic arch, and not
proportionally with its increased length. The inner line of the
sockets of the molar teeth in the upper jaw of the wild rabbit
forms a perfectly straight line; but in some of the largest skulls
of the lop-eared this line was plainly bowed inwards. In one
specimen there was an additional molar tooth on each side of the
upper jaw, between the molars and premolars; but these two teeth
did not correspond in size; and as no rodent has seven molars, this
is merely a monstrosity, though a curious one.

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 rabbit,
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
rabbit, 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 at all from that of the wild rabbit. In
the Porto Santo feral rabbits the supra-orbital plates are
generally narrower and more pointed than in our wild
rabbits.

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

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

Fig. 12—Atlas Vertebræ of Rabbits.

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

Fig. 13—Third Cervical Vertebræ, of natural size,
of—A. Wild Rabbit; B. Hare-coloured, large, Lop-eared Rabbit.

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.

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.

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.

Ninth and tenth dorsal vertebræ.—In the wild rabbit the neural
spine of the ninth vertebra is just perceptibly thicker than that of the
eighth; and the neural spine of the tenth is plainly thicker and shorter
than those of all the anterior vertebræ. In the large lop-eared rabbits the
neural spines of the tenth, ninth, and eighth vertebræ, and even in a
slight degree that of the seventh, 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 from the common wild rabbit, in a direction exactly
opposite to that assumed by the large lop-eared rabbits, the neural spines
of the ninth and tenth vertebræ were not at all larger than those of the
several anterior vertebra. In this same Porto Santo specimen there was no
trace in the ninth vertebra of the anterior lateral processes (see fig.
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,^[26] a haemal spine was moderately well
developed on the under side of the twelfth dorsal vertebra, and I have seen
this in no other specimen.

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

Fig. 15—Terminal bone of Sternum of Rabbits.

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

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

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

Scapula.—The
acromion sends out a rectangular bar, ending in an oblique knob,
which latter in the wild rabbit (fig. 16, A) varies a little in
shape and size, as does the apex of the acromion in sharpness, and
the part just below the rectangular bar in breadth. But the
variations in these respects in the wild rabbit are very slight:
whilst in the large lop-eared rabbits they are considerable. Thus
in some specimens (B) the oblique terminal knob is developed into a
short bar, forming an obtuse angle with the rectangular bar. In
another specimen (C) these two unequal bars form nearly a straight
line. The apex of the acromion varies much in breadth and
sharpness, as may be seen by comparing figures 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 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 length 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 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 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 rabbits. If we
had possessed a large number of domesticated rabbits of nearly the same
size with the wild rabbits, 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.^[27] Now,
after weighing the bodies of four wild rabbits, and of four large but not
fattened lop-eared rabbits, I find that on an average the wild are to the
lop-eared in weight as 1 to 2·17; in average length of body as 1 to
1·41; whilst in capacity of skull they are 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 Table 3 I
have given the measurements of the skull 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 observations
were made. The fourth column gives the capacity of the skull by the
weight of small shot with which the skulls were 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 grains 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 Table 3: the first
four wild rabbits have skulls of the same length, and these differ
but little in capacity. The Sandon rabbit (No. 4) is interesting,
as, though now wild, it is known to be descended from a
domesticated breed, as is still shown by its peculiar colouring and
longer body; nevertheless the skull has recovered its normal length
and full capacity. The next three rabbits are wild, but of small
size, and they all have skulls with slightly lessened capacities.
The three Porto Santo feral rabbits (Nos. 8 to 10) offer a
perplexing case; their bodies are greatly reduced in size, as in a
lesser degree are their skulls in length and in actual capacity, in
comparison with the skulls of wild English rabbits. But when we
compare the capacities of the skull in the three Porto Santo
rabbits, we observe a surprising difference, which does not stand
in any relation to the slight difference in the length of their
skulls, nor, as I believe, to any difference in the size of their
bodies; but I neglected weighing 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 Table 3, 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 the 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 ·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-eared 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-eared skulls, that the latter
would have to be increased 21 per cent to come up to the standard of the
hare.^[28]

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

From the several facts above
given,—namely, firstly, that the actual capacity of the skull
in the Himalayan, Moscow, and Angora breeds, is less than in the
wild rabbit, though they are in all their dimensions rather larger
animals; secondly, that the capacity of the skull of the large
lop-eared rabbits has not been increased in nearly the same ratio
as the capacity of the skull of the smaller wild rabbits has been
decreased; and thirdly, that the capacity of the skull in these
same large lop-eared rabbits is very inferior to that of the hare,
an animal of nearly the same size,—I conclude,
notwithstanding the remarkable differences in capacity in the
skulls of the small Porto 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 organisation 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 taken together have increased in weight, in due proportion
with the increased weight of body, but the hind legs have increased
less than the front legs; 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
has assumed characters proper to the fourth cervical vertebra; 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 process of the
frontal bones and the free end of the malar bones have increased in
breadth; and in the larger breeds the occipital foramen is
generally much less deeply notched than in wild rabbits. Certain
parts of the scapula and the terminal sternal bones have become
highly variable in shape. The ears have been increased enormously
in length and breadth through continued selection; their weight,
conjoined probably with the disuse of their muscles, has caused
them to lop downwards; and this has affected the position and form
of the bony auditory meatus; and this again, by correlation, the
position in a slight degree of almost every bone in the upper part
of the skull, and even the position of the condyles of the lower
jaw.

REFERENCES

[1]
M. P. Gervais, ‘Hist. Nat. des Mammifères,’ 1854, tom. i., p. 288.

[2]
U. Aldrovandi ‘De Quadrupedibus digitatis,’ 1637, p. 383. For Confucius and G.
Markham see a writer who has studied the subject in ‘Cottage Gardener,’
Jan. 22, 1861, p. 250.

[3]
Owen, ‘British Fossil Mammals,’ p. 212.

[4]
Bechstein, ‘Naturgesch. Deutschlands,’ 1801, B. i. p. 1133. I have received
similar accounts with respect to England and Scotland.

[5]
‘Pigeons and Rabbits,’ by E. S. Delamer, 1854, p. 133. Sir J. Sebright
(‘Observations on Instinct,’ 1836, p. 10.) speaks most strongly on the
difficulty. But this difficulty is not invariable, as I have received two
accounts of perfect success in taming and breeding from the wild rabbit. See
also Dr. P. Broca in ‘Journal de la Physiologie,’ tom. ii. p. 368.