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The Variation of Animals and Plants under Domestication by Charles Darwin

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This etext was prepared by Sue Asscher asschers@dingoblue.net.au

THE VARIATION OF

ANIMALS AND PLANTS

UNDER DOMESTICATION

BY

CHARLES DARWIN, M.A., F.R.S., ETC.

IN TWO VOLUMES

VOLUME I.

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 re-
modelled; 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.

CONTENTS.

INTRODUCTION.

CHAPTER 1.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.

CHAPTER 1.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.

CHAPTER 1.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.

CHAPTER 1.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--VERTEBRAE--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.

CHAPTER 1.V.

DOMESTIC PIGEONS.

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

CHAPTER 1.VI.

PIGEONS--continued.

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

CHAPTER 1.VII.

FOWLS.

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

CHAPTER 1.VIII.

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

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

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

PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.
TURKEY, BREEDS OF--CROSSED WITH THE UNITED STATES SPECIES--EFFECTS OF
CLIMATE ON.

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

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

CHAPTER 1.IX.

CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.

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

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

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

CHAPTER 1.X.

PLANTS continued--FRUITS--ORNAMENTAL TREES--FLOWERS.

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

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

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

CHAPTER 1.XI.

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

BUD-VARIATION IN THE PEACH, PLUM, CHERRY, VINE, GOOSEBERRY, CURRANT, AND
BANANA, AS SHOWN BY THE MODIFIED FRUIT--IN FLOWERS: CAMELLIAS, AZALEAS,
CHRYSANTHEMUMS, ROSES, ETC.--ON THE RUNNING OF THE COLOUR IN CARNATIONS
--BUD-VARIATIONS IN LEAVES--VARIATIONS BY SUCKERS, TUBERS, AND BULBS--ON
THE BREAKING OF TULIPS--BUD-VARIATIONS GRADUATE INTO CHANGES CONSEQUENT ON
CHANGED CONDITIONS OF LIFE--GRAFT-HYBRIDS--ON THE SEGREGATION OF THE
PARENTAL CHARACTERS IN SEMINAL HYBRIDS BY BUD-VARIATION--ON THE DIRECT OR
IMMEDIATE ACTION OF FOREIGN POLLEN ON THE MOTHER-PLANT--ON THE EFFECTS OF A
PREVIOUS IMPREGNATION ON THE SUBSEQUENT OFFSPRING OF FEMALE ANIMALS--
CONCLUSION AND SUMMARY.

CHAPTER 1.XII.

INHERITANCE.

WONDERFUL NATURE OF INHERITANCE--PEDIGREES OF OUR DOMESTICATED ANIMALS--
INHERITANCE NOT DUE TO CHANCE--TRIFLING CHARACTERS INHERITED--DISEASES
INHERITED--PECULIARITIES IN THE EYE INHERITED--DISEASES IN THE HORSE--
LONGEVITY AND VIGOUR--ASYMMETRICAL DEVIATIONS OF STRUCTURE--POLYDACTYLISM
AND REGROWTH OF SUPERNUMERARY DIGITS AFTER AMPUTATION--CASES OF SEVERAL
CHILDREN SIMILARLY AFFECTED FROM NON-AFFECTED PARENTS--WEAK AND FLUCTUATING
INHERITANCE: IN WEEPING TREES, IN DWARFNESS, COLOUR OF FRUIT AND FLOWERS--
COLOUR OF HORSES--NON-INHERITANCE IN CERTAIN CASES--INHERITANCE OF
STRUCTURE AND HABITS OVERBORNE BY HOSTILE CONDITIONS OF LIFE, BY
INCESSANTLY RECURRING VARIABILITY, AND BY REVERSION--CONCLUSION.

LIST OF ILLUSTRATIONS.

FIGURE.

1. DUN DEVONSHIRE PONY, WITH SHOULDER, SPINAL, AND LEG STRIPES.

2. HEAD OF JAPAN OR MASKED PIG.

3. HEAD OF WILD BOAR, AND OF "GOLDEN DAYS," A PIG OF THE YORKSHIRE LARGE
BREED.

4. OLD IRISH PIG WITH JAW-APPENDAGES.

5. HALF-LOP RABBIT.

6. SKULL OF WILD RABBIT.

7. SKULL OF LARGE LOP-EARED RABBIT.

8. PART OF ZYGOMATIC ARCH, SHOWING THE PROJECTING END OF THE MALAR BONE OF
THE AUDITORY MEATUS, OF RABBITS.

9. POSTERIOR END OF SKULL, SHOWING THE INTER-PARIETAL BONE, OF RABBITS.

10. OCCIPITAL FORAMEN OF RABBITS.

11. SKULL OF HALF-LOP RABBIT.

12. ATLAS VERTEBRAE OF RABBITS.

13. THIRD CERVICAL VERTEBRAE OF RABBITS.

14. DORSAL VERTEBRAE, FROM SIXTH TO TENTH INCLUSIVE, OF RABBITS.

15. TERMINAL BONE OF STERNUM OF RABBITS.

16. ACROMION OF SCAPULA OF RABBITS.

17. THE ROCK-PIGEON, OR COLUMBA LIVIA.

18. ENGLISH POUTER.

19. ENGLISH CARRIER.

20. ENGLISH BARB.

21. ENGLISH FANTAIL.

22. AFRICAN OWL.

23. SHORT-FACED ENGLISH TUMBLER.

24. SKULLS OF PIGEONS, VIEWED LATERALLY.

25. LOWER JAWS OF PIGEONS, SEEN FROM ABOVE.

26. SKULL OF RUNT, SEEN FROM ABOVE.

27. LATERAL VIEW OF JAWS OF PIGEONS.

28. SCAPULAE OF PIGEONS.

29. FURCULA OF PIGEONS.

30. SPANISH FOWL.

31. HAMBURGH FOWL.

32. POLISH FOWL.

33. OCCIPITAL FORAMEN OF THE SKULLS OF FOWLS.

34. SKULLS OF FOWLS, VIEWED FROM ABOVE, A LITTLE OBLIQUELY.

35. LONGITUDINAL SECTIONS OF SKULLS OF FOWLS, VIEWED LATERALLY.

36. SKULL OF HORNED FOWL, VIEWED FROM ABOVE, A LITTLE OBLIQUELY.

37. SIXTH CERVICAL VERTEBRAE OF FOWLS, VIEWED LATERALLY.

38. EXTREMITY OF THE FURCULA OF FOWLS, VIEWED LATERALLY.

39. SKULLS OF DUCKS, VIEWED LATERALLY, REDUCED TO TWO-THIRDS OF THE
NATURAL SIZE.

40. CERVICAL VERTEBRAE OF DUCKS, OF NATURAL SIZE.

41. PODS OF THE COMMON PEA.

42. PEACH AND ALMOND STONES, OF NATURAL SIZE, VIEWED EDGEWAYS.

43. PLUM STONES, OF NATURAL SIZE, VIEWED LATERALLY.

TABLE 1: PRINCIPAL ADDITIONS AND CORRECTIONS IN THIS (SECOND) EDITION.

First Edition, Volume I., Page 34.
Second Edition Volume I., Page 35.
Dr. Burt Wilder's observations on the brains of different breeds of the
Dog.

First Edition, Volume I., Page 38.
Second Edition Volume I., Page 40.
Degeneracy of Dogs imported into Guinea.

First Edition, Volume I., Page 51.
Second Edition Volume I., Page 54.
Difference in the number of the lumbar vertebrae in the races or species of
the Horse.

First Edition, Volume I., Page 102.
Second Edition Volume I., Page 106.
Hairy appendages to the throats of Goats.

First Edition, Volume I., Page 162.
Second Edition Volume I., Page 170.
Sexual differences in colour in the domestic Pigeon.

First Edition, Volume I., Page 217.
Second Edition Volume I., Page 228.
Movements like those of the Tumbler-pigeon, caused by injury to the brain.

First Edition, Volume I., Page 290.
Second Edition Volume I., Page 306.
Additional facts with respect to the Black-shouldered Peacock.

First Edition, Volume I., Page 296.
Second Edition Volume I., Page 312.
Ancient selection of Gold-fish in China.

First Edition, Volume I., Page 314.
Second Edition Volume I., Page 332.
Major Hallett's 'Pedigree Wheat.'

First Edition, Volume I., Page 326.
Second Edition Volume I., Page 345.
The common radish descended from Raphanus raphanistrum.

First Edition, Volume I., Page 374.
Second Edition Volume I., Page 398.
Several additional cases of bud-variation given.

First Edition, Volume I., Page 396.
Second Edition Volume I., Page 420.
An abstract of all the cases recently published of graft-hybrids in the
potato, together with a general summary on graft-hybridisation.

First Edition, Volume I., Page 399.
Second Edition Volume I., Page 429.
An erroneous statement with respect to the pollen of the date-palm
affecting the fruit of the Chamaerops omitted.

First Edition, Volume I., Page 400.
Second Edition Volume I., Page 430.
New cases of the direct action of pollen on the mother-plant.

First Edition, Volume I., Page 404.
Second Edition Volume I., Page 435.
Additional and remarkable instances of the action of the male parent on the
future progeny of the female.

First Edition, Volume II., Page 14.
Second Edition Volume I., Page 459.
An erroneous statement corrected, with respect to the regrowth of
supernumerary digits after amputation.

First Edition, Volume II., Page 23.
Second Edition Volume I., Page 467.
Additional facts with respect to the inherited effects of circumcision.

First Edition, Volume II., Page 23.
Second Edition Volume I., Page 467.
Dr. Brown-Sequard on the inherited effects of operations on the Guinea-pig.

First Edition, Volume II., Page 24.
Second Edition Volume I., Page 469.
Other cases of inherited mutilations.

First Edition, Volume II., Page 43.
Second Edition Volume II., Page 17.
An additional case of reversion due to a cross.

First Edition, Volume II., Page 72.
Second Edition Volume II., Page 48.
Inheritance as limited by sex.

First Edition, Volume II., Page 105.
Second Edition Volume II., Page 83.
Two varieties of maize which cannot be crossed.

First Edition, Volume II., Page 120.
Second Edition Volume II., Page 99.
Some additional facts on the advantages of cross-breeding in animals.

First Edition, Volume II., Page 123.
Second Edition Volume II., Page 103.
Discussion on the effects of close interbreeding in the case of man.

First Edition, Volume II., Page 135 to 141.
Second Edition Volume II., Page 117 to 122.
Additional cases of plants sterile with pollen from the same plant.

First Edition, Volume II., Page 149.
Second Edition Volume II., Page 131.
Mr. Sclater on the infertility of animals under confinement.

First Edition, Volume II., Page 152.
Second Edition Volume II., Page 134.
The Aperea a distinct species from the Guinea-pig.

First Edition, Volume II., Page 230.
Second Edition Volume II., Page 215.
Professor Jager on hawks killing light-coloured pigeons.

First Edition, Volume II., Page 273.
Second Edition Volume II., Page 262.
Professor Weismann on the effects of isolation in the development of
species.

First Edition, Volume II., Page 281.
Second Edition Volume II., Page 271.
The direct action of the conditions of life in causing variation.

First Edition, Volume II., Page 317.
Second Edition Volume II., Page 309.
Mr. Romanes on rudimentary parts.

First Edition, Volume II., Page 324 to 328.
Second Edition Volume II., Page 316 to 327.
Some additional cases of correlated variability.

First Edition, Volume II., Page 339.
Second Edition Volume II., Page 333.
On Geoffroy St.-Hilaire's law of "soi pour soi."

First Edition, Volume II., Page 357 to 404.
Second Edition Volume II., Page 349 to 399.
The chapter on Pangenesis has been largely altered and re-modelled; but the
essential principles remain the same.

THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION.

INTRODUCTION.

The object of this work is not to describe all the many races of animals
which have been domesticated by man, and of the plants which have been
cultivated by him; even if I possessed the requisite knowledge, so gigantic
an undertaking would be here superfluous. It is my intention to give under
the head of each species only such facts as I have been able to collect or
observe, showing the amount and nature of the changes which animals and
plants have undergone whilst under man's dominion, or which bear on the
general principles of variation. In one case alone, namely in that of the
domestic pigeon, I will describe fully all the chief races, their history,
the amount and nature of their differences, and the probable steps by which
they have been formed. I have selected this case, because, as we shall
hereafter see, the materials are better than in any other; and one case
fully described will in fact illustrate all others. But I shall also
describe domesticated rabbits, fowls, and ducks, with considerable 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.
(Here shown with [].) 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. (Introduction/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.) 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. (Introduction/2. M. Pouchet has recently
('Plurality of Races' English Translation 1864 page 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.) 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
(Introduction/3. Leon Dufour in 'Annales des Science. Nat.' (3rd series,
Zoolog.) tome 5 page 6.)) 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. (Introduction/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 1.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. Palaeontology (1/1. Owen 'British Fossil Mammals'
pages 123 to 133. Pictet 'Traite de Pal.' 1853 tome 1 page 202. De
Blainville in his 'Osteographie, Canidae' page 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 page 131 etc.
and his other publications. Jeitteles has discussed in great detail the
character of the breeds of pre-historic dogs: 'Die vorgeschichtlichen
Alterthumer der Stadt Olmutz' II. Theil, 1872 page 44 to end.) 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. (1/2. Pallas, I
believe, originated this doctrine in 'Act. Acad. St. Petersburgh' 1780 Part
2. Ehrenberg has advocated it, as may be seen in De Blainville's
'Osteographie' page 79. It has been carried to an extreme extent by Col.
Hamilton Smith in the 'Naturalist Library' volumes 9 and 10. 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 page 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. Gen.' 1860 tome 3 page 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 tome 2 page
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.") 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. (1/3. 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 'Beitrage zur Kulturgeschichte' Leipzig 1852 s. 115, 'Youatt
on the Dog' 1845 page 6. A very full history is given by De Blainville in
his 'Osteographie, Canidae.') 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 (1/4. 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 page 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.) 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 (1/5. 'Proc. Zoolog. Soc.' July 12, 1831.) 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 (1/6. 'Sporting in Algeria' page 51.) 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. (1/7. 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 page 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.) 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. (1/8. These, and the following facts on the
Danish remains, are taken from M. Morlot's most interesting memoir in 'Soc.
Vaudoise des Sc. Nat.' tome 6 1860 pages 281, 299, 320.) 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. Rutimeyer (1/9. 'Die
Fauna der Pfahlbauten' 1861 s. 117, 162.), 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). Rutimeyer
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 (1/10. De Blainville 'Osteographie, Canidae.'); 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
Rutimeyer, 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. (1/11. Sir R. Schomburgk has
given me information on this head. See also 'Journal of R. Geographical
Soc.' volume 13 1843 page 65.)

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 (1/12. 'Domestication of Animals' Ethnological Soc.
December 22, 1863.) 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 Canidae 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 (1/13. 'Journal of Researches' etc. 1845 page
393. With respect to Canis antarcticus, see page 193. For the case of the
antelope, see 'Journal Royal Geographical Soc.' volume 23 page 94.) 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." (1/14. The authorities for the foregoing statements are as
follow:--Richardson in 'Fauna Boreali-Americana' 1829 pages 64, 75; Dr.
Kane 'Arctic Explorations' 1856 volume 1 pages 398, 455; Dr. Hayes 'Arctic
Boat Journey' 1860 page 167. Franklin's 'Narrative' volume 1 page 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 volume 2 page 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.' tome 7 1860 page 148, gives a good account of the half-
bred Esquimaux dogs.)

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. (1/15. 'Fauna
Boreali-Americana' 1829 pages 73, 78, 80. Nott and Gliddon, 'Types of
Mankind' page 383. The naturalist and traveller Bartram is quoted by
Hamilton Smith in 'Naturalist Lib.' volume 10 page 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 volume 2 page 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.)

Turning to the southern parts of the new world, Columbus found two kinds of
dogs in the West Indies; and Fernandez (1/16. 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 page 329.) 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 (1/17. 'Naturgeschichte der Saugethiere von Paraguay' 1830 s.
151.), 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 (1/18. Quoted in Humboldt 'Aspects of Nature'
(English translation) volume 1 page 108.) 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 ingae, 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 Canidae.

Turning to the Old World, some European dogs closely resemble the wolf;
thus the shepherd dog of the plains of Hungary is white or reddish-brown,
has a sharp nose, short, erect ears, shaggy coat, and bushy tail, and so
much resembles a wolf that Mr. Paget, who gives this description, says he
has known a Hungarian mistake a wolf for one of his own dogs. Jeitteles,
also, remarks on the close similarity of the Hungarian dog and wolf.
Shepherd dogs in Italy must anciently have closely resembled wolves, for
Columella (vii. 12) advises that white dogs be kept, adding, "pastor album
probat, ne pro lupo canem feriat." Several accounts have been given of dogs
and wolves crossing naturally; and Pliny asserts that the Gauls tied their
female dogs in the woods that they might cross with wolves. (1/19. Paget
'Travels in Hungary and Transylvania' volume 1 page 501. Jeitteles 'Fauna
Hungariae Superioris' 1862 s. 13. See Pliny 'History of the World' (English
translation) 8th book ch. 40 about the Gauls crossing their dogs. See also
Aristotle 'Hist. Animal.' Lib. 8 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 2 page 94.) 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. (1/20. I give this on excellent
authority, namely Mr. Blyth (under the signature of Zoophilus) in the
'Indian Sporting Review' October 1856 page 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.)

With respect to Jackals, Isidore Geoffroy Saint-Hilaire (1/21. For numerous
and interesting details on the resemblance of dogs and jackals see Isid.
Geoffroy St.-Hilaire 'Hist. Nat. Gen.' 1860 tome 3 page 101. See also
'Hist. Nat. des Mammiferes' par Prof. Gervais, 1855 tome 2 page 60.) 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. (1/22. Also Guldenstadt 'Nov. Comment. Acad. Petrop.' tome 20 pro
anno 1775 page 449. Also Salvin in 'Land and Water' October 1869.) A number
of excellent naturalists, from the time of Guldenstadt 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 etonnamment a des chacals." Ehrenberg (1/23. Quoted
by De Blainville in his 'Osteographie, Canidae' pages 79, 98.) 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.
(1/24. See Pallas in 'Act. Acad. St. Petersburgh' 1780 part 2 page 91. For
Algeria, see Isid. Geoffroy St.-Hilaire 'Hist. Nat. Gen.' tome 3 page 177.
In both countries it is the male jackal which pairs with female domestic
dogs.) 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. (1/25. John Barbut 'Description of the Coast of
Guinea in 1746.') On the east coast of Africa, between latitude 4 deg and 6
deg 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 (1/26. 'Travels in
South Africa' volume 2 page 272.) 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. (1/27. Selwyn, Geology of Victoria;
'Journal of Geolog. Soc.' volume 14 1858 page 536 and volume 16 1860 page
148; and Prof. M'Coy in 'Annals and Mag. of Nat. Hist.' (3rd series) volume
9 1862 page 147. The Dingo differs from the dogs of the central Polynesian
islands. Dieffenbach remarks ('Travels' volume 2 page 45) that the native
New Zealand dog also differs from the Dingo.)

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 Canidae 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. (1/28. 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
volume 2 page 295.)

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 primaevus of India were tamed by Mr. Hodgson (1/29. 'Proceedings
Zoological Soc.' 1833 page 112. See also on the taming of the common wolf,
L. Lloyd 'Scandinavian Adventures' 1854 volume 1 page 460. With respect to
the jackal, see Prof. Gervais 'Hist. Nat. Mamm.' tome 2 page 61. With
respect to the aguara of Paraguay see Rengger's work.), 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
(130. Roulin, in 'Mem. present. par divers Savans' tome 6 page 341.) 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 (1/31. Martin 'History of the Dog' page 14.) "made his voice sound
as loudly as any other dog of the same age and size." According to
Professor Nillson (1/32. Quoted by L. Lloyd in 'Field Sports of North of
Europe' volume 1 page 387.), 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. (1/33. Quatrefages 'Soc. d'Acclimat.' May 11, 1863 page
7.) An interesting account has been given by Mr. G. Clarke (1/34. 'Annals
and Mag of Nat. Hist.' volume 15 1845 page 140.) 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. (1/35. Azara 'Voyages dans l'Amer. Merid.' tome 1 page 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 page 3.) 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. (1/36. With respect
to wolves burrowing holes see Richardson 'Fauna Boreali-Americana' page 64;
and Bechstein 'Naturgeschichte Deutschlands' b. 1 s. 617.) These feral dogs
have not become uniform in colour on Juan Fernandez, Juan de Nova, or La
Plata. (1/37. See Poeppig 'Reise in Chile' b. 1 s. 290; Mr. G. Clarke, as
above; and Rengger, s. 155.) 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 (1/38. Dogs, 'Nat. Library' volume 10
page 121; an endemic South American dog seems also to have become feral in
this island. See Gosse 'Jamaica' page 340.) 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 (1/39. Low
'Domesticated Animals' page 650.) 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 (1/40. 'The Naturalist
Library' Dogs, volume 10 pages 4, 19.) 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 maniere a
rappeler la tache terminale de meme couleur, qui caracterise la plupart des
Canides sauvages." (1/41. Quoted by Prof. Gervais 'Hist. Nat. Mamm.' tome 2
page 66.) 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. (1/42. 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 page 353). Hunter found that the gestation of a
mongrel from wolf and dog ('Phil. Transact.' 1789 page 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.' tome 4 page 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. Gen.' tome 3 page 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.) 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 (1/43. See Isid. Geoffroy St.-Hilaire 'Hist. Nat. Gen.' tome 3 page
112, on the odour of jackals. Col. Ham. Smith in 'Nat. Lib.' volume 10 page
289.); 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 (1/44. Quoted by Quatrefages
in 'Bull. Soc. d'Acclimat.' May 11, 1863.) 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
Canidae, 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
(1/45. 'Journal de la Physiologie' tome 2 page 385.), 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
(1/46. See Mr. R. Hill's excellent account of this breed in Gosse's
'Jamaica' page 338; Rengger 'Saugethiere von Paraguay' s. 153. With respect
to Spitz dogs, see Bechstein's 'Naturgesch. Deutschlands' 1801 b. 1 s. 638.
With respect to Dr. Hodgkin's statement made before Brit. Assoc. see 'The
Zoologist' volume 4 for 1845-46 page 1097.) 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 (1/47. 'Acta Acad. St. Petersburgh' 1780
part 2 pages 84, 100.) 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 Canidae 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. (1/48. M. Broca has shown ('Journal de Physiologie' tome 2 page
353) that Buffon's experiments have been often misrepresented. Broca has
collected (pages 390-395) many facts on the fertility of crossed dogs,
wolves, and jackals.) 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. (1/49. 'De la Longevite
Humaine' par M. Flourens 1855 page 143. Mr. Blyth says ('Indian Sporting
Review' volume 2 page 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. Gen.' tome 3 page 217, who speaks of the
hybrid offspring of the jackal as perfectly fertile for three generations.)
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. (1/50. On authority of F. Cuvier quoted in Bronn's 'Geschichte der
Natur' b. 2 s. 164.) Some hounds from Central Africa, brought home by Major
Denham, never bred in the Town of London (1/51. W.C.L. Martin 'History of
the Dog' 1845 page 203. Mr. Philip P. King, after ample opportunities of
observation, informs me that the Dingo and European dogs often cross in
Australia.); 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 Canidae; 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
(1/52. Ruppel 'Neue Wirbelthiere von Abyssinien' 1835-40 'Mammif.' s. 39
pl. 14. There is a specimen of this fine animal in the British Museum.),
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 (1/53. Even
Pallas admits this; see 'Act. Acad. St. Petersburgh' 1780 page 93.) 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 Canidae 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 (1/54. Quoted
by I. Geoffroy 'Hist. Nat. Gen.' tome 3 page 453.) that in form the
differences are "plus fortes que celles d'aucunes especes sauvages d'un
meme 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.
(1/55. F. Cuvier in 'Annales du Museum' tome 18 page 337; Godron 'De
l'Espece' tome 1 page 342; and Col. H. Smith in 'Nat. Library' volume 9
page 101. See also some observations on the degeneracy of the skull in
certain breeds, by Prof. Bianconi 'La Theorie Darwinienne' 1874 page 279.)
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." (1/56. Dr. Burt Wilder 'American Assoc. Advancement of
Science' 1873 pages 236, 239.) 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 (1/57. Isid.
Geoffroy Saint-Hilaire 'Hist. des Anomalies' 1832 tome 1 page 660, Gervais
'Hist. Nat. des Mammiferes' tome 2 1855 page 66. De Blainville
('Osteographie, Canidae' page 137) has also seen an extra molar on both
sides.); and Professor Gervais says that there are dogs "qui ont sept
paires de dents superieures et huit inferieures." De Blainville (1/58.
'Osteographie, Canidae' page 137.) 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. Muller (1/59. Wurzburger 'Medecin.
Zeitschrift' 1860 b. 1 s. 265.), 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 (1/60. Mr. Yarrell in 'Proc. Zoological Soc.' October 8, 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.),--sometimes having
none except one molar on each side; but this, though characteristic of the
breed, must be considered as a monstrosity. M. Girard (1/61. Quoted in 'The
Veterinary' London volume 8 page 415.), 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." (1/62. This is quoted from Stonehenge, a great authority,
'The Dog' 1867 page 187.)

With respect to minor differences little need be said. Isidore Geoffroy has
shown (1/63. 'Hist. Nat. General' tome 3 page 448.) 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. (1/64. W. Scrope
'Art of Deer-Stalking' page 354.) 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. (1/65.
Quoted by Col. Ham. Smith in 'Nat. Lib.' volume 10 page 79.) 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 (1/66. De Blainville 'Osteographie,
Canidae' page 134. F. Cuvier 'Annales du Museum' tome 18 page 342. In
regard to mastiffs, see Col. H. Smith 'Nat. Lib.' volume 10 page 218. For
the Thibet mastiff, see Mr. Hodgson in 'Journal of As. Soc. of Bengal'
volume 1 1832 page 342.) 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 (1/67. 'The Dog' 1845
page 186. With respect to diseases Youatt asserts (page 167) that the
Italian greyhound is "strongly subject" to polypi in the matrix or vagina.
The spaniel and pug (page 182) are most liable to bronchocele. The
liability to distemper (page 232) is extremely different in different
breeds. On the distemper, see also Col. Hutchinson on 'Dog Breaking' 1850
page 279.) "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. (1/68. See 'Youatt on the
Dog' page 15; 'The Veterinary' London volume 11 page 235.) The Rev R.
Everest (1/69. 'Journal of As. Soc. of Bengal' volume 3 page 19.) believes
that no one has succeeded in keeping the Newfoundland dog long alive in
India; so it is, according to Lichtenstein (1/70. 'Travels' volume 2 page
15.), even at the Cape of Good Hope. The Thibet mastiff degenerates on the
plains of India, and can live only on the mountains. (1/71. Hodgson in
'Journal of As. Soc. of Bengal' volume 1 page 342.) Lloyd (1/72. 'Field
Sports of the North of Europe' volume 2 page 165.) 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 Canidae 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 (1/73. 'Hist. Nat. des Mammif.' 1855 tome 2
pages 66, 67.), admits "si l'on prenait sans controle les alterations dont
chacun de ces organes est susceptible, on pourrait croire qu'il y a entre
les chiens domestiques des differences plus grandes que celles qui separent
ailleurs les especes, quelquefois meme 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 (1/74. 'History of Quadrupeds' 1793
volume 1 page 238.) 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 (1/75. 'Oriental Field Sports' quoted by
Youatt 'The Dog' page 15.), 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." (1/76. A. Murray gives this passage in his 'Geographical
Distribution of Mammals' 4to 1866 page 8.) 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 (1/77. Quoted by Mr. Galton 'Domestication of Animals'
page 13.), 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 (1/78.
'Hist. Nat. Gen.' tome 3 page 450.), 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." (1/79. Mr.
Greenhow on the Canadian Dog in Loudon's 'Mag. of Nat. Hist.' volume 6 1833
page 511.) 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. (1/80. See Mr. C.O.
Groom-Napier on the webbing of the hind feet of Otterhounds in 'Land and
Water' October 13, 1866 page 270.) 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
(1/81. 'Fauna Boreali-Americana' 1829 page 62.), 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 (1/82. 'The
Horse in all his Varieties, etc.' 1829 pages 230, 234.), 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 (1/83. 'The
Dog' 1845 pages 31, 35; with respect to King Charles' spaniel page 45; for
the setter page 90.), 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 (1/84. In the 'Encyclop. of Rural
Sports' page 557.) 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

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