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THE VARIATION
OF
ANIMALS AND PLANTS
UNDER DOMESTICATION.
BY
CHARLES DARWIN, M.A., F.R.S., &c.
AUTHORIZED EDITION, WITH A PREFACE
BY
PROFESSOR ASA GRAY.
IN TWO VOLUMES. VOL. I.
With Yllustrations.
NEW-YORK:
ORANGE JUDD & COMPANY,
2 45 BROADWAY.

Entered, according to Act of Congress, in the year 1868, by
ORANGE JUDD & COMPANY,
In the Clerk's Office of the District Court of the United States for the Southern
District of New York.

AUTHOR'S PREFACE TO THE AMERICAN EDITION.
An edition of this book was reprinted immediately
after its first publication, and I thus had the opportu-
nity of inserting various corrections and some important
additions. These are included in the present American
edition, together with some new corrections. It is a
great gratification to me that my work should be thought
worthy of republication in the United States, which con-
tains so large a body of intelligent readers. The details
given in the first volume will probably be too numerous
and minute for most readers; but they appeared to me
worth publishing, as different persons might be interested
in different classes of animals and plants; and the facts
taken together shew in the clearest manner how largely
organic beings vary when subjected to domestication. I
venture to call the reader's attention to the chapter on
Pangenesis. The view there propounded is simply hypo-
thetical, but it has appeared to me, and I have the satis-
faction to know that it has likewise thus appeared to
some capable judges in England, to be no small gain to
seize on a material bond, by which the various forms of
reproduction, inheritance, development, etc., can be con-
nected together. We thus get rid of such vague terms
as spermatic force, the vivification of the ovule, sexual
potentiality, and the diffusion of mysterious essences or
properties from either parent, or from both, to the child.
Whatever may be thought of the conclusions at which I
have arrived on various points, I hope that the student
will find my work of use, as giving to him a larger body
of methodically arranged facts on certain subjects than
can be found, as I believe, in any other work.
CHARLES DARWIN.
Down BROMLEY, KENT, March 28, 1868.

ADDITIONS AND CORRECTIONS.
PUBLISHERS' NOTE TO THE READER.—The first English edition
of this work was taken up at once, and a second called for. In
the reprint Mr. Darwin made a number of changes and correc-
tions, and sent us the advance sheets containing them. He also has
given us a number of manuscript corrections which do not even
appear in the latest English reprint. If the reader will mark the
passages indicated below, he will have his copy revised up to the
author's latest views. These corrections, as well as Mr. Darwin's
preface, were received after the book was printed ; and we were
obliged, by force of circumstances, to insert both in extra pages.
CORRECTIONS TO VOL. I.
Page 86, and wherever Sus Indica occurs, read Sus Indicus.
Page 104, 16 lines from top. Instead of " It has been found in
England associated with the remains of the elephant and rhino-
ceros," read “It apparently did not exist in England before the
Neolithic period, though a greater age was formerly assigned to it."
Page 104, 2d line from bottom. After B. longifrons add, " and
according to Mr. Boyd Dawkins is identical with it."
Page 104, foot-note 40. Strike out reference to Owen, British
Mammalia, and insert “Mr. Boyd Dawkins on the British Fossil
Oxen, Journal of the Geol. Soc. Aug. 1867, p. 182."
Page 127, 2d line from bottom. For “during the early stone
period," read " during the early part of the Neolithic period.”
Page 200, foot-note 35. Strike out that part of the note begin-
ning "but it is stated," and ending with "than in the female. .
Page 223, 12 lines from top. For (Dendrocygna viduata) read
(Anas moschata).
Page 351, 18 lines from top. After word “strain” insert “ Again,
Mr. T. Jenner Wier informs me that a peacock at Blackheath,
whilst young, was white, but as it became older it assumed the cha-
racter of the black-shouldered variety ; both its parents were com-
mon peacocks. Here we have six distinct cases," etc.
Page 352, 9 lines from top. For "as it did to Sir R. Heron, to pre-
ponderate strongly in favour," read "evidence seems to me to be
decisive in favour of the black-shouldered," etc.
Page 355, 8 lines from top. For "by naturalists," read “ by some
naturalists."
Page 475, 19 lines from bottom. Insert, after "no success,
Dr. Hildebrand informs me, in a letter dated Jan. 2d, 1868, that he
has recently succeeded with the potato. He removed all the eyes
from a white, smooth-skinned potato, and all from a red, scaly po-
tato, and inserted them reciprocally into each other. From these
eyes he raised only two plants; and of the tubers formed by them
two were red and scaly at one end, and white and smooth-skinned
at the other; the middle part being white with red streaks. Hence
the possibility of a graft hybrid may be looked at as established."

ADDITIONS AND CORRECTIONS.
11
Mr. Darwin states that this case of Dr. Hildebrand has modified
his belief in the possibility of making a graft hybrid.--PUB.]
Page 475, 18 lines from bottom. After instance known to me,”
insert “(with the exception of the case just given)."
Page 476, 13 lines from bottom. Strike out all of the sentence
after "above described."
Page 480. To foot-note 126 add : " Dr. Hildebrand, of Bonn, in
a letter dated Jan. 2, 1868, informs me that he has recently crossed
yellow and red maize and obtained the same results as Dr. Sovi,
with the important addition, that in one case the axis which sup-
ports the seeds was stained of a brownish color ; Dr. Hildebrand
also gives me some striking cases with respect to the apple-tree,
like those recorded further on. These valuable facts will soon be
recorded in the 'Bot. Zeitung.'
Page 485. Add to foot-note: "Dr. Bowerbank has given us the
following striking case :- A black, hairless Barbary bitch was first
impregnated by a mongrel spaniel with long brown hair, and she
produced five puppies, three of which were hairless and two cov-
ered with short brown hair. The next time she was put to a full
black, hairless Barbary dog ; but the mischief had been implanted
in the mother, and again about half the litter looked like pure
Barbarys, and the other half like the short-haired progeny of the
first father."
Page 486, 5 lines from bottom. For " There is a considerable but
insufficient body of evidence," read " There is sufficient evidence."
CORRECTIONS TO VOL. II.
Page 25. To the paragraph ending with "operated on," add
"since the publication of the first edition of this work, I have re-
ceived an account of another instance of the regrowth of a super-
numerary digit."
Page 32, 5 lines from bottom. For “ 73" insert “ 72."
Page 56, 2d paragraph. After "always produced," add "I hear
from Mr. Blyth that the hybrids from the canary and goldfinch al-
most invariably have streaked feathers on their backs; and this
streaking must be derived from the aboriginal wild canary."
Page 88, 14 lines from top. For“ (Tadnora Ægyptiuca),” read
" (Anser Ægyptiacus)."
Page 170, 6 lines from top. After " P. edulis," insert “ In a third
instance, however, P. quadrangularis fruited freely, when arti-
ficially fertilized with its own pollen.”
Page 184, 5 lines from top. Erase all of the sentence after "tur.
key," and add "and fowl are kept and bred by various remote
tribes."
Page 221, 2d line from bottom. Strike out all the sentence after
"result," and insert "so the same thing occurs with trimorphic
plants, for instance, the mid-styled form of Lythrum salicariu
could be illegitimately fertilized with the greatest ease by pollen
from the longer stamens of the short-styled form, and yielded many
seeds, but the latter form did not yield a single seed when fertilized
by the longer stamens of the mid-styled form.'
Page 341, 342, and wherever " Lucaze Duthiers” occurs, read " La-
caze," and alter the same in the index.

iv
ADDITIONS AND CORRECTIONS.
22
Page 428, 3d line from bottom. Strike out all of the sentence
after "amputation," and insert "how it comes that organic beings
identical in every respect are habitually produced by such widely
different processes as budding and true seminal generation."
Page 431, 11 lines from bottom. After "nature," add "and in
the case of Daphnia, Sir J. Lubbock first showed that ova and
pseud-ova are identical in structure.
Page 431, foot-note 5. For Cecydomyide" read " Cecidomyde."
Page 437, paragraph Graft-hybrids. In the 2d line of the
paragraph, cut out after Cytisus adami and make it read “ Cytisus
adami, it was shown that, after the tissues of two plants," etc. In
the sixth line of paragraph, for "are intimately united," read "have
become intimately united.” In the Sth line, “it is certain,"
should read “it is also certain." The closing sentence of the para-
graph, “Should it ever be proved," etc., is to be modified to read,
The possibility of the production of hybridized buds by the union
of two distinctive vegetative tissues is an important fact, as it shows
us that sexual and asexual production are essentially the same: for
the power," etc.
Page 442, bottom line. For "inserted into the eye,” read “inserted
into the ear of an ox, lived for eight years, and acquired, according to
Prof. Mantegazza, a weight of 396 grammes, and the astonishing
length of 24 centimetres, or about 9 English inches; so that the
head of the ox appeared to bear three horns."
Page 443. For foot-note 22, substitute" Degli innesti animali, etc.
Milano, 1865, p. 51, Tab. 3."
Page 449, 5th line from top. Strike out the sentence beginning
“Nearly similar views," and insert " Views in some respects similar
have been propounded, as I find by other authors."
Page 453, 3d line from bottom. For * 4,872,000," read “ 6,867,
840.
Page 453. Cancel the foot-note 34, as far as the word “Harmer,"
and insert "Mr. F. Buckland carefully calculated, by weighing, the
above number of eggs in a codfish ; see · Land and Water,' 1868, P.
62. In a previous instance he found the number to be 4,872,000."
Page 463, 17 lines from top. Strike out the paragraph be-
ginning "It was shown," and substitute the following: “As by
our hypothesis budding or fission differs from seminal generation
only in the manner in which the gemmules are first aggregated, we
can understand the possibility of the formation of graft-hybrids, and
these graft-hybrids, which combine the character of the two forms
of which the tissues have become united, connect in the closest and
most interesting manner gemmation and sexual reproduction."
Page 518, 16 lines from top. "Anas moschata," add “i., p. 223"
Page 522. After “Bowen, Prof.," insert - BOWERBANK, DR., on
the effects of a first impregnation, i., p. 485.”
Page 530. Dendrocygna viduata, strike out reference to vol. i.
Page 539. After
After “HILDEBRAND, DR.," insert "on graft hybrids
with the potato, i., p. 475. On the influence of the pollen on the
mother plant, i., p. 480."
Page 550. Under * Owen, Prof. R.," strike out reference to Bos
longifrons.
Page 562. For " Sus Indica" read “Sus Indicus."

PREFACE TO THE AMERICAN EDITION.
This work is here reprinted from the English edi-
tion, under an arrangement with the author, upon the
recommendation of the subscriber. It is a perfect trea-
sury of facts relative to domesticated animals and some
of the more important cultivated plants; of the princi-
ples which govern the production, improvement, and
preservation of breeds and races; of the laws of inheri-
tance, upon which all origination of improved varieties
depends; of the ill effects of breeding in-and-in, neces-
sary though this be to the full development and perpetu-
ation of a choice race or breed, and of the good effects of
an occasional cross, by which, rightly managed, a breed
may be invigorated or improved. These and various
kindred subjects are discussed scientifically, with rare
ability, acuteness, and impartiality, by one who has de-
voted most of his life to this class of inquiries, and who
discusses them in a way and style equally interesting
and instructive to the professional naturalist or phy-
siologist and to the general reader. To the intelligent
agriculturist and breeder, these volumes will be especi-
ally valuable, and it is in the interest of such practical
men and amateurs that they are here reprinted.

iy
PREFACE.
The subject is, of course, connected with the theory
which has made Mr. Darwin's name so famous; a
theory which, extending the application and range of
these facts into past ages, regards the present species of
plants and animals as older and stronger-marked varie-
ties, originated under a natural selection of the sorts best
adapted to the circumstances and conditions of each place
and time, in a way which may fairly be compared with
the development of our domesticated animals and plants
under artificial selection and care. It was the study of
domesticated races that suggested the theory. Whether
that stand or fall, the facts in respect to breeds and
races, which are here so faithfully collected and discuss-
ed, are none the worse for having served as the basis
of ingenious speculations; and they have an interest of
their own, irrespective of all such theories, as well as a
practical importance which should commend them to
general attention. The curious speculations toward the
close of the second volume, upon the way in which
peculiarities may be supposed to be transmitted from
parents to offspring, or from grandparents to a second
or later generation and the like, are entirely independent
of the Darwinian theory.
The English edition being quite beyond the reach of
the majority of readers in this country, the publishers
of The American Agriculturist have done well in mak-
ing these volumes generally accessible.
A. GRAY.
CAMBRIDGE, MASS., March, 1868.

CONTENTS OF VOLUME I.
INTRODUCTION
·
Page 11
CHAPTER I.
DOMESTIC DOGS AND CATS.
ANCIENT VARIETIES OF THE DOG RESEMBLANCE OF DOMESTIC DOGS IN
VARIOUS COUNTRIES TO NATIVE CANINE SPECIES - ANIMALS NOT AC-
QUAINTED 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 SEV-
ERAL RACES IN PART DUE TO DESCENT FROM DISTINCT SPECIES DIF-
FERENCES 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 - EXTINC-
TION 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
.. 27
CHAPTER II.
HORSES AND ASSES.
HORSE. —DIFFERENCES IN THE BREEDS — INDIVIDUAL VARIABILITY OF -
DIRECT EFFECTS OF THE CONDITIONS OF LIFE CAN WITHSTAND NUCH
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
66

VI
CONTENTS.
CHAPTER III.
PIGS — CATTLE - SHEEP - GOATS.
PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICA - TORF-
SCHWEIN JAPAN PIG FERTILITY OF CROSSED PIGS - CHANGES IN
THE SKULL OF THE HIGHLY CULTIVATED RACES CONVERGENCE OF
CHARACTER - GESTATION - SOLID-HOOFED SWINE - CURIOUS APPENDA GES
TO THE JAWS - DECREASE IN SIZE OF THE TUSKS - YOUNG PIGS LONGI-
TUDINALLY STRIPED FERAL PIGS CROSSED BREEDS.
CATTLE. -- ZEBU A DISTINCT SPECIES - EUROPEAN CATTLE PROBABLY DE-
SCENDED FROM THREE WILD FORMS - ALL THE RACES NOW FERTILE
TOGETHER BRITISH PARK CATTLE - ON THE COLOUR OF THE ABO-
RIGINAL 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 ..
85
CHAPTER IV.
DOMESTIC RABBITS.
DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RABBIT - - ANCIENT
DOMESTICATION ANCIENT SELECTION LARGE LOP-EARED RABBITS -
VARIOUS BREEDS FLUCTUATING CHARACTERS ORIGIN OF THE HIMA-
LAYAN 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 DIFFER-
ENT SPECIES OF HARES · VERTEBRA 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 ..
.. 130
CHAPTER V.
DOMESTIC PIGEONS.
ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS INDIVIDUAL
VARIABILITY — VARIATIONS OF A REMARKABLE NATURE OSTEOLOGICAL
CHARACTERS: SKULL, LOWER JAW, NUMBER OF VERTEBRÆ CORRELA-
TION OF GROWTH: TONGUE WITH BEAK, EYELIDS AND NOSTRILS WITH

CONTENTS.
vii
WATTLED SKIN – NUMBER OF WING-FEATHERS, AND LENGTH OF WING
COLOUR AND DOWN WEBBED AND FEATHERED FEET - ON THE EF-
FECTS OF DISUSE-LENGTH OF FEET IN CORRELATION WITH LENGTH OF
BEAK —LENGTH OF STERNUM, SCAPULA, AND FURCULA — LENGTH OF
WINGS SUMMARY ON THE POINTS OF DIFFERENCE IN THE SEVERAL
BREEDS
.. 163
CHAPTER VI.
PIGEONS-continued.
ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES -
HABITS OF LIFE — WILD RACES OF THE ROCK-PIGEON DOVECOT PIGEONS
PROOFS OF THE DESCENT OF THE SEVERAL RACES FROM COLUMBA
LIVIA -- FERTILITY OF THE RACES WHEN CROSSED REVERSION TO THE
PLUMAGE OF THE WILD ROCK-PIGEON CIRCUMSTANCES FAVOURABLE TO
THE FORMATION OF THE RACES ANTIQUITY AND HISTORY OF THE
PRINCIPAL RACES - - MANNER OF THEIR FORMATION SELECTION UN-
CONSCIOUS 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 .. 221
CHAPTER VII.
FOWLS.
BRIEF DESCRIPTIONS OF THE CHIEF BREEDS ARGUMENTS IN FAVOUR OF
THEIR DESCENT FROM SEVERAL SPECIES - ARGUMENTS IN FAVOUR OF
ALL THE BREEDS HAVING DESCENDED FROM GALLUS BANKIVA --- REVER-
SION TO THE PARENT-STOCK IN COLOUR - ANALOGOUS VARIATIONS
ANCIENT HISTORY OF THE FOWL EXTERNAL DIFFERENCES BETWEEN
THE SEVERAL BREEDS - EGGS - CHICKENS SECONDARY SEXUAL CHA-
RACTERS WING- AND TAIL- FEATHERS, VOICE, DISPOSITION, ETC.
OSTEOLOGICAL DIFFERENCES IN THE SKULL, VERTEBRÆ, ETC. — EF-
FECTS OF USE AND DISUSE OF CERTAIN PARTS CORRELATION OF
GROWTH
.. 273
CHAPTER VIII.
DUCKS -- GOOSE - PEACOCK - TURKEY - GUINEA-FOWL-
CANARY-BIRD - GOLD-FISH --- HIVE-BEES -- SILK-MOTHS.
DUCKS, SEVERAL BREEDS OF — PROGRESS OF DOMESTICATION ---ORIGIN
OF, FROM THE COMMON WILD-DUCK DIFFERENCES IN THE DIFFERENT

viii
CONTENTS.
BREEDS – OSTEOLOGICAL DIFFERENCES — EFFECTS OF USE AND DIS-
USE ON THE LIMB-BONES.
GOOSE, ANCIENTLY DOMESTICATED — LITTLE VARIATION OF+SEBASTAPOL
BREED.
PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.
TURKEY, BREEDS OF — CROSSED WITH THE UNITED STATES SPECIES
EFFECTS OF CLIMATE ON.
GUINEA-FOWL, CANARY-BIRD, GOLD-FISH, HIVE-BEES.
SILK-MOTHS, SPECIES AND BREEDS OF — ANCIENTLY DOMESTICATED
CARE IN THEIR SELECTION DIFFERENCES IN THE DIFFERENT RACES
IN THE EGG, CATERPILLAR, AND COCOON STATES — INHERITANCE OF
CHARACTERS IMPERFECT WINGS - LOST INSTINCTS — CORRELATED CHA-
RACTERS
333
CHAPTER IX.
CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.
PRELIMINARY REMARKS ON THE NUMBER AND PARENTAGE OF CUL-
TIVATED PLANTS FIRST STEPS IN CULTIVATION GEOGRAPHICAL DIS-
TRIBUTION OF CULTIVATED PLANTS.
CEREALIA.— DOUBTS ON THE NUMBER OF SPECIES. - WHEAT : VARI-
ETIES 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 INTER CROSS. BEANS. — POTATOES :
NUMEROUS VARIETIES OF — DIFFERING LITTLE, EXCEPT IN THE TUBERS
CHARACTERS INHERITED
368
CHAPTER X.
PLANTS continued — FRUITS - ORNAMENTAL TREES — FLOW-
ERS.
FRUITS. - GRAPES - VARY IN ODD AND TRIFLING PARTICULARS.
MULBERRY. • THE ORANGE GROUP - SINGULAR RESULTS FROM CROSS-
ING. PEACH AND NECTARINE BUD-VARIATION - ANALOGOUS VARI-
ATION -- - RELATION TO THE ALMOND. APRICOT. - PLUS — VARI-
ATION IN THEIR STONES. CHERRIES SINGULAR VARIETIES OF.
APPLE.
PEAR. - STRAWBERRY INTERBLENDING OF THE ORIGINAL

CONTENTS.
ix
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 CON-
STITUTIONAL PECULIARITIES - KIND OF VARIATION. · ROSES - SEVE-
RAL SPECIES CULTIVATED. PANSY. DAHLIA. HYACINTH,
HISTORY AND VARIATION OF
400
CHAPTER XI.
ON BUD-VARIATION, AND ON CERTAIN ANOMALOUS MODES
OF REPRODUCTION AND VARIATION.
BUD-VARIATIONS IN THE PEACH, PLUM, CHERRY, VINE, GOOSEBERRY, CUR-
RANT, AND BANANA, AS SHOWN BY THE MODIFIED FRUIT - IN FLOW-
ERS: CAMELLIAS, AZALEAS, CHRYSANTHEMUMS, ROSES, ETC. - ON THE
RUNNING OF THE COLOUR IN CARNATIONS BUD-VARIATIONS IN LEAVES
VARIATIONS BY SUCKERS, TUBERS, AND BULBS — ON THE BREAKING
OF TULIPS BUD-VARIATIONS GRADUATE INTO CHANGES CONSEQUENT ON
CHANGED CONDITIONS OF LIFE - CYTISUS ADAMI, ITS ORIGIN AND TRANS-
FORMATION - ON THE UNION OF TWO DIFFERENT EMBRYOS IN ONE
SEED THE TRIFACIAL ORANGE - ON REVERSION BY BUDS IN HY-
BRIDS AND MONGRELS ON THE PRODUCTION OF MODIFIED BUDS BY
THE GRAFTING OF ONE VARIETY OR SPECIES ON ANOTHER — ON THE
DIRECT OR IMMEDIATE ACTION OF FOREIGN POLLEN ON THE MOTHER-
PLANT — ON THE EFFECTS IN FEMALE ANIMALS OF A FIRST IMPREGNA-
TION ON THE SUBSEQUENT OFFSPRING - CONCLUSION AND SUMMARY 448
LIST OF ILLUSTRATIONS.
..
PAGE
1. DUN DEVONSHIRE PONY, WITH SHOULDER, SPINAL, AND LEG
STRIPES..
.. 75
2. HEAD OF JAPAN OR MASKED PIG..
.. 90
3. HEAD OF WILD BOAR, AND OF “GOLDEN DAYS," A PIG OF
THE YORKSHIRE LARGE BREED
93
4. OLD IRISH PIG, WITH JAW-APPENDAGES
97
5. HALF-LOP RABBIT ..
136
6. SKULL OF WILD RABBIT
. 147
7. SKULL OF LARGE LOP-EARED RABBIT
117
.

Х
CONTENTS.
. 148
.
.. 168
.
..
..
8. PART OF ZYGOMATIC ARCH, SHOWING THE PROJECTING END
OF THE MALAR-BONE, AND THE AUDITORY MEATUS, OF RAB-
BITS
148
9. POSTERIOR END OF SKULL, SHOWING THE INTER-PARIETAL BONE
OF RABBITS
10. OCCIPITAL FORAMEN OF RABBITS
148
11. SKULL OF HALF-LOP RABBIT
149
12. ATLAS VERTEBRÆ OF RABBITS ..
151
13. THIRD CERVICAL VERTEBRÆ OF RABBITS
.. 152
14. DORSAL VERTEBRÆ, FROM SIXTH TO TENTH INCLUSIVE, OF RAB-
BITS
153
15. TERMINAL BONE OF STERNUM OF RABBITS
154
16. ACROMION OF SCAPULA OF RABBITS
. 154
17. THE ROCK-PIGEON, OR COLUMBA LIVIA
18. ENGLISH POUTER
170
19. ENGLISH CARRIER
174
20. ENGLISH BARB ..
. 180
21. ENGLISH FANTAIL
. 182
22. AFRICAN OWL ..
. 185
23. SHORT-FACED ENGLISH TUMBLER
. 188
24. SKULLS OF PIGEONS, VIEWED LATERALLY
. 201
25. LOWER JAWS OF PIGEONS, SEEN FROM ABOVE
202
26. SKULL OF RUNT, SEEN FROM ABOVE
203
27. LATERAL VIEW OF JAWS OF PIGEONS
203
28. SCAPULÆ OF PIGEONS.
206
29. FURCULÆ OF PIGEONS ..
206
30. SPANISH FOWL
274
31. HAMBURGI FOWL ..
. 277
32. POLISII FOWL ..
279
33. OCCIPITAL FORAMEN OF THE SKULLS OF FOWLS
.. 316
34. SKULLS OF FOWLS, VIEWED FROM ABOVE, A LITTLE OBLIQUELY 316
35. LONGITUDINAL SECTIONS OF SKULLS OF FOWLS, VIEWED LAT-
ERALLY .. ..
. 318
36. SKULL OF HORNED FOWL, VIEWED FROM ABOVE, A LITTLE
OBLIQUELY ....
.. 320
37. SIXTH CERVICAL VERTEBRÆ OF FOWLS, VIEWED LATERALLY.. 323
38. EXTREMITY OF THE FURCULA OF FOWLS, VIEWED LATERALLY 324
39. SKULLS OF DUCKS, VIEWED LATERALLY, REDUCED TO TWO
THIRDS OF THE NATURAL SIZE
340
40. CERVICAL VERTEBRÆ OF DUCKS, OF NATURAL SIZE ..
342
41. PODS OF THE COMMON PEA
395
42. PEACH AND ALMOND STONES, OF NATURAL SIZE, VIEWED EDGE-
WAYS
.. 406
43. PLUM STONES, OF NATURAL SIZE, VIEWED LATERALLY
416

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 in-
tention 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 describ-
ed will in fact illustrate all others. But I shall also de-
scribe domesticated rabbits, fowls, and ducks, with con-
siderable.fulness.
The subjects discussed in this volume are so connected
that it is not a little difficult to decide how they can be

12
INTRODUCTION.
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 numerou details, in support of any
proposition or conclusion, small type has been used. The
reader will, I think, find this plan a convenience, for, if
he does not doubt the conclusion or care about the de-
tails, he can easily pass them over; yet I may be permit-
ted 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 spe-
cies. 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 “tam-
pering with nature” and causing variability. If organic
beings had not possessed an inherent tendency to vary,
man could have done nothing. He unintentionally ex-
poses his animals and plants to various conditions of life,
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 publish-
ing this first work has been caused by
continued ill-health.
2 M. Pouchet has recently (Plurality
of Races,' Eng. Translat., 1864, p. 83,
&c.) insisted that variation under do-
mestication throws no light on the natu-
ral modification of species. I cannot
perceive the force of his arguments, or,
to speak more accurately, of his asser-
tions to this effect.

NATURAL SELECTION.
13
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 proba-
bly 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 va-
rieties. 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 inevi-
tably 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 noth-
ing, 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 na-
ture 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 spe-
cies in a state of nature undergo change.
I shall in this volume treat, as fully as my inaterials
permit, the whole subject of variation under domestica-
tion. We may thus hope to obtain some light, little

14
INTRODUCTION.
though it be, on the causes of variability,ấon 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 domes-
ticated organisms are liable. We shall learn something
on the laws of inheritance, on the effects of crossing dif-
ferent breeds, and on that sterility which often supervenes
when organic beings are removed from their natural con-
ditions of life, and likewise when they are too closely
interbred. During this investigation we shall see that
the principle of Selection is all important. Although man
does not cause variability and cannot even prevent it, he
can select, preserve, and accumulate the variations given
to him by the hand of nature in any way which he
chooses; and thus he can certainly produce a great result.
Selection
may be followed either methodically and inten-
tionally, 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 indivi-
duals which he prizes most, and by destroying the worth-
less 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 cha-
racter, as compared with natural species; for they have
been modified not for their own benefit, but for that of
man.
In a second work I shall discuss the variability of or-
ganic beings in a state of nature; namely, the individual

NATURAL SELECTION,
15
races.
differences presented by animals and plants, and those
slightly greater and generally inherited differences which
are ranked by naturalists as varieties or geographical
We shall see how difficult, or rather how impos-
sible 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 in-
clude 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 organi-
zation is in some slight degree plastic; granting that
many animals and plants have varied greatly under do-
mestication, 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 differ-
ences 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 contri-
vances, which have justly excited in the mind of every
observer the highest admiration. There is, for instance,
a fly (Cecidomyia)' which deposits its eggs within the
Léon Dufour in 'Annales des Scienc. Nat.' (3rd series, Zoolog.), tom. v. p. 6.

16
INTRODUCTION.
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 hymenopte-
rous insect depends on a dipterous insect, and this de-
pends 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 cha-
racteristic of varieties into the greater differences charae-
teristic of species and genera, including the admirable
adaptations of each being to its complex organic and in-
organic conditions of life,—will form the main subject of
my
second work. We shall therein see that all organic
beings, without exception, tend to increase at so high a
ratio, that no district, no station, not even the whole sur-
face of the land or the whole ocean, would hold the pro-
geny
of a single pair after a certain number of genera-
tions. The inevitable result is an ever-recurrent Strug-
gle 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 dis-
appeared 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 varia-
tions ever arise in any way advantageous to any being
under its excessively complex and changing relations of
and it would be a strange fact if beneficial varia-
tions 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
life;

NATURAL SELECTION.
17
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 familiar-
ity. 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 varie-
ties 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 at-
traction 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 de-
pends in some manner on the action of the surrounding
circumstances on the organism. I have, also, often per-
sonified the word Nature; for I have found it difficult to
avoid this ambiguity; but I mean by nature only the ag-
gregate action and product of many natural laws,--and
by laws only the ascertained sequence of events.
In the chapter devoted to natural selection I shall
show from experiment and from a multitude of facts,
that the greatest amount of life can be supported on
each spot by great diversification or divergence in the
structure and constitution of its inhabitants. We shall,

18
INTRODUCTION. .
also, see that the continued production of new forms
through natural selection, which implies that each new
variety has some advantage over others, almost inevita-
bly leads to the extermination of the older and less im-
proved forms. These latter are almost necessarily inter-
mediate in structure as well as in descent between the
last-produced forms and their original parent-species.
Now, if we suppose a species to produce two or more
varieties, and these in the course of time to produce
other varieties, the principle of good being derived from
diversification of structure will generally lead to the
preservation of the most divergent varieties; thus the
lesser differences characteristic of varieties come to be
augmented into the greater differences characteristic of
species, and, by the extermination of the older inter
mediate forms, new species come to be distinctly defined
objects. Thus, also, we shall see how it is that organic
beings can be classed by what is called a natural method
in distinct groups-species under genera, and genera
under families.
As all the inhabitants of each country may be said,
owing to their high rate of reproduction, to be striving
to increase in numbers; as each form is related to many
other forms in the struggle for life,—for destroy any one
and its place will be seized by others; as every part of
the organization occasionally varies in some slight de-
gree, and as natural selection acts exclusively by the
preservation of variations which are advantageous under
the excessively complex conditions to which each being
is exposed, no limit exists to the number, singularity, and
perfection of the contrivances and co-adaptations which
may thus be produced. An animal or a plant may thus
slowly become related in its structure and habits in the
most intricate manner to many other animals and plants,
and to the physical conditions of its home. Variations
in the organization will in some cases be aided by habit,
or by the use and disuse of parts, and they will be gov-

NATURAL SELECTION.
-19
erned by the direct action of the surrounding physical
conditions and by correlation of growth.
On the principles here briefly sketched out, there is
no innate or necessary tendency in each being to its own
advancement in the scale of organization. We are almost
compelled to look at the specialization or differentiation
of parts or organs for different functions as the best or
even sole standard of advancement; for by such division
of labour each function of body and mind is better
performed. And, as natural selection acts exclusively
through the preservation of profitable modifications of
structure, and as the conditions of life in each area gen-
erally become more and more complex, from the increas-
ing number of different forms which inhabit it and from
most of these forms acquiring a more and more perfect
structure, we may confidently believe, that, on the whole,
organization advances. Nevertheless a very simple form
fitted for very simple conditions of life might remain for
indefinite ages unaltered or unimproved; for what would
it profit an infusorial animalcule, for instance, or an
intestinal worm, to become highly organized ? Members
of a high group might even become, and this apparently
has occurred, fitted for simpler conditions of life; and
in this case natural selection would tend to simplify or de-
grade the organization, for complicated mechanism for
simple actions would be useless or even disadvanta-
geous.
In a second work, after treating of the Variation of
organisms in a state of nature, of the Struggle for Exist-
ence and the principle of Natural Selection, I shall dis-
cuss the difficulties which are opposed to the theory.
These difficulties may be classed under the following
heads :—the apparent impossibility in some cases of a
very simple organ graduating by small steps into a
highly perfect organ; the marvellous facts of Instinct;
the whole question of Hybridity; and, lastly, the ab-
sence, at the present time and in our geological forma-

20
INTRODUCTION.
tions, 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 inexplica-
ble.
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 hypoth-
esis, 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 inev-
itable 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 homol-
ogies. If the principle of natural selection does explain
these and other large bodies of facts, it ought to be re-
ceived. On the ordinary view of each species having
been independently created, we gain no scientific expla-
nation 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.

NATURAL SELECTION.
21
In a third work I shall try the principle of natural se-
lection by seeing how far it will give a fair explanation
of the several classes of facts just alluded to. It was the
consideration of these facts which first led me to take up
the present subject. When I visited, during the voyage
of H.M.S. Beagle, the Galapagos Archipelago, situated
in the Pacific Ocean about 500 miles from the shore of
South America, I found myself surrounded by peculiar
species of birds, reptiles, and plants, existing nowhere
else in the world. Yet they nearly all bore an American
stamp. In the song of the mocking-thrush, in the harsh
cry of the carrion-hawk, in the great candlestick-like
opuntias, I clearly perceived the neighbourhood of Ame-
rica, though the islands were separated by so many miles
of ocean from the mainland, and differed much from it
in their geological constitution and climate.
Still more
surprising was the fact that most of the inhabitants of
each separate island in this small archipelago were speci-
fically different, though most closely related to each other.
The archipelago, with its innumerable craters and bare
streams of lava, appeared to be of recent origin; and thus
I fancied myself brought near to the very act of creation.
I often asked myself how these many peculiar animals
and plants had been produced: the simplest answer
seemed to be that the inhabitants of the several islands
had descended from each other, undergoing modification
in the course of their descent; and that all the inhabitants
of the archipelago had descended from those of the near-
est land, namely America, whence colonists would na-
turally have been derived. But it long remained to me
an inexplicable problem how the necessary degree of
modification could have been effected, and it would have
thus remained for ever, had I not studied domestic
productions, and thus acquired a just idea of the power
of selection. As soon as I had fully realized this idea,
I saw, on reading Malthus on Population, that Natural
Selection was the inevitable result of the rapid increase

22
INTRODUCTION.
of all organic beings; for I was prepared to appreciate
the struggle for existence by having long studied the
habits of animals.
Before visiting the Galapagos I had collected many
animals whilst travelling from north to south on both
sides of America, and everywhere, under conditions of
life as different as it is possible to conceive, American
forms were met with-species replacing species of the
same peculiar genera. Thus it was when the Cordilleras
were ascended, or the thick tropical forests penetrated, or
the fresh waters of America searched. Subsequently I
visited other countries, which in all the conditions of life
were incomparably more like to parts of South America,
than the different parts of that continent were to each
other; yet in these countries, as in Australia or Southern
Africa, the traveller cannot fail to be struck with the en-
tire difference of their productions. Again the reflection
was forced on me that community of descent from the
early inhabitants or colonists of South America would
alone explain the wide prevalence of American types of
structure throughout that immense area.
To exhume with one's own hands the bones of extinct
and gigantic quadrupeds brings the whole question of the
succession of species vividly before one's mind; and I had
found in South America great pieces of tessellated armour
exactly like, but on a magnificent scale, that corering
the pigmy armadillo; I had found great teeth like those
of the living sloth, and bones like those of the
cavy.
An
analogous succession of allied forms had been previously
observed in Australia. Here then we see the prevalence,
as if by descent, in time as in space, of the same types in
the same areas; and in neither case does the similarity of
the conditions by any means seem sufficient to account for
the similarity of the forms of life. It is notorious that
the fossil remains of closely consecutive formations are
closely allied in structure, and we can at once understand
the fact if they are likewise closely allied by descent.

NATURAL SELECTION.
23
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 spe-
cies come in gradually one by one. Ancient and extinct
forms of life often show combined or intermediate cha-
racters, like the words of a dead language with respect to
its several offshoots or living tongues. All these and
other such facts seemed to me to point to descent with
modification as the method of production of new groups
of species.
The innumerable past and present inhabitants of the
world are connected together by the most singular and
complex affinities, and can be classed in groups under
groups, in the same manner as varieties can be classed
under species and sub-varieties under varieties, but with
much higher grades of difference. It will be seen in my
third work that these complex affinities and the rules for
classification receive a rational explanation on the prin-
ciple of descent, together with modifications acquired
through natural selection, entailing divergence of cha-
racter and the extinction of intermediate forms. How
inexplicable is the similar pattern of the hand of a man,
the foot of a dog, the wing of a bat, the flipper of a
seal, on the doctrine of independent acts of creation !
how simply explained on the principle of the natural
selection of successive slight variations in the diverging
descendants from a single progenitor ! So it is, if we
look to the structure of an individual animal or plant,
when we see the fore and hind limbs, the skull and ver-
tebræ, the jaws and legs of a crab, the petals, stamens,
and pistils of a flower, built on the same type or pat-
tern. During the many changes to which in the course
of time all organic beings have been subjected, certain
organs or parts have occasionally become at first of lit-
tle use and ultimately superfluous; and the retention of
such parts in a rudimentary and utterly useless condition
can, on the descent-theory, be simply understood. On

24
INTRODUCTION.
the principle of modifications being inherited at the
same age in the child, at which each successive varia-
tion first appeared in the parent, we shall see why rudi-
mentary parts and organs are generally well developed
in the individual at a very early age. On the same prin-
ciple of inheritance at corresponding ages, and on the
principle of variations not generally supervening at a
very early period of embryonic growth (and both these
principles can be shown to be probable from direct evi-
dence), that most wonderful fact in the whole round of
natural history, namely, the similarity of members of the
same great class in their embryonic condition,-the em-
bryo, for instance, of a mammal, bird, reptile, and fish
being barely distinguishable, — becomes simply intelli-
gible.
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 explana-
tion on the theory of independent Creations; they can-
not 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 conelude that at least all the
members of the same class have descended from a sin-
gle ancestor. A number of organic beings are included

NATURAL SELECTION.
25
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 de-
scent from a common form ; therefore it may be safely
admitted that all the members of the same class have
descended from one progenitor. But as the members
of quite distinct classes have something in common in
structure and much in common in constitution, analogy
and the simplicity of the view would lead us one step
further, and to infer as probable that all living creatures
have descended from a single prototype.
I hope that the reader will pause before coming to any
final and hostile conclusion on the theory of natural se-
lection. It is the facts and views to be hereafter given
which have convinced me of the truth of the theory.
The reader may consult my Origin of Species,' for a
general sketch of the whole subject; but in that work
he has to take many statements on trust. In consider-
ing the theory of natural selection, he will assuredly
meet with weighty difficulties, but these difficulties re-
late chiefly to subjects—such as the degree of perfec-
tion of the geological record, the means of distribution,
the possibility of transitions in organs, &c.-on which
we are confessedly ignorant; nor do we know how ig-
norant we are. If we are much more ignorant than is
generally supposed, most of these difficulties wholly dis-
appear. Let the reader reflect on the difficulty of look-
ing 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 ac-
count for all that we see in its past history. The pre-
sent action of natural selection may seem more or less
probable; but I believe in the truth of the theory, be-
2.

26
INTRODUCTION.
cause it collects under one point of view, and gives a
rational explanation of, many apparently independent
classes of facts.
4 In treating the several subjects in-
cluded in the present and succeeding
works I have continually been led to
ask for information from many zoolo-
gists, 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 re-
peatedly applied for information and
specimens to foreigners, and to British
merchants and officers of the Govern-
ment residing in distant lands, and,
with the rarest exceptions, I have re-
ceived prompt, open-handed, and valu-
able assistance. I cannot express too
strongly my obligations to the many
persons who have assisted me, and
who, I am convinced, would be equal-
ly willing to assist others in any scien-
tific investigation,

CHAP. I.
27
DOGS: THEIR PARENTAGE.
CHAPTER I.
DOMESTIC DOGS AND CATS
ANCIENT VARIETIES OF THE DOG - RESEMBLANCE OF DOMESTIC
DOGS IN VARIOUS COUNTRIES TO NATIVE CANINE SPECIES -- ANI-
MALS NOT ACQUAINTED WITH MAN AT FIRST FEARLESS — DOGS
RESEMBLING WOLVES AND JACKALS -HABIT OF BARKING AC-
QUIRED 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 CON-
STITUTION - 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 VARIA
BILITY,
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 to-
gether. We shall probably never be able to as

28
CHAP. I.
DOGS.
their origin with certainty. Paleontology' 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 charac-
ter of some of the breeds; and it almost necessarily as-
sumes, that a large number of species have become extinct
since man domesticated the dog; whereas we plainly see
that the members of the dog-family are extirpated by
human agency
with much difficulty; even so recently as
1710 the wolf existed in so small an island as Ireland.
The reasons which have led various authors to infer
that our dogs have descended from more than one wild
species are as follows. Firstly, the great difference be-
1 Owen, British Fossil Mammals,' p.
129 to 133. Pictet's Traité de Pal.,'
1853, tom. i. p. 202. De Blainville, in
his Ostéographie, Canidæ,' p. 142, has
largely discussed the whole subject and
concludes that the extinct parent of all
domesticated dogs came nearest to the
wolf in organization, and to the jackal in
habits.
2 Pallas, I believe, originated this doc-
trine in Act. Acad. St. Petersburgh,'
1780, Part ii. Ehrenberg has advocated
it, as may be seen in De Blainville's
Ostéographie,' p. 79. It has been carried
to an extreme extent by Col. Hamilton
Smith in the Naturalist Library,' vol. ix.
and x.
Mr. W. C. Martin adopts it in
his excellent History of the Dog,' 1815;
as does Dr. Morton, as well as Nott and
Gliddon, in the United States. Prof. Low,
in his 'Domesticated Animals,' 1845, P.
666, comes to this same conclusion. No
one has argued on this side with more
clearness and force than the late James
Wilson, of Edinburgh, in various papers
read before the Highland Agricultural
and Wernerian Societies. Isidor Geof-
froy St. Hilaire (Hist. Nat. Gén.,' 1860,
tom. iii. p. 107), though he believes that
most dogs have descended from the jack-
al, yet inclines to the belief that some are
descended from the wolf. Prof. Gervais
(Hist. Nat. Mamm.,'1855, tom. ii. p. 69),
referring to the view that all the domestic
races are the modified descendants of a
species, after a long discussion,
says, "Cette opinion est, suivant nous du
moins, la moins probable."

CHAP. I.
29
THEIR PARENTAGE.
tween the several breeds; but this will appear of com-
paratively little weight, after we shall have seen how
great are the differences between the several races of
various domesticated animals which certainly have de-
scended from a single parent-form. Secondly, the more
important fact that, at the most anciently known histori-
cal 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 four-
teenth century and the Roman classical period. At this
earlier period various breeds, namely hounds, house-dogs,
lapdogs, &c., existed; but as Dr. Walther has remarked
it is impossible to recognise the greater number with any
certainty. Youatt, however, gives a drawing of a beau-
tiful sculpture of two greyhound puppies from the Villa
of Antoninus. On an Assyrian monument, about 640 B.C.,
an enormous mastiff is figured ; and according to Sir H.
Rawlinson (as I was informed at the British Museum),
similar dogs are still imported into this same country. I
have looked through the magnificent works of Lepsius
and Rosellini, and on the monuments from the fourth to
the twelfth dynasties (i.e. from about 3400 B.C. to 2100
B.C.) several varieties of the dog are represented; most
of them are allied to greyhounds; at the later of these
periods a dog resembling a hound is figured, with droop-
ing ears, but with a longer back and more pointed head
3 Berjeau, The Varieties of the Dog;
in old Sculptures and Pictures, 1863.
Der Hund,' von Dr. F. L. Walther, s. 48,
Giessen, 1817: this author seems care-
fully to have studied all classical works
on the subject. See also 'Volz, Beiträge
zur Kulturgeschichte,' Leipzig, 1852, s.
115. Youаtt on the Dog,' 1815, p. 6. A
very full history is given by De Blainville
in his Ostéographie, Canidae.
4 I have seen drawings of this dog
from the tomb of the son of Esar Had-
don, and clay models in the British Mu-
seum. Nott and Gliddon, in their
Types of Mankind,' 1854, p. 393, give a
copy of these drawings. This dog has
been called a Thibetan mastiff, but Mr.
H. A. Oldfield, who is familiar with the
so-called Thibet mastiff, and has examin-
ed the drawings in the British Museum,
informs me that he considers them dif-
ferent.

30
CHAP. I.
DOGS.
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 also has described an Indian
Pariah dog as presenting the same monstrous character.
The most ancient dog represented on the Egyptian mo-
numents is one of the most singular; it resembles a grey-
hound, but has long pointed ears and a short curled tail:
a closely allied variety still exists in Northern Africa ; for
Mr. E. Vernon Harcourt 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, lap-
dogs, and turnspits, existed, more or less closely resem-
bling our present breeds. But there is not sufficient
evidence that any of these ancient dogs belonged to the
same idential sub-varieties with our present dogs. 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
5 Proc. Zoolog. Soc.,' July 12th, 1831.
& Sporting in Algeria,' p. 51.
7 Berjeau gives fac-similes of the
Egyptian drawings. Mr. C. L. Martin,
in his "History of the Dog,' 1845, copies
several figures from the Egyptian monu-
ments, and speaks with much confidence
with respect to their identity with still
living dogs. Messrs. Nott and Gliddon
(Types of Mankind,' 1854, p. 388) give
still more numerous figures. Mr. Glid-
don 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.

CHAP. I.
31
THEIR PARENTAGE.
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 bear-
ing in mind that the most barbarous nations possess do-
mestic dogs, the argument from insufficient time falls
away greatly in value.
Long before the period of any historical record the dog
was domesticated in Europe. In the Danish Middens of
the Neolithic or Newer Stone period, bones of a canine
animal are imbedded, and Steenstrup ingeniously argues
that these belonged to a domestic dog; for a very large
proportion of the bones of birds preserved in the refuse,
consists of long bones, which it was found on trial dogs
cannot devour. This ancient dog was succeeded in Den-
mark during the Bronze period by a larger kind, present-
ing certain differences, and this again during the Iron
period, by a still larger kind. In Switzerland we hear
from Prof. Rütimeyer, that during the Neolithic period
a domesticated dog of middle size existed, which in its
skull was about equally remote from the wolf and jackal,
and partook of the characters of our hounds and setters
or spaniels (Jagdhund und Wachtelhund). Rütimeyer
insists strongly on the constancy of form during a very
long period of time of this the most ancient known dog.
During the Bronze period a larger dog appeared, and this
closely resembled in its jaw a dog of the same age in
Denmark. Remains of two notably distinct varieties of
the dog were found by Schmerling in a cave;
10 but their
age cannot be positively determined.
The existence of a single race, remarkably constant in
8 These, and the following facts on the
Danish remains, are taken from M. Mor-
lot's most interesting memoir in 'Soc.
Vaudoise des Sc. Nat.,' tom. vi., 1860,
pp. 281, 299, 320.
9 Die Fauna der Pfahlbauten,' 1861,
s. 117, 162.
10 De Blainville, Ostéographie, Ca-
nidæ,

32
CHAP. I.
DOGS.
form during the whole Neolithic period, is an interesting
fact in contrast with what we see of the changes which
the races underwent during the period of the successive
Egyptian monuments, and in contrast with our existing
dogs. The character of this animal during the Neolithic
period, as given by Rütimeyer, supports De Blainville's
view that our varieties have descended from an unknown
and extinct form. But we should not forget that we
know nothing with respect to the antiquity of man in the
warmer parts of the world. The succession of the differ-
ent kinds of dogs in Switzerland and Denmark is thought
to be due to the immigration of conquering tribes bring-
ing with them their dogs; and this view accords with
the belief that different wild canine animals were domes-
ticated in different regions. Independently of the im-
migration 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 pos-
sess a large breed, which they barter at a high price with
other tribes. 11
The main argument in favour of the several breeds of
the dog being the descendants of distinct wild stocks, is
their resemblance in various countries to distinct species
still existing there. It must, however, be admitted that
the comparison between the wild and domesticated ani-
mal has been made but in few cases with sufficient exact-
ness. Before entering on details, it will be well to show
that there is no a priori difficulty in the belief that seve-
ral canine species have been domesticated; for there is
much difficulty in this respect with some other domestic
11 Sir R. Schomburgk has given me
information on this head. See also
Journal of R. Geograph. Soc.,' vol.
xiii., 1843, p. 65.

CHAP. I.
33
THEIR PARENTAGE.
quadrupeds and birds. Members of the dog family in-
habit nearly the whole world; and several species agree
pretty closely in habits and structure with our several
domesticated dogs. Mr. Galton has shown how fond
savages are of keeping and taming animals of all kinds.
Social animals are the most easily subjugated by man,
and several species of Canidæ hunt in packs. It deserves
notice, as bearing on other animals as well as on the dog,
that at an extremely ancient period, when man first en-
tered 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 an-
tarcticus) 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 hold-
ing a piece of meat in one hand and a knife in the other,
could sometimes stick them at night. On an island in the
Sea of Aral, when first discovered by Butakoff, the saigak
antelopes, which are “generally very timid and watchful,
did not fly from us, but on the contrary looked at us with
a sort of curiosity." So, again, on the shores of the Mau-
ritius, the manatee was not at first in the least afraid of
man, and thus it has been in several quarters of the world
with seals and the morse. I have elsewhere shown 13 how
slowly the native birds of several islands have acquired
and inherited a salutary dread of man: at the Galapagos
Archipelago I pushed with the muzzle of my gun hawks
from a branch, and held out a pitcher of water for other
birds to alight on and drink. Quadrupeds and birds
which have seldom been disturbed by man, dread him
12 Domestication of Animals :' Eth-
nological Soc., Dec. 22nd, 1863.
13 Journal of Researches,' &c., 1845,
p. 393. With respect to Canis antarc.
ticus, see p. 193. For the case of the an-
telope, see 'Journal Royal Geograph.
Soc.,' vol. xxiii. p. 94.

34
CHAP, I.
DOGS.
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 differ-
ent 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 sa-
gacious Richardson says,
" The resemblance between the
Northern American wolves (Canis lupus, var. occidenta-
lis) 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

CHAP. I.
35
THEIR PARENTAGE.
drooping tail, and scared look of the wolf. In disposi-
tion the Esquimaux dogs differ little from wolves, and,
according to Dr. Hayes, they are capable of no attach-
ment to man, and are so savage, that when hungry they
will attack even their masters. According to Kane they
readily become feral. Their affinity is so close with
wolves that they frequently cross with them, and the
Indians take the whelps of wolves" to improve the breed
of their dogs.” The half-bred wolves sometimes (Lamare-
Picquot) cannot be tamed, “though this case is rare;" but
they do not become thoroughly well broken in till the
second or third generation. These facts show that there
can be but little, if any, sterility between the Esquimaux
dog and the wolf, for otherwise they would not be used
to improve the breed. As Dr. Hayes says of these dogs,
“reclaimed wolves they doubtless are." 14
North America is inhabited by a second kind of wolf,
the prairie-wolf (Canis latrans), which is now looked at
by all naturalists as specifically distinct from the com-
mon 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
14 The authorities for the foregoing
statements are as follow :-Richardson,
in Fauna Boreali-Americana,' 1829, pp.
64, 75; Dr. Kane, 'Arctic Explorations,
1856, vol. i. pp. 398, 455; Dr. Hayes,
Arctic Boat Journey,' 1860, p. 167.
Franklin's Narrative,' vol. i. p. 269,
gives the case of three whelps of a black
wolf being carried away by the Indians.
Parry, Richardson, and others, give ac-
counts of wolves and dogs naturally cross-
ing in the eastern parts of North Ameri-
ca. Seeman, in his 'Voyage of H.M.S.
Herald,' 1853, vol. ii. p. 26, says the wolf
is often caught by the Esquimaux for the
purpose of crossing with their dogs, and
thus adding to their size and strength.
M. Lamare-Picquot, in 'Bull, de la. Soc.
d'Acclimat.,' tom. vii., 1860, p. 148, gives
a good account of the half-bred Esqui-
maux dogs.

36
CHAP. I.
DOGS.
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. 16
Turning to the southern parts of the New World,
Columbus found two kinds of dogs in the West Indies;
and Fernandez 16 describes three in Mexico: some of
these native dogs were dumb—that is, did not bark. In
Guiana it has been known since the time of Buffon that
the natives cross their dogs with an aboriginal species,
apparently the Canis cancrivorus. Sir R. Schomburgk,
who has so carefully explored these regions, writes to me,
“I have been repeatedly told by the Arawaak Indians,
who reside near the coast, that they cross their dogs with
a wild species to improve the breed, and individual dogs
have been shown to me which certainly resembled the C.
cancrivorus much more than the common breed. It is
but seldom that the Indians keep the C. cancrivorus for
domestic
purposes, nor is the Ai, another species of wild
dog, and which I consider to be identical with the Dusi-
cyon silvestris of H. Smith, now much used by the Are-
cunas 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 abo-
riginal 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 observ-
15 Fauna Boreali-Americana, 1829,
pp. 73, 78, 80. Nott and Gliddon, 'Types
of Mankind,' p. 383. The naturalist and
traveller Bartram is quoted by Hamilton
Smith, in 'Nat. Hist. Lib.,' vol. x. p. 156.
A Mexican domestic dog seems also to
resemble a wild dog of the same country;
but this may be the prairie-wolf. Another
capable judge, Mr. J. K. Lord (The
Naturalist in Vancouver Island,' 1866,
vol. ii. p. 218), says that the Indian dog
of the Spokans, near the Rocky Moun-
tains, " is beyond all question nothing
more than a tamed Cayote or prairie-
wolf," or Canis latrans.
16 I quote this from Mr. R. Hill's
excellent account of the Alco or domes-
tic dog of Mexico, in Gosse's Natural-
ist's Sojourn in Jamaica,' 1851, p. 329.

CHAP. I.
37
THEIR PARENTAGE.
er, Rengger," gives reasons for believing that a hairless
dog was domesticated when America was first visited by
Europeans: some of these dogs in Paraguay are still
dumb, and Tschudi 18 states that they suffer from cold in
the Cordillera. This naked dog is, however, quite distinct
from that found preserved in the ancient Peruvian burial-
places, and described by Tschudi, under the name of
Canis Ingo, 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. Ameri-
can species of Canidæ.
Turning to the Old World, some European dogs closely
resemble the wolf; thus the shepherd dog of the plains
of Hungary is white or reddish brown, has a sharp nose,
short, erect ears, shaggy coat, and bushy tail, and so much
resembles a wolf that Mr. Paget, who gives this descrip-
tion, 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. Shep-
herd 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. The European wolf differs
17 Naturgeschichte der Saeugethiere
von Paraguay,' 1830, s. 151.
18 Quoted in Humboldt's 'Aspects
of Nature,' Eng. transl., vol. i. p.
108.
19 Paget's Travels in Hungary and
Transylvania,' vol. i. p. 501. Jeitteles,
Fauna Hungariæ Superioris,' 1962, s.
13. See Pliny, Hist. of the World'
(Eng. transl.), 8th book, ch. xl., about
the Gauls crossing their dogs. See also
Aristotle, Hist. Animal.' lib. viii. c.
28. For good evidence about wolves
and dogs naturally crossing near the

38
CHAP. I.
DOGS.
slightly from that of North America, and has been ranked
by many naturalists as a distinct species. The common
wolf of India is also by some esteemed as a third species,
and here again we find a marked resemblance between
the pariah dogs of certain districts of India and the
Indian wolf.20
With respect to Jackals, Isidore Geoffroy Saint Hi-
laire 21
says that not one constant difference can be pointed
out between their structure and that of the smaller races
of dogs. They agree closely in habits: jackals, when
tamed and called by their master, wag their tails, crouch,
and throw themselves on their backs; they smell at the
tails of dogs, and void their urine sideways. A number
of excellent naturalists, from the time of Güldenstädt to
that of Ehrenberg, Hemprich, and Cretzschmar, have
expressed themselves in the strongest terms with respect
to the resemblance of the half-domestic dogs of Asia
and Egypt to jackals. M. Nordmann, for instance, says,
“Les chiens d’Awhasie ressemblent étonnamment à des
chacals.” Ehrenberg 2 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
Pyrenees, see M. Mauduyt, 'Du Loup
et de ses Races,' Poitiers, 1851 ; also
Pallas, in Acta Acad. St. Petersburgh,'
1780, part ii. p. 94.
20 I give this on excellent authority,
namely, Mr. Blyth (under the signature
of Zoophilus), in the 'Indian Sporting
Review,' Oct. 1856, p. 134. Mr. Blyth
states that he was struck with the
resemblance between a brush-tailed race
of pariah-dogs, north-west of Cawn pore,
and the Indian wolf. He gives corro-
borative evidence with respect to the
dogs of the valley of the Nerbudda.
21 For numerous and interesting de-
tails on the resemblance of dogs and
jackals, see Isid. Geoffroy St. Hilaire,
* Hist. Nat. Gén.,' 1860, tom. iii. p. 101.
See also Hist. Nat. des Mammifères,
par Prof. Gervais, 1855, tom. ii. p. 60.
22 Güldenstädt, 'Nov. Comment. Acad.
Petrop.,' tom. xx., pro anno 1775, p. 449.
23 Quoted by De Blainville in his
Ostéographie, Canidæ,' pp. 79, 98.

CHAP. I.
39
THEIR PARENTAGE.
on record in Algeria. 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 Gui-
nea are foxlike animals, and are dumb. On the east
coast of Africa, between lat. 4° and 6° south, and about
ten days' journey in the interior, a semi-domestic dog, as
the Rev. S. Erhardt informs me, is kept, which the na-
tives assert is derived from a similar wild animal. Lich-
tenstein 26
says
that the dogs of the Bosjemans present a
striking resemblance even in colour (excepting the black
stripe down the back) with the C. mesomelas of South
Africa. Mr. E. Layard informs me that he has seen a
Caffre dog which closely resembled an Esquimaux dog.
In Australia the Dingo is both domesticated and wild;
though this animal may have been introduced aboriginally
by man, yet it must be considered as almost an endemic
form, for its remains have been found in a similar state of
preservation and associated with extinct mammals, so that
its introduction must have been ancient.27
From this resemblance in several countries of the half-
domesticated dogs to the wild species still living there,
---from the facility with which they can often be crossed
together,-from even half-tamed animals being so much
valued by savages,-and from the other circumstances
previously remarked on which favour their domestication,
it is highly probable that the domestic dogs of the world
have descended from two good species of wolf (viz. C.
lupus and C. latrans), and from two or three other doubt-
24 See Pallas, in . Act. Acad. St. Pe-
tersburgh,' 1780, part ii. p. 91. For
Algeria, see Isid. Geoffroy St. Hilaire,
Hist. Nat. Gén.,' tom. iii. p. 177. In
both countries it is the male jackal
which pairs with female domestic dogs.
25 John Barbut's 'Description of the
Coast of Guinea in 1746.'
26 Travels in South Africa,' vol. ii. p.
272.
27 Selwyn, Geology of Victoria ; 'Jour-
nal of Geolog. Soc.,' vol. xiv., 1855, p.
536, and vol. xvi., 1860, p. 148; and
Prof. M'Coy, in 'Annals and Mag. of Nat.
Hist.' (3rd series), vol. ix., 1862, p. 147.
The Dingo differs from the dogs of the
central Polynesian islands. Dieffenbach
remarks ('Travels,' vol. ii. p. 45) that
the native New Zealand dog also differs
from the Dingo.

40
CHAP. I.
DOGS.
28
ful species of wolves (namely, the European, Indian, and
North African forms); from at least one or two South
American canine species; from several races or species
of the jackal; and perhaps from one or more extinct spe-
cies. Those authors who attribute great influence to the
action of climate by itself may thus account for the re-
semblance of the domesticated dogs and native animals
in the same countries; but I know of no facts supporting
the belief in so powerful an action of climate.
It cannot be objected to the view of several canine
species having been anciently domesticated, that these
animals are tamed with difficulty: facts have been already
given on this head, but I may add that the young of the
Canus primævus of India were tamed by Mr. Hodgson,
and became as sensible to caresses, and manifested as
much intelligence, as any sporting dog of the same age.
There is not much difference, as we have already shown
and shall immediately further see, in habits between the
domestic dogs of the North American Indians and the
wolves of that country, or between the Eastern pariah
dogs and jackals, or between the dogs which have run
wild in various countries and the several natural species
of the family. The habit of barking, however, which is
almost universal with domesticated dogs, and which does
not characterise a single natural species of the family,
seems an exception; but this habit is soon lost and soon
reacquired. The case of the wild dogs on the island of
Juan Fernandez having become dumb has often been
quoted, and there is reason to believe 2' that the dumb-
ness 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
28 Proceedings Zoolog. Soc.,'1833, p. - History Nat. Mamm.,' tom. ii. p. 61.
112. See, also, on the taming of the With respect to the aguara of Paraguay,
common wolf, L. Lloyd, Scandinavian see Rengger's work.
Adventures,' vol. i. p. 460, 1854. With 29 Roulin, in 'Mém. présent. par di-
respect to the jackal, see Prof. Gervais, vers Savans,' tom, vi. p. 341.

CHAP. I.
41
THEIR PARENTAGE.
dogs, of the Canis latrans type, when brought to Eng-
land, never learned to bark properly; but one born in
the Zoological Gardens 50 made his voice sound as loud-
ly as any other dog of the same age and size.” Accord-
ing to Professor Nillson, 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. An
interesting account has been given by Mr. G. Clarke 33
of some dogs run wild on Juan de Nova, in the Indian
Ocean; "they had entirely lost the faculty of barking;
they had no inclination for the company of other dogs,
nor did they acquire their voice," during a captivity of
several months. On the island they “congregate in vast
packs, and catch sea-birds with as much address as foxes
could display.” The feral dogs of La Plata have not
become dumb; they are of large size, hunt single or in
packs, and burrow holes for their young. In these ha-
bits the feral dogs of La Plata resemble wolves and
jackals; both of which hunt either singly or in packs,
and burrow holes. These feral dogs have not become
uniform in colour on Juan Fernandez, Juan de Nova, or
La Plata.36 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
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
37
32
30 Martin, History of the Dog,' p. 14.
31 Quoted by L. Lloyd in 'Field Sports
of North of Europe,' vol. i. p. 387.
Quatrefages, Soc. d'Acclimat.,'
May 11th, 1863, p. 7.
Discours, Exposition des Races Ca-
nines,' 1865, p. 3.
35 With respect to wolves burrowing
holes, see Richardson, 'Fauna Boreali-
Americana,' p. 64; and Bechstein, Na-
turgesch. Deutchlands,' b. i. s. 617.
33 Annals and Mag. of Nat. Hist.,'
vol. xv., 1845, p. 140.
34 Azara, Voyages dans l'Amér. Mé-
rid.,' tom. i. p. 381 ; his account is fully
confirmed by Rengger. Quatrefages
gives an account of a bitch brought
from Jerusalem to France which bur-
rowed a hole and littered in it. See
36 See Poeppig, 'Reise in Chile,' b.i.
s. 290; Mr. G. Clarke, as above; and
Rengger, s. 155.
37 Dogs, Nat. Library,' vol. x. p.
121: an endemic South American dog
seems also to have become feral in this
island. See Gosse's Jamaica,' p. 340.

42
CHAP. I.
DOGS.
naturalised in Australia,“ varies considerably in colour,"
as I am informed by Mr. P. P. King: a half-bred Dingo
reared in England 38 showed signs of wishing to bur-
l'OW.
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 grey-
hounds 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 mon-
grel, and some fox-hounds, coloured black and white, with not a
trace of tan-colour, excepting the spots over the eyes, and some-
times 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 gra-
dation, from the whole face being tan-coloured, to a complete ring
round the eyes, to a minute spot over the inner and upper corners.
The spots occur in various sub-breeds of terriers and spaniels; in
setters; in hounds of various kinds, including the turnspit-like
German badger-hound ; in shepherd dogs; in a mongrel, of which
neither parent had the spots; in one pure bulldog, though the spots
were in this case almost white; and in greyhounds - but true black-
and-tan greyhounds are excessively rare; nevertheless I have been
assured by Mr. Warwick, that one ran at the Caledonian Champion
meeting of April, 1860, and was marked precisely like a black-
and-tan terrier.” Mr. Swinhoe at my request looked at the dogs
38 Low, 'Domesticated Animals,' p. 650.

CHAP. I.
43
THEIR PARENTAGE.
in China, at Amoy, and he soon noticed a brown dog with yellow
spots over the eyes. Colonel H. Smith 39 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 pa-
rent-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 tol-
erably plain correlation between tan-coloured legs and face; and
less frequently between black legs and a black face; and this gene-
ral 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 0. mesomelas, C. aureus, and (judging from Colonel
Ham. Smith's drawing) in C. alopex and C. thaleb. Other species have
a trace of a black line over the corners of the eyes, as in C. variega-
tus, cinereo-variegatus, and fulvus, and the wild Dingo. Hence I
am inclined to conclude that a tendency for tan-coloured spots to ap-
pear over the eyes in the various breeds of dogs, is analogous to the
case observed by Desmarest, namely, that when any white appears
on a dog the tip of the tail is always white," de manière à rappeler
la tache terminale de même couleur, qui caractérise la plupart des
Canidés sauvages." 40
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
89 The Naturalist Library,' Dogs, vol.
x. pp. 4, 19.
40 Quoted by Prof. Gervais Hist. Nat.
Mamm.,' tom. ii. p. 66.

44
CHAP. I.
DOGS.
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 de-
gree variable." 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-
ninė, sixty-two, and sixty-seven days. The average pe-
riod is sixty-three days; but Bellingeri states that this
holds good only with large dogs; and that for small races
it is from sixty to sixty-three days; Mr. Eyton of Eyton,
who has had much experience with dogs, also informs me
that the time is apt to be longer with large than with
small dogs.
F. Cuvier has objected that the jackal would not have
been domesticated on account of its offensive smell; but
savages are not sensitive in this respect. The degree of
odour, also, differs in the different kinds of jackal; 42 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 ter-
riers---differ much in this respect; and M. Godron states
that the hairless so-called Turkish dog is more odoriferous
41 J. Hunter shows that the long pe-
riod of seventy-three days given by Buf-
fon is easily explained by the bitch hav-
ing received the dog many times during
a period of sixteen days ("Phil, Trans-
act.,' 1787, p. 253). Hunter found that
the gestation of a mongrel from wolf and
dog ("Phil. Transact.,' 1789, p. 160) ap-
parently was sixty-three days, for she
received the dog more than once. The
period of a mongrel dog and jackal was
fifty-nine days. Fred. Cuvier found the
period of gestation of the wolf to be
("Dict. Class d'Hist. Nat.,' tom. iv. p. 8)
two months and a few days, which agrees
with the dog. Isid. G. St. Hilaire, who
has discussed the whole subject, and from
whom I quote Bellingeri, states (Hist.
Nat. Gén.,' tom. iii. p. 112) that in the
Jardin des Plantes the period of the
jackal has been found to be from sixty
to sixty-three days, exactly as with the
dog.
42 See Isid. Geoffroy St. Hilaire, 'Hist.
Nat. Gén.,' tom. iii. p. 112, on the odour
of jackals. Col. Ham. Smith, in 'Nat.
Hist. Lib.,' vol. x. p. 289.

CHAP. I.
45
THEIR PARENTAGE.
than other dogs. Isidore Geoffroy * gave to a dog the
same odour as that from a jackal by feeding it on raw
flesh.
The belief that our dogs are descended from wolves,
jackals, South American Canidæ, and other species sug-
gests 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 inter-
crossed ; and such sterility will be admitted as almost cer-
tain by all those who believe that the lessened fertility of
crossed forms is an infallible criterion of specific distinct-
ness. 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, mu-
tually fertile together. But, as Broca has well remarked,
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 re-
productive powers differ in the several races of the dog
when crossed are (passing over mere size as rendering pro-
pagation difficult) as follows: the Mexican Alco 45
appa-
rently dislikes dogs of other kinds, but this perhaps is not
strictly a sexual feeling; the hairless endemic deg of Par-
aguay, according to Rengger, mixes less with the Euro-
pean 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
48 Quoted by Quatrefages in Bull. Soc.
d'Acclimat.,' May 11th, 1863.
14 Journal de la Physiologie,' tom. ii.
guay,' s. 153. With respect to Spitz
dogs, see Bechstein's Naturgesch.
Deutschlands,' 1801, b. i. s. 638. With
respect to Dr. Hodgkin's statement made
before Brit. Assoc., see "The Zoologist,
vol. iv., for 1845-46, p. 1097.
p. 355.
45 See Mr. R. Hill's excellent account
of this breed in Gosse's 'Jamaica,' p.
838; Rengger's 'Saeugethiere von Para-

46
CAAP. I.
DOGS.
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 46 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 seve-
ral wild stocks, that I am led to admit the truth of this
hypothesis.
There is another and closely allied difficulty consequent
on the doctrine of the descent of our domestic dogs from
several wild species, namely, that they do not stem to be
perfectly fertile with their supposed parents. But the
experiment has not been quite fairly tried; the Hungarian
dog, for instance, which in external appearance so closely
resembles the European wolf, ought to be crossed with
this wolf; and the pariah-dogs of India with Indian
wolves and jackals; and so in other cases. That the
sterility is very slight between certain dogs and wolves
and other Canidæ is shown by savages taking the trouble
to cross them. Buffon got four successive generations
from the wolf and dog, and the mongrels were perfectly
fertile together. 47 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.48 But these animals
were closely confined; and many wild animals, as we
46 Acta Acad. St. Petersburgh,' 1780,
part ii. pp. 84, 100.
47 M. Broca has shown (Journal de
Physiologie,' tom. ii. p. 353) that Buffon's
experiments have been often misrepre-
sented. Broca has collected (pp. 390-395)
many facts on the fertility of crossed
dogs, wolves, and jackals.
48 De la Longévité Humaine,' par M.
Flourens, 1855, p. 143. Mr. Blyth says
(Indian Sporting Review,' vol. ii. p. 137)
that he has seen in India several hybrids

CHAP. I.
47
THEIR PARENTAGE.
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. 49 Some hounds from Central Africa, brought
home by Major Denham, never bred in the Tower of Lon-
don; and a similar tendency to sterility might be trans-
mitted to the hybrid offspring of a wild animal. Moreover,
it appears that in M. Flourens' experiments the hybrids
were closely bred in and in for three or four generations;
but this circumstance, although it would almost certainly
increase the tendency to sterility, would hardly account
for the final result, even though aided by close confine-
ment, unless there had been some original tendency to
lessened fertility. Several years ago I saw confined in the
Zoological Gardens of London a female hybrid from an
English dog and jackal, which even in this the first gene-
ration was so sterile that, as I was assured by her keeper,
she did not fully exhibit her proper periods; but this case,
from the numerous instances of fertile hybrids from these
two animals, was certainly exceptional. In almost all ex-
periments 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.
from the pariah-dog and jackal; and be-
tween one of these hybrids and a terrier.
The experiments of Hunter on the jackal
are well known. See also Isid. Geoffroy
St. Hilaire, Hist. Nat. Gén.,' tom. iii. p.
217, who speaks of the hybrid offspring
of the jackal as perfectly fertile for three
generations.
49 On authority of F. Cuvier, quoted
in Bronn's Geschichte der Natur, ' B. ii.
s. 164.
50 W. C. L. Martin, History of the
Dog,' 1845, p. 203. Mr. Philip P. King,
after ample opportunities of observation,
informs me that the Dingo and European
dogs often cross in Australia.

48
CHAP. L.
DOGS.
Notwithstanding the difficulties in regard to fertility
given in the last two paragraphs, when we reflect on
the inherent improbability of man having domesticated
throughout the world one single species alone of so
widely distributed, so easily tamed, and so useful a
group as the Canidæ; when we reflect on the extreme
antiquity of the different breeds; and especially when
we reflect on the close similarity, both in external struc-
ture 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 si-
mensis, with its elongated muzzle; that of the larger
dogs from the larger wolves, and the smaller and slighter
dogs from jackals: and thus perhaps we may account
for certain constitutional and climatal differences. But it
would be a great error to suppose that there has not been
in addition a large amount of variation. The inter-
crossing 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 ex-
plain by crossing the origin of such extreme forms as
thoroughbred greyhounds, bloodhounds, bulldogs, Blen-
heim spaniels, terriers, pugs, &c., unless we believe that
forms equally or more strongly characterised in these
different respects once existed in nature. But hardly any
61 Rüppel, 'Neue Wirbelthiere von fine animal in the British Museum
Abyssinien,' 1835-40 ; Mammif.' s. 39. 52 Even Pallas admits this: see Act
pl. xiv. There is a specimen of this Acad. St. Petersburgh,' 1780, p. 93.

CHAP. I.
49
DIFFERENCES OF BREEDS.
one has been bold enough to suppose that such unnatural
form:s ever did or could exist in a wild state. When
compared with all known members of the family of Ca-
nidæ they betray a distinct and abnormal origin. No
instance is on record of such dogs as bloodhounds,
spaniels, true greyhounds having been kept by savages:
they are the product of long-continued civilization.
The number of breeds and sub-breeds of the dog is great : Youatt,
for instance, describes twelve kinds of greyhounds. I will not at
tempt to enumerate or describe the varieties, for we cannot discri-
minate 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 63 that in form the differences are "plus
fortes que celles d'aucunes espèces sauvages d'un même genre na-
turel.” 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 zygoma-
tic arch, and of the temporal fossæ; the position of the occiput-all
vary considerably. 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 ; 55 and Pro-
fessor Gervais says that there are dogs" qui ont sept paires de dents
supérieures et huit inférieures." De Blainville 56 has given full par-
ticulars on the frequency of these deviations in the number of the
teeth, and has shown that it is not always the same tooth which is
supernumerary. In short-muzzled races, according to H. Müller, 57
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, _sometimes having none except one molar on each side;
53 Quoted by I. Geoffroy, Hist. Nat.
Gén.,' tom. iii. p. 453.
54 F. Cuvier, in 'Annales du Muséum,'
tom. xviii. p. 337; Godron, De l'Espèce,
tom, i. p. 342; and Col. Ham. Smith, in
Naturalist's Library,' vol. ix. p. 101.
56 Isid. Geoffroy Saint Hilaire, Hist.
des Anomalies,' 1832, tom. i. p. 660.
Gervais, Hist. Nat. des Mammifères,
tom. ii., 1855, p. 66. De Blainville, ("Os-
téographie, Canidæ,' p.137) has also seen
an extra molar on both sides.
56 Ostéographie, Canidæ,' p. 137.
57 Würzburger, 'Medecin. Zeitschrift,
1860, B. i. s. 265.
58 Mr. Yarrell, in Proc. Zoolog. Soc.,'
Oct. 8th, 1833. Mr. Waterhouse showed
me a skull of one of these dogs, which
had only a single molar on each side and
some imperfect incisors.
3

50
CHAP. I.
DOGS.
but this, though characteristic of the breed, must be considered as a
monstrosity. M. Girard,59 who seems to have attended closely to the
subject, says that the period of the appearance of the permanent
teeth differs in different dogs, being earlier in large dogs; thus the
mastiff assumes its adult teeth in four or five months, whilst in the
spaniel the period is sometimes more than seven or eight months.
With respect to minor differences little need be said. Isidore
Geoffroy has shown 60 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.61 Every
one knows how the ears vary in size in different breeds, and with
their great development their muscles become atrophied. Certain
breeds of dogs are described as having a deep furrow between the
nostrils and lips. The caudal vertebræ, according to F. Cuvier, on
whose authority the two last statements rest, vary in number; and
the tail in shepherd dogs is almost absent. The mammæ vary from
seven to ten in number: Daubenton, having examined twenty-one
dogs, found eight with five mamme on each side; eight with four
on each side; and the others with an unequal number on the two
sides.62 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 cha-
racters, varies. These modifications, however, in the feet of dogs
are not important, because they ought to be ranked, as De Blain-
ville has shown,63 as monstrosities. Nevertheless they are interest-
ing 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
59 Quoted in The Veterinary,' Lon-
don, vol. viii. p. 415.
60 Hist. Nat. Général,' tom. iii. p.
448.
61 W. Scrope, 'Art of Deer-Stalking,'
63 De Blainville, Ostéographie Cani-
dæ,' p. 134. F. Cuvier, Annales du
Muséum,' tom. xviii. p. 342. In regard
to mastiffs, see Col. Ham. Smith, Nat.
Lib.,' vol. x. p. 218. For the Thibet mas-
tiff, see Mr. Hodgson in Journal of As.
Soc. of Bengal,' vol. i., 1832, p. 342.
p. 354.
62 Quoted by Col. Ham. Smith in 'Na-
uralist's Library,' vol. x, p. 79.

CHAP. I.
51
DIFFERENCES OF BREEDS.
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 consti-
tutional differences: the pulse, says Youatt, 64 varies materially
according to the breed, as well as to the size of the animal.” Dif-
ferent breeds of dogs are subject in different degrees to various dis-
eases. 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. The Rev. R Everest 66 be-
lieves that no one has succeeded in keeping the Newfoundland dog
long alive in India; so it is, according to Lichtenstein,67 even at the
Cape of Good Hope. "The Thibet mastiff degenerates on the plains
of India, and can live only on the mountains.68 Lloyd "' 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 admis-
sion that their skulls differ more than do those of the
species of
genus, and bearing in mind how
closely the bones of wolves, jackals, foxes, and other
Canidæ agree, it is remarkable that we meet with the
statement, repeated over and over again, that the races
of the dog differ in no important characters. A highly
competent judge, Prof. Gervais, º admits, “ si l'on prenait
sans contrôle les altérations dont chacun de ces organes
est susceptible, on pourrait croire qu'il y a entre les
chiens domestiques des différences plus grandes que
celles qui séparent ailleurs les espèces, quelquefois même
les genres.” Some of the differences above enumerated
any
natural
na.
64 The Dog,' 1845, p. 156. With re-
spect to diseases, Youatt asserts (p. 167)
that the Italian greyhound is strongly
subject" to polypi in the matrix or vagi-
The spaniel and pug (p. 182) are
most liable to bronchocele. The liability
to distemper (p. 232) is extremely differ-
ent in different breeds On the distem-
per, see also Col. Hutchinson on 'Dog
Breaking,' 1850, p. 279.
65 See Youаtt on the Dog, p. 15; "The
Veterinary,' London, vol. xi. p. 235.
68 6Journal of As. Soc. of Bengal,' vol.
iii. p. 19.
67 Travels,' vol. ii. p. 15.
68 Hodgson, in 'Journal of As. Soc. of
Bengal,' vol. i. p. 342.
69 Field Sports of the North of Eu-
rope,' vol. ii. p. 165.
70 Hist. Nat. des Mammif.,'1855, tom:
ii. pp. 66, 67.

52
CHAP. I.
DOGS.
are in one respect of comparatively little value, for they
are not characteristic of distinct breeds: no one pretends
that such is the case with the additional molar teeth or with
the number of mammæ ; the additional digit is generally
present with mastiffs, and some of the more important dif-
ferences in the skull and lower jaw are more or less cha-
racteristic of various breeds. But we must not forget that
the predominant power of selection has not been applied
in any of these cases; we have variability in important
parts, but the differences have not been fixed by selection.
Man cares for the form and fleetness of his greyhounds,
for the size of his mastiffs, for the strength of the jaw in
his bulldogs, &c.; but he cares nothing about the num-
ber of their molar teeth or mammæ or digits; nor do we
know that differences in these organs are correlated with,
or owe their development to differences in other parts of
the body about which man does care. Those who have
attended to the subject of selection will admit that, na-
ture 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 addition-
al 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 mak-
ing 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, profound-
ly ignorant. We may attribute part of the difference in
external form and constitution to inheritance from dis-
tinct 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

CHAP. I.
53
MEANS OF MODIFICATION.
have already seen how often savages cross their dogs
with wild native species; and Pennant gives a curious
account" of the manner in which Fochabers, in Scotland,
was stocked with a multitude of curs of a most wolf-
ish aspect” from a single hybrid-wolf brought into that
district.
It would appear that climate to a certain extent di-
rectly 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 men-
tal faculties, but in form. Captain Williamson," 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 pre-
vent their crossing with native dogs ; so that the de-
terioration 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 con-
tracted, their noses more pointed, their size inferior, and
71 History of Quadrupeds,' 1793, vol.
1. p. 238.
72 Oriental Field Sports,' quoted by
Youatt, The Dog,' p. 15.

54
CHAP. I.
DOGS.
their limbs more slender. This remarkable tendency to
rapid deterioration in European dogs subjected to the
climate of India, may perhaps partly be accounted for by
the tendency to reversion to a primordial condition which
many animals exhibit, as we shall see in a future chapter,
when exposed to new conditions of life.
Some of the peculiarities characteristic of the several
breeds of the dog have probably arisen suddenly, and,
though strictly inherited, may be called monstrosities;
for instance, the shape of the legs and body in the turn-
spit 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 cours-
ing hares, as with water-dogs in swimming—and the
want of exercise, in the case of lap-dogs-must have pro-
duced 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 methodi-
eal and unconscious, of slight individual differences,-the
latter kind of selection resulting from the occasional pre-
servation, during hundreds of generations, of those indi-
vidual dogs which were the most useful to man for cer-
tain purposes and under certain conditions of life. In a
future chapter on Selection I shall show that even bar-
barians 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,“ the dogs are sometimes com-
pelled by want to leave their masters and provide for
themselves; but in a few days they generally return.
73 Quoted by Mr. Galton, 'Domestication of Animals,' p. 13.

CHAP. I.
55
MEANS OF MODIFICATION.
And we may infer that dogs of different shapes, sizes,
and habits, would have the best chance of surviving un-
der 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,
extends to the third phalanges, whilst in ordinary dog's
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.” 76 English
otter-hounds are said to have webbed feet; a friend ex-
amined for me the feet of two, in comparison with the
feet of some harriers and bloodhounds; he found the
skin variable in extent in all, but more developed in the
otter than in the other hounds. As aquatic animals
which belong to quite different orders have webbed feet,
there can be no doubt that this structure would be ser-
74 Hist. Nat. Gén.,' tom. iii. p. 450.
75 Mr. Greenhow on the Canadian
Dog, in Loudon's 'Mag. of Nat. Hist.,'
vol vi. 1833, p. 511.
78 See Mr. C. O. Groom-Napier on the
webbing of the bind feet of Otter-hounds
in Land and Water,' Oct. 13th, 1866, p.
270.

56
CHAP. I.
DOGS.
viceable to dogs that frequent the water. We may confi-
dently infer that no man ever Selected his water-dogs by
the extent to which the skin was developed between
their toes; but what he does, is to preserve and breed
from those individuals which hunt best in the water, or
best retrieve wounded game, and thus he unconsciously
selects dogs with feet slightly better webbed. Man
thus closely imitates Natural Selection. We have an
excellent illustration of this same process in North
America, where, according to Sir J. Richardson," all the
wolves, foxes, and aboriginal domestic dogs have their
feet broader than in the corresponding species of the Old
World, and “well calculated for running on the snow.'
Now, in these Arctic regions, the life or death of every
animal will often depend on its success in hunting over
the snow when softened; and this will in part depend on
the feet being broad; yet they must not be so broad as
to interfere with the activity of the animal when the
ground is sticky, or with its power of burrowing holes
or with other habits of life.
As changes in domestic breeds which take place so
slowly as not to be noticed at any period, whether due to
the selection of individual variations or of differences re-
sulting from crosses, are most important in understand-
ing 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, who paid particular atten-
tion 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
77 Fauna Boreali-Americana,' 1829,
78 The Horse in all his Varieties,
&c., 1829, pp. 230, 234.
P. 62.

CHAP. I.
57
MEANS OF MODIFICATION.
was effected by a cross with the greyhound. With re-
spect to this latter dog, Youаtt 79 who is generally cau-
tious 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 80
believes that our English greyhounds are the descend-
ants, progressively improved, of the large rough grey-
hounds 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 grey-
hounds, which failed in courage, with a bulldog-this
breed being chosen from being deficient in the power of
scent; “after the sixth or seventh generation,” says
Youatt, "there was not a vestige left of the form of the
bulldog, but his courage and indomitable perseverance
remained."
Youatt infers, from a comparison of an old picture of
King Charles's spaniels with the living dog, that “the
breed of the present day is materially altered for the
'worse :" the muzzle has become shorter, the forehead
more prominent, and the eyes larger: the changes in this
case have probably been due to simple selection. The
setter, as this author remarks in another place, "is evi-
dently 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 satis-
79 The Dog,' 1845, pp. 31, 35; with 80 In the Encyclop. of Rural Sports,'
respect to King Charles's spaniel, p. 45; p. 557.
for the setter, p. 90.

58
CHAP. I.
DOGS.
factory on this point, we might have recourse to history :")
he then refers to a document dated 1685 bearing on this
subject, and adds that the pure Irish setter shows no signs
of a cross with the pointer, which some authors suspect
has been the case with the English setter. Another
writer 81 remarks that, if the mastiff and English bulldog
had formerly been as distinct as they are at the present
time (i.e. 1828), so accurate an observer as the poet Gay
(who was the author of 'Rural Sports' in 1711) would
have spoken in his Fable of the Bull and the Bulldog, and
not of the Bull and the Mastiff. There can be no doubt
that the fancy bulldogs of the present day, now that they
are not used for bull-baiting, have become greatly reduced
in size, without any express intention on the part of the
breeder. Our pointers are certainly descended from a
Spanish breed, as even their names, Don, Ponto, Carlos,
&c., would show: it is said that they were not known in
England before the Revolution in 1688; 82 but the breed
since its introduction has been much modified, for Mr.
Borrow, who is a sportsman and knows Spain intimately
well, informs me that he has not seen in that country any
breed “corresponding in figure with the English pointer;
but there are genuine pointers near Xeres which have
been imported by English gentlemen.” A nearly parallel
case is offered by the Newfoundland dog, which was cer-
tainly brought into England from that country, but which
has since been so much modified that, as several writers
have observed, it does not now closely resemble any ex-
isting native dog in Newfoundland.83
These several cases of slow and gradual changes in our
English dogs possess some interest; for though the
changes have generally, but not invariably, been caused
81 The Farrier,' 1829, vol. i. p. 337.
82 See Col. Hamilton Smith on the an-
tiquity of the Pointer, in Naturalist's
Library,' vol. x. p. 196.
83 The Newfoundland dog is believed
to have originated from a cross between
the Esquimaux dog and a large French
hound. See Dr. Hodgkin, Brit Assoc.,'
1844; Bechstein's 'Naturgesch. Deutsch-
lands, Band i. s. 574; Naturalist's
Library,' vol. x. p. 132; also Mr. Jukes'
Excursion in and about Newfoundland.'

CHAP. I.
59
DOMESTIC CATS: THEIR PARENTAGE.
by one or two crosses with a distinct breed, yet we may
feel sure, from the well-known extreme variability of
crossed breeds, that rigorous and long-continued selection
must have been practised, in order to improve them in a
definite manner. As soon as any strain or family became
slightly improved or better adapted to altered circum-
stances, it would tend to supplant the older and less im-
proved strains. For instance, as soon as the old foxhound
was improved by a cross with the greyhound, or by sim-
ple selection, and assumed its present character-and the
change was probably required by the increased fleetness
of our hunters-it rapidly spread throughout the country,
and is now everywhere nearly uniform. But the process
of improvement is still going on, for every one tries to
improve his strain by occasionally procuring dogs from
the best kennels. Through this process of gradual substi-
tution the old English hound has been lost; and so it has
been with the old Irish greyhound and apparently with
the old English bulldog. But the extinction of former
breeds is apparently aided by another cause; for when-
ever a breed is kept in scanty numbers, as at present with
the bloodhound, it is reared with difficulty, and this ap-
parently is due to the evil effects of long-continued close
interbreeding. As several breeds of the dog have been
slightly but sensibly modified within so short a period as
the last one or two centuries, by the selection of the best
individual dogs, modified in many cases by crosses with
other breeds; and as we shall hereafter see that the breed-
ing of dogs was attended to in ancient times, as it still is
by savages, we may conclude that we have in selection,
even if only occasionally practised, a potent means of
modification.
DOMESTIC CATS.
Cats have been domesticated in the East from an ancient
period; Mr. Blyth informs me that they are mentioned
in a Sanskrit writing 2000 years old, and in Egypt their

60
CHAP. I.
DOMESTIC CATS.
antiquity is known to be even greater, as shown by mon-
umental drawings and their mummied bodies. These
mummies, according to De Blainville, 64 who has particu-
larly studied the subject, belong to no less than three
species, namely, F. caligulata, bubastes, and chaus. The
two former species are said to be still found, both wild
and domesticated, in parts of Egypt. F. caligulata
presents a difference in the first inferior milk molar tooth,
as compared with the domestic cats of Europe, which
makes De Blainville conclude that it is not one of the
parent-forms of our cats. Several naturalists, as Pallas,
Temminck, Blyth, believe that domestic cats are the de-
scendants of several species commingled : it is certain
that cats cross readily with various wild species, and it
would
appear that the character of the domestic breeds
has, at least in some cases, been thus affected. Sir W.
Jardine has no doubt that, “in the north of Scotland,
there has been occasional crossing with our native species
(F. sylvestris), and that the result of these crosses has
been kept in our houses. I have seen,” he adds,“ many
cats very closely resembling the wild cat, and one or two
that could scarcely be distinguished from it.” Mr.
Blyth® remarks on this passage, “but such cats are
never seen in the southern parts of England ; still, as
compared with any Indian tame cat, the affinity of the
ordinary British cat to F. sylvestris is manifest; and due
I suspect to frequent intermixture at a time when the
tame cat was first introduced into Britain and continued
rare, while the wild species was far more abundant than
at present.” In Hungary, Jeitteles was assured on
trustworthy authority that a wild male cat crossed with
86
84 De Blainville, Ostéographie, Felis,'
p. 65, on the character of F. caligulata,
pp. 85, 89, 90, 175, on the other mum-
mied species. He quotes Ehrenberg on
F. maniculata being mummied.
85 Asiatic Soc. of Calcutta; Curator's
Report, Aug. 1856. The passage from
Sir W. Jardine is quoted from this Re-
port. Mr. Blyth, who has especially at-
tended to the wild and domestic cats of
India, has given in this Report a very
interesting discussion on their origin.
88 Fauna Hungariæ Sup.,' 1862, s. 12.

CHAP. I.
61
THEIR VARIATION.
a female domestic cat, and that the hybrids long lived in
a domesticated state. In Algiers the domestic cat has
crossed with the wild cat (F. Lybica) of that country. 87
In South Africa, as Mr. E. Layard informs me, the do-
mestic cat intermingles freely with the wild F. caffra;
he has seen a pair of hybrids which were quite tame and
particularly attached to the lady who brought them up;
and Mr. Fry has found that these hybrids are fertile. In
India the domestic cat, according to Mr. Blyth, has cross-
ed with four Indian species. With respect to one of
these species, F. chaus, an excellent observer, Sir W.
Elliot, informs me that he once killed, near Madras, a
wild brood, which were evidently hybrids from the do-
mestic cat; these young animals had a thick lynx-like
tail and the broad brown bar on the inside of the fore-
arm characteristic of F. chaus. Sir W. Elliot adds that
he has often observed this same mark on the forearms of
domestic cats in India. Mr. Blyth states that domestic
cats coloured nearly like F. chaus, but not resembling
that species in shape, abound in Bengal; he adds,“ such
a colouration is utterly unknown in European cats, and
the proper tabby markings (pale streaks on a black
ground, peculiarly and symmetrically disposed) so com-
mon in English cats, are never seen in those of India.”
Dr. D. Short has assured Mr. Blythe' that at Hansi hy-
brids between the common cat and F. ornata (or tor-
quata) occur, “and that many of the domestic cats of
that part of India were undistinguishable from the wild
F. ornata.” Azara states, but only on the authority of
the inhabitants, that in Paraguay the cat has crossed
with two native species. From these several cases we
see that in Europe, Asia, Africa, and America, the com-
mon cat, which lives a freer life than most other domesti-
cated animals, has crossed with various wild species;
87 Isid. Geoffroy Saint Hilaire, Hist. Nat. Gén., ' tom. iii. p. 177.
88 Proc. Zoolog. Soc., ' 1863, p. 184.

62
CHAP. I.
DOMESTIC CATS.
and that in some instances the crossing has been suffi-
ciently frequent to affect the character of the breed.
Whether domestic cats have descended from several
distinct species, or have only been modified by occa-
sional crosses, their fertility, as far as is known, is unim-
paired. The large Angora or Persian cat is the most dis-
tinct in structure and habits of all the domestic breeds;
and is believed by Pallas, but on no distinct evidence, to
be descended from the F. manul of middle Asia; but I
am assured by Mr. Blyth that this cat breeds freely with
Indian cats, which, as we have already seen, have appa-
rently been much crossed with F. chaus. In England
half-bred Angora cats are perfectly fertile with the com-
mon cat; I do not know whether the half-breeds are fer-
tile one with another; but as they are common in some
parts of Europe, any marked degree of sterility could
hardly fail to have been noticed.
Within the same country we do not meet with distinct
races of the cat, as we do of dogs and of most other
domestic animals; though the cats of the same country
present a considerable amount of fluctuating variability.
The explanation obviously is that, from their nocturnal
and rambling habits, indiscriminate crossing cannot with-
out much trouble be prevented. Selection cannot be
brought into play to produce distinct breeds, or to keep
those distinct which have been imported from foreign
lands. On the other hand, in islands and in countries
completely separated from each other, we meet with
breeds more or less distinct; and these cases are worth
giving as showing that the scarcity of distinct races in
the same country is not caused by a deficiency of varia-
bility in the animal. The tailless cats of the Isle of Man
are said to differ from common cats not only in the want
of a tail, but in the greater length of their hind legs, in
the size of their heads, and in habits. The Creole cat of
Antigua, as I am informed by Mr. Nicholson, is smaller,
and has a more elongated head, than the British cat. In

CHAP. I.
63
THEIR VARIATION.
Ceylon, as Mr. Thwaites writes to me, every one at first
notices the different appearance of the native cat from
the English animal; it is of small size, with closely lying
hairs; its head is small, with a receding forehead; but
the ears are large and sharp; altogether it has what is
there called a “low-caste" appearance. Rengger 80 says
that the domestic cat, which has been bred for 300 years
in Paraguay, presents a striking difference from the Eu-
ropean cat; it is smaller by a fourth, has a more lanky
body, its hair is short, shining, scanty, and lies close,
especially on the tail: he adds that the change has been
less at Ascension, the capital of Paraguay, owing to the
continual crossing with newly imported cats; and this
fact well illustrates the importance of separation. The
conditions of life in Paraguay appear not to be highly
favourable to the cat, for, though they have run half-
wild, they do not become thoroughly feral, like so many
other European animals. In another part of South Ame-
rica, according to Roulin,'' the introduced cat has lost
the habit of uttering its hideous nocturnal howl. The
Rev. W. D. Fox purchased a cat in Portsmouth, which
he was told came from the coast of Guinea ; its skin was
black and wrinkled, fur bluish-grey and short, its ears
rather bare, legs long, and whole aspect peculiar. This
negro cat was fertile with common cats. On the
posite coast of Africa, at Mombas, Captain Owen, R.N.,01
states that all the cats are covered with short stiff hair
instead of fur: he gives a curious account of a cat from
Algoa Bay, which had been kept for some time on board
and could be identified with certainty; this animal was
left for only eight weeks at Mombas, but during that
short period it “underwent a complete metamorphosis,
“having parted with its sandy-coloured fur.''
” A cat
op-
89 Saeugethiere von Paraguay,' 1830,
s. 212.
90 Mem. présentés par divers Sa-
vans: Acad. Roy. des Sciences,' tom. vi.
p. 346. Gomara first noticed this fact in
1554.
91 Narrative of Voyages,' vol. ii. p.
150.

64
CHAP. I.
DOMESTIC CATS.
from the Cape of Good Hope has been described by
Desmarest as remarkable from a red stripe extending
along the whole length of its back. Throughout an im-
mense area, namely, the Malayan archipelago, Siam, Pe-
gu, and Burmah, all the cats have truncated tails about
half the proper length, often with a sort of knot at the
end. In the Caroline archipelago the cats have very long
legs, and are of a reddish-yellow colour." In China a
breed has drooping ears. At Tobolsk, according to
Gmelin, there is a red-coloured breed. In Asia, also,
we find the well-known Angora or Persian breed.
The domestic cat has run wild in several countries, and
everywhere assumes, as far as can be judged by the short
recorded descriptions a uniform character. Near Mal-
donado, in La Plata, I shot one which seemed perfectly
wild; it was carefully examined by Mr. Waterhouse, 94
who found nothing remarkable in it, excepting its great
size. In New Zealand, according to Dieffenbach, the feral
cats assume a streaky grey colour like that of wild cats;
and this is the case with the half-wild cats of the Scotch
Highlands.
We have seen that distant countries possess distinct do-
mestic races of the cat. The difference may be in part due
to descent from several aboriginal species, or at least to
crosses with them. In some cases, as in Paraguay, Mombas,
and Antigua, the differences seem due to the direct action
of different conditions of life. In other cases some slight
effect may possibly be attributed to natural selection, as
cats in many cases have largely to support themselves
and to escape diverse dangers. But man, owing to the
difficulty of pairing cats, has done nothing by methodi-
cal selection ; and probably very little by unintentional
p. 308.
92 J. Crawfurd, 'Descript. Dict. of the
Indian Islands,' p. 255. The Madagas-
car cat is said to have a twisted tail : see
Desmarest, in 'Encyclop. Nat. Mamm.,'
1820, p. 233, for some of the other breeds.
03 Admiral Lutké's Voyage, vol. iii.
94 'Zoology of the Voyage of the Bea-
gle, Mammalia,' p. 20. Dieffenbach,
Travels in New Zealand,' vol. ii. p. 185.
Ch. St. John, 'Wild Sports of the High-
lands,' 1846, p. 40.

CHAP. I.
65
THEIR VARIATION.
selection; though in each litter he generally saves the
prettiest, and values most a good breed of mouse or rat-
catchers. Those cats which have a strong tendency to
prowl after game, generally get destroyed by traps. As
cats are so much petted, a breed bearing the same rela-
tion to other cats, that lapdogs bear to larger dogs, would
have been much valued ; and if selection could have been
applied, we should certainly have had many breeds in
each long-civilized country, for there is plenty of variabil-
ity to work upon.
We see in this country considerable diversity in size,
some in the proportions of the body, and extreme variabili-
ty in colouring. I have only lately attended to this subject,
but have already heard of some singular cases of varia-
tion; one of a cat born in the West Indies toothless, and
remaining so all its life. Mr. Tegetmeier has shown me
the skull of a female cat with its canines so much devel-
oped that they protruded uncovered beyond the lips; the
tooth with the fang being .95, and the part projecting
from the gum 6 of an inch in length. I have heard of a
family of six-toed cats. The tail varies greatly in length;
I have seen a cat which always carried its tail flat on its
back when pleased. The ears vary in shape, and certain
strains, in England, inherit a pencil-like tuft of hairs,
above a quarter of an inch in length, on the tips of their
ears; and this same peculiarity, according to Mr. Blyth,
characterises some cats in India. The great variability
in the length of the tail and the lynx-like tufts of hairs on
the ears are apparently analogous to differences in certain
wild species of the genus. A much more important dif-
ference, according to Daubenton, is that the intestines
of domestic cats are wider, and a third longer, than in
wild cats of the same size; and this apparently has been
caused by their less strictly carnivorous diet.
95 Quoted by Isid. Geoffroy, 'Hist. Nat. Gén.,' tom. iii. p. 427.

66
CHAP. II,
HORSES.
CHAPTER II.
HORSES AND ASSES.
HORSE. DIFFERENCES IN THE BREEDS INDIVIDUAL VARIA-
BILITY OF DIRECT EFFECTS OF THE CONDITIONS OF LIFE
CAN WITHSTAND MUCH COLD — BREEDS MUCH MODIFIED BY SE-
LECTION - COLOURS OF THE HORSE - DAPPLING - DARK STRIPES
ON THE SPINE, LEGS, SHOULDERS, AND FOREHEAD - DUN-COLOUR-
ED HORSES MOST FREQUENTLY STRIPED STRIPES PROBABLY
DUE TO REVERSION TO THE PRIMITIVE STATE OF THE HORSE.
ASSES. -BREEDS OF COLOUR OF — LEG- AND SHOULDER-STRIPES
SHOULDER-STRIPES SOMETIMES ABSENT, SOMETIMES FORKED.
The history of the Horse is lost in antiquity. Remains
of this animal in a domesticated condition have been
found in the Swiss lake-dwellings, belonging to the lat-
ter part of the Stone period. At the present time the
number of breeds is great, as may be seen by consulting
any treatise on the Horse. Looking only to the native
ponies of Great Britain, those of the Shetland Isles,
Wales, the New Forest, and Devonshire are distinguish-
able; and so it is with each separate island in the great
Malay archipelago. Some of the breeds present great
differences in size, shape of ears, length of mane, pro-
1 Rütimeyer, 'Fauna der Pfahlbauten,'
1861, s. 122.
2 See Youаtt on the Horse: J. Law-
rence on the Horse, 1829 : W. C. L. Mar-
tin, History of the Horse,' 1845 : Col.
Ham. Smith, in Naturalist's Library,
Horses,' 1841, vol. xii. : Prof. Veith,
'Die Naturgesch. Haussäugethiere,' 1856.
3 Crawfurd, 'Descript. Dict. of Indian
Islands, 1856, p. 153.
* There are
many different breeds, every island har-
ing at least one peculiar to it." Thus in
Sumatra there are at least two breeds ;
in Achin and Batubara one; in Java sev-
eral breeds; one in Bali, Lomboc, Sum-
bawa (one of the best breels), Tambora,
Bima, Gunung-api, Celebes, Sumba, and
Philippines. Other breeds are specified
by Zollinger in the Journal of the In-
dian Archipelago,' vol. v. p. 343, &c.

CHAP. II.
67
THEIR VARIATION.
portions of the body, form of the withers and hind quar-
ters, and especially in the head. Compare the race-horse,
dray-horse, and a Shetland pony in size, configuration, and
disposition; and see how much greater the difference is
than between the six or seven other living species of
genus Equus.
Of individual variations not known to characterise
particular breeds, and not great or injurious enough to
be called monstrosities, I have not collected many cases.
Mr. G. Brown, of the Cirencester Agricultural College,
who has particularly attended to the dentition of our do-
mestic animals, writes to me that he has "several times
noticed eight permanent incisors instead of six in the
jaw.” Male horses alone properly have canines, but
they are occasionally found in the mare, though of small
size. The number of ribs is properly eighteen, but
Youatt. asserts that not unfrequently there are nineteen
on each side, the additional one being always the poste-
rior rib. I have seen several notices of variations in the
bones of the leg; thus Mr. Price speaks of an addition-
al bone in the hock, and of certain abnormal appearances
between the tibia and astragalus, as quite common in
Irish horses, and not due to disease. Horses have often
been observed, according to M. Gaudry,' to possess a
trapezium and a rudiment of a fifth metacarpal bone, so
that “one sees appearing by monstrosity, in the foot of
the horse, structures which normally exist in the foot of
the Hipparion,”-an allied and extinct animal. In vari-
ous countries horn-like projections have been observed
on the frontal bones of the horse: in one case described
by Mr. Percival they arose about two inches above the
orbital processes, and were “very like those in a calf
from five to six months old," being from half to three-
4 The Horse,' &c., by John Lawrence,
1829, p. 14.
5 The Veterinary,' London, vol, v. p.
543.
6 Proc. Veterinary Assoc., in The Ve-
terirary,' vol. xiii. p. 42.
7 Bulletin de la Soc. Géolog.,' tom.
xxii., 1866, p. 22.

68
CHAP. II.
HORSES.
quarters of an inch in length.Azara has described
two cases in South America in which the projections
were between three and four inches in length: other in-
stances have occurred in Spain.
That there has been much inherited variation in the
horse cannot be doubted, when we reflect on the number
of the breeds existing throughout the world or even
within the same country, and when we know that they
have largely increased in number since the earliest
known records. Even in so fleeting a character as co-
lour, Hofacker found that, out of two hundred and six-
teen cases in which horses of the same colour were paired,
only eleven pairs produced foals of a quite different co-
lour. As Professor Low 11 has remarked, the English
race-horse offers the best possible evidence of inheritance.
The pedigree of a race-horse is of more value in judging
of its probable success than its
appearance : King He-
gained in prizes 201,5051. sterling, and begot 497
winners; "Eclipse" begot 334 winners.
Whether the whole amount of difference between the
various breeds be due to variation is doubtful. From the
fertility of the most distinct breeds when crossed, nat-
uralists have generally looked at all the breeds as having
descended from a single species. Few will agree
with
Colonel H. Smith, who believes that they have descend-
ed from no less than five primitive and differently colour-
ed stocks, But as several species and varieties of the
rod”
13
8 Mr. Percival, of the Enniskillen Dra-
goons, in 'The Veterinary,' vol. i. p. 224:
see Azara, 'Des Quadrupèdes du Para-
guay,' tom. ii. p. 313. The French trans-
lator of Azara refers to other cases men-
tioned by Huzard as occurring in Spain.
9 Godron, 'De l'Espèce,' tom. i. p. 378.
10 'Ueber die Eigenschafter,' &c.,
1828, s. 10.
11 Domesticated Animals of the Brit-
ish Islands,' pp. 527, 532. In all the ve-
terinary treatises and papers which I
have read, the writers insist in the
strongest terms on the inheritance by
the horse of all good and bad tendencies
and qualities. Perhaps the principle of
inheritance is not really stronger in the
horse than in any other animal; but,
from its value, the tendency has been
more carefully observed.
12 Andrew Knight crossed breeds so
different in size as a dray-horse and Nor-
wegian pony ; see A. Walker on Inter-
marriage,' 1838, p. 205.
13 Naturalist's Library,' Horses, vol.
xii. p. 208.

CHAP. II.
69
THEIR VARIATION.
horse existed 14 during the later tertiary periods, and as
Rütimeyer found differences in the size and form of the
skull in the earliest known domesticated horses, 16 we
ought not to feel sure that all our breeds have descended
from a single species. As we see that the savages of
North and South America easily reclaim the feral horses,
there is no improbability in savages in various quarters
of the world having domesticated more than one native
species or natural race. No aboriginal or truly wild horse
is positively known now to exist; for it is thought by
some authors that the wild horses of the East are escaped
domestic animals. 16 If our domestic breeds have descend-
ed from several species or natural races, these apparently
have all become extinct in the wild state.
With our
present knowledge, the common view that all have de-
scended from a single species is, perhaps, the most probable.
With respect to the causes of the modifications which
horses have undergone, the conditions of life seem to
produce a considerable direct effect. Mr. D. Forbes, who
has had excellent opportunities of comparing the horses
of Spain with those of South America, informs me that
the borses of Chile, which have lived under nearly the
same conditions as their progenitors in Andalusia, remain
unaltered, whilst the Pampas horses and the Puno ponies
are considerably modified. There can be no doubt that
horses become greatly reduced in size and altered in ap-
pearance by living on mountains and islands; and this
apparently is due to want of nutritious or varied food.
Every one knows how small and rugged the ponies are on
the Northern islands and on the mountains of Europe.
Corsica and Sardinia have their native ponies; and there
14 Gervais, Hist. Nat. Mamm.,' tom.
ii. p. 143. Owen, British Fossil Mam-
mals,' p. 353.
15 Kenntniss der fossilen Pferde,'
1863, s. 131
16 Mr. W. C. L. Martin (The Horse,'
1845, p. 34), in arguing against the belief
that the wild Eastern horses are merely
feral, has remarked on the improbability
of man in ancient times having extirpated
a species in a region where it can now
exist in numbers.

70
CHAP. II.
HORSES.
were," or still are, on some islands on the coast of Virgi-
nia, ponies like those of the Shetland Islands, which are
believed to have originated through exposure to unfa-
vourable conditions. The Puno ponies, which inhabit the
lofty regions of the Cordillera, are, as I hear from Mr. D.
Forbes, strange little creatures, very unlike their Spanish
progenitors. Further south, in the Falkland Islands, the
offspring of the horses imported in 1764 have already so
much deteriorated in size 18 and strength that they are
unfitted for catching wild cattle with the lasso; so that
fresh horses have to be brought for this purpose from La
Plata at a great expense. The reduced size of the horses
bred on both southern and northern islands, and on sev-
eral mountain-chains, can hardly have been caused by the
cold, as a similar reduction has occurred on the Virginian
and Mediterranean islands. The horse can withstand in-
tense cold, for wild troops live on the plains of Siberia
under lat. 56°,10 and aboriginally the horse must have in-
habited countries annually covered with snow, for he long
retains the instinct of scraping it away to get at the herb-
age beneath. . The wild tarpans in the East have this in-
stinct; and, as I am informed by Admiral Sulivan, this is
likewise the case with the horses which have run wild on
the Falkland Islands; now this is the more remarkable as
the progenitors of these horses could not have followed
this instinct during many generations in La Plata : the
wild cattle of the Falklands never scrape away the snow,
and perish when the ground is long covered. In the
northern parts of America the horses, descended from
those introduced by the Spanish conquerors of Mexico,
have the same habit, as have the native bisons, but not so
the cattle introduced from Europe.20
17 Transact. Maryland Academy,'
vol. i. part i. p. 28.
18 Mr. Mackinnon on The Falkland
Islands,' p. 25. The average height of
the Falkland horses is said to be 14
hands 2 inches. See also my 'Journal of
Researches.'
19 Pallas, Act. Acad. St. Peters-
burgh,' 1777, part ii. p. 265. With re-
spect to the tarpans scraping away the
snow, see Col. Hamilton Smith in Nat.
Lib.,' vol. xii. p. 165.
20 Franklin's 'Narrative,' vol. i. p.
87; note by Sir J. Richardson.

CHAP. II.
71
THEIR COLOURS AND STRIPES.
22
The horse can flourish under intense heat as well as
under intense cold, for he is known to come to the high-
est perfection, though not attaining a large size, in Arabia
and northern Africa. Much humidity is apparently more
injurious to the horse than heat or cold. In the Falk-
land Islands, horses suffer much from the dampness; and
this same circumstance may perhaps partly account for
the singular fact that to the eastward of the Bay of Ben-
gal, over an enormous and humid area, in Ava, Pegu,
Siam, the Malayan archipelago, the Loo Choo Islands,
and a large part of China, no full-sized horse is found.
When we advance as far eastward as Japan, the horse
reacquires his full size.?
With most of our domesticated animals, some breeds
are kept on account of their curiosity or beauty; but the
horse is valued almost solely for its utility. Hence semi-
monstrous breeds are not preserved; and probably all the
existing breeds have been slowly formed either by the
direct action of the conditions of life, or through the se-
lection of individual differences. No doubt semi-mon-
strous breeds might have been formed; thus Mr. Water-
ton records as the case of a mare which produced succes-
sively three foals without tails, so that a tailless race
might have been formed like the tailless races of dogs and
cats. A Russian breed of horses is said to have frizzled
hair, and Azara 24 relates that in Paraguay horses are oc-
casionally born, but are generally destroyed, with hair
like that on the head of a negro; and this peculiarity is
transmitted even to half-breeds: it is a curious case of
correlation that such horses have short manes and tails,
21 Mr. J. H. Moor, Notices of the In-
dian Archipelago :' Singapore, 1887, p.
189. A pony from Java was sent ('Athe-
næum,' 1842, p. 718) to the Queen only
28 inches in height. For the Loo Choo
Islands, see Beechey's Voyage,' 4th edit.
vol. i. p. 499.
22 J. Crawford, 'History of the Horse;'
Journal of Royal United Service Institu-
tion,' vol. iv.
Essays on Natural History,' 2nd
series, p. 161.
"Quadrupèdes du Paraguay,' tom.
ii. p. 333.
234
24 4

72
CHAP. II.
HORSES.
and their hoofs are of a peculiar shape like those of a
mule.
It is scarcely possible to doubt that the long-continued
selection of qualities serviceable to man has been the
chief agent in the formation of the several breeds of the
horse. Look at a dray-horse, and see how well adapted
he is to draw heavy weights, and how unlike in appear-
ance to any allied wild animal. The English race-horse
is known to have proceeded from the commingled blood
of Arabs, Turks, and Barbs; but selection and training
have together made him a very different animal from his
parent-stocks.
As a writer in India, who evidently
knows the pure Arab well, asks, who now, “looking at
our present breed of race-horses, could have conceived that
they were the result of the union of the Arab horse and
African mare?” The improvement is so marked that in
running for the Goodwood Cup “the first descendants of
Arabian, Turkish, and Persian horses, are allowed a dis-
count of 18 lbs. weight; and when both parents are of
these countries a discount of 36 lbs.25. It is notorious
that the Arabs have long been as careful about the pedi-
gree of their horses as we are, and this implies great and
continued care in breeding. Seeing what has been done
in England by careful breeding, can we doubt that the
Arabs must likewise have produced during the course of
centuries a marked effect on the qualities of their horses?
But we may go much farther back in time, for in the most
ancient known book, the Bible, we hear of studs carefully
kept for breeding, and of horses imported at high prices
from various countries. We may therefore conclude
26
p. 181.
25 Prof. Low, 'Domesticated Animals,
p. 546. With respect to the writer in In-
dia, see 'India Sporting Review, vol. ii.
As Lawrence has remarked
("The Horse,' p. 9), "perhaps no in-
stance has ever occurred of a three-part
bred horse (i.e. a horse, one of whose
grand-parents was of impure blood) sav-
ing his distance in running two miles
with thoroughbred racers.” Some few
instances are on record of seven-eighths
racers having been successful.
26 Prof. Gervais (in his Hist. Nat.
Mamm.,' tom. ii. p. 144) has collected
many facts on this head. For instance,
Solomon (Kings, b.i.ch. x. v. 28) bought
horses in Egypt at a high price.

CHAP. II.
73
THEIR COLOURS AND STRIPES.
that, whether or not the various existing breeds of the
horse have proceeded from one or more aboriginal stocks,
yet that a great amount of change has resulted from the
direct action of the conditions of life, and probably a still
greater amount from the long-continued selection by man
of slight individual differences.
With several domesticated quadrupeds and birds, cer-
tain coloured marks are either strongly inherited or tend
to reappear after having long been lost. As this subject
will hereafter be seen to be of importance, I will give a
full account of the colouring of horses. All English
breeds, however unlike in size and appearance, and sev-
eral of those in India and the Malay archipelago, present
a similar range and diversity of colour. The English
race-horse, however, is said a' never to be dun-coloured ;
but as dun and cream-coloured horses are considered by
the Arabs as worthless, “and fit only for Jews to ride,"
these tints may have been removed by long-continued
selection. Horses of every colour, and of such widely
different kinds as dray-horses, cobs, and ponies, are all
occasionally dappled, in the same manner as is so con-
spicuous with grey horses. This fact does not throw
any clear light on the colouring of the aboriginal horse,
but is a case of analogous variation, for even asses are
sometimes dappled, and I have seen, in the British
Museum, a hybrid from the ass and zebra dappled
on its hinder quarters. By the expression analogous
variation (and it is one that I shall often have occasion
to use) I mean a variation occurring in a species or vari-
ety which resembles a normal character in another and
9 28
27 The Field,' July 13th, 1861, p. 42.
28 E. Vernon Harcourt, Sporting in
Algeria,' p. 26.
29 I state this from my own observa-
tions made during several years on the
colours of horses. I have seen cream-co-
loured, light-dun and mouse-dun horses
dappled, which I mention because it has
been stated (Martin, History of the
Horse,' p. 134) that duns are never dap-
pled. Martin (p. 205) refers to dap-
pled asses. In The Farrier' (London,
1828, pp. 453, 455) there are some good
remarks on the dappling of horses; and
likewise in Col, Hamilton Smith on The
Horse.'
4

74
CHAP. II.
HORSES.
distinct species or variety. Analogous variations may
arise, as will be explained in a future chapter, from two
or more forms with a similar constitution having been
exposed to similar conditions,-or from one of two forms
having reacquired through reversion a character inherited
by the other form from their common progenitor, -or
from both forms having reverted to the same ancestral
character. We shall immediately see that horses oc-
casionally exhibit a tendency to become striped over
a large part of their bodies; and as we know that stripes
readily pass into spots and cloudy marks in the varieties
of the domestic cat and in several feline species-even
the cubs of the uniformly-coloured lion being spotted
with dark marks on a lighter ground—we may suspect
that the dappling of the horse, which has been noticed
by some authors with surprise, is a modification or
vestige of a tendency to become striped.
This tendency in the horse to become striped is in several respects
an interesting fact. Horses of all colours, of the most diverse
breeds, in various parts of the world, often have a dark stripe ex-
tending along the spine, from the mane to the tail ; but this is so
common that I need enter into no particulars.30 Occasionally
horses are transversely barred on the legs, chiefly on the under
side; and more rarely they have a distinct stripe on the shoulder,
like that on the shoulder of the ass, or a broad dark patch repre-
senting a stripe. Before entering on any details I must premise that
the term dun-coloured is vague, and includes three groups of colour,
viz. that between cream-colour and reddish brown, which graduates
into light-bay or light-chesnut-this, I believe, is often called
fallow-dun ; secondly, leaden or slate-colour or mouse-dun, which
graduates into an ash-colour; and, lastly, dark-dun, between brown
and black. In England I have examined a rather large, lightly-
built, fallow-dun Devonshire pony (fig. 1), with a conspicuous stripe
along the back, with light transverse stripes on the under sides of
its front legs, and with four parallel stripes on each shoulder.
30 Some details are given in 'The Far-
rier; '1828, pp. 452, 455. One of the least
ponies I ever saw, of the colour of a
mouse, had a conspicuous spinal stripe.
A small Indian chesnut pony had the
same stripe, as had a remarkably heavy
chesnut cart-horse. Race-horses often
have the spinal stripe.

CHAP. II.
75
THEIR COLOURS AND STRIPES.
.
Fig. 1.-Dun Devonshire Pony, with shoulder, spinal, and leg stripes.
Of these four stripes the posterior one was very minute and faint;
the anterior one, on the other hand, was long and broad, but in-
terrupted in the middle, and truncated at its lower extremity, with
the anterior angle produced into a long tapering point. I mention
this latter fact because the shoulder-stripe of the ass occasionally
presents exactly the same appearance. I have had an outline and
description sent to me of a small, purely-bred, light fallow-dun
Welch pony, with a spinal stripe, a single transverse stripe on each
leg, and three shoulder-stripes ; the posterior stripe corresponding
with that on the shoulder of the ass was the longest, whilst the
two anterior parallel stripes, arising from the mane, decreased in
length, in a reversed manner as compared with the shoulder-stripes
on the above-described Devonshire pony. I have seen a bright
fallow-dun, strong cob, with its front legs transversely barred on
the under sides in the most conspicuous manner; also a dark-leaden
mouse-coloured pony with similar leg stripes, but much less conspi-
cuous; also a bright fallow-dun colt, fully three-parts thoroughbred,
with very plain transverse stripes on the legs; also a chesnut-dun
cart-horse with a conspicuous spinal stripe, with distinct traces of
shoulder-stripes, but none on the legs; I could add other cases.
My son made a sketch for me of a large, heavy, Belgian cart-horse,
of a fallow-dun, with a conspicuous spinal stripe, traces of leg.
stripes, and with two parallel (three inches apart) stripes about
seven or eight inches in length on both shoulders. I have seen
another rather light cart-horse, of a dirty dark cream-colour, with
striped legs, and on one shoulder a large ill-defined dark cloudy

76
CHAP. II.
HORSES.
patch, and on the opposite shoulder two parallel faint stripes. All
the cases yet mentioned are duns of various tints; but Mr. W. W.
Edwards has seen a nearly thoroughbred chesnut horse which had
the spinal stripe, and distinct bars on the legs; and I have seen
two bay carriage-horses with black spinal stripes; one of these
horses had on each shoulder a light shoulder-stripe, and the other
had a broad black ill-defined stripe, running obliquely half-way
down each shoulder; neither had leg-stripes.
The most interesting case which I have met with occurred in a
colt of my own breeding. A bay mare (descended from a dark-brown
Flemish mare by a light grey Turcoman horse) was put to Hercules,
a thoroughbred dark bay, whose sire (Kingston) and dam were both
bays. The colt ultimately turned out brown; but when only a fort-
night old it was a dirty bay, shaded with mouse-grey, and in parts
with a yellowish tint: it had only a trace of the spinal stripe, with
a few obscure transverse bars on the legs; but almost the whole
body was marked with very narrow dark stripes, in most parts so
obscure as to be visible only in certain lights, like the stripes which
may be seen on black kittens. These stripes were distinct on the
hind-quarters, where they diverged from the spine, and pointed a
little forwards; many of them as they diverged from the spine be-
came a little branched, exactly in the same manner as in some
zebrine species. The stripes were plainest on the forehead between
the ears, where they formed a set of pointed arches, one under the
other, decreasing in size downwards towards the muzzle ; exactly
similar marks may be seen on the forehead of the quagga and
Burchell's zebra. When this foal was two or three months old all
the stripes entirely disappeared. I have seen similar marks on the
forehead of a fully grown, fallow-dun, cob-like horse, having a con-
spicuous spinal stripe, and with its front legs well barred.
In Norway the colour of the native horse or pony is dun, varying
from almost cream-colour to dark mouse-dun ; and an animal is not
considered purely bred unless it has the spinal and leg stripes.31 In
one part of the country my son estimated that about a third of the
ponies had striped legs; he counted seven stripes on the fore-legs
and two on the hind-legs of one pony ; only a few of them exhibit-
ed traces of shoulder-stripes; but I have heard of a cob imported
from Norway which had the shoulder as well as the other stripes
well developed. Colonel Ham. Smith 82 alludes to dun-horses with
81 I have received information, through
the kindness of the Consul-General, Mr.
J. R. Crowe, from Prof. Boeck, Rasck, and
Esmarck, on the colours of the Norwegian
ponies. See, also, The Field,' 1861, p.
431.
32 Col. Ham. Smith, Nat. Lib.,' vol,
xii. p. 275.

CEAP. II.
77
THEIR COLOURS AND STRIPES.
the spinal stripe in the Sierras of Spain ; and the horses originally
derived from Spain, in some parts of South America, are now duns.
Sir W. Elliot informs me that he inspected a herd of 300 South
American horses imported into Madras, and many of these had
transverse stripes on the legs and short shoulder-stripes; the most
strongly marked individual, of which a coloured drawing was sent
me, was a mouse-dun, with the shoulder-stripes slightly forked.
In the North-Western parts of India striped horses of more than
one breed are apparently commoner than in any other part of the
world ; and I have received information respecting them from sev-
eral officers, especially from Colonel Poole, Colonel Curtis, Major
Campbell, Brigadier St. John, and others. The Katty war horses
are often fifteen or sixteen hands in height, and are well but lightly
built. They are of all colours, but the several kinds of duns pre-
vail ; and these are so generally striped, that a horse without stripes
is not considered pure. Colonel Poole believes that all the duns
have the spinal stripe, the leg-stripes are generally present, and he
thinks that about half the horses have the shoulder-stripe; this
stripe is sometimes double or treble on both shoulders. Colonel
Poole has often seen stripes on the cheeks and sides of the nose.
He has seen stripes on the grey and bay Kattywars when first foaled,
but they soon faded away. I have received other accounts of cream-
coloured, bay, brown, and grey Kattywar horses being striped.
Eastward of India, the Shan (north of Burmah) ponies, as I am in-
formed by Mr. Blyth, have spinal, leg, and shoulder stripes. Sir
W. Elliot informs me that he saw two bay Pegu ponies with leg-
stripes. Burmese and Javanese ponies are frequently dun-coloured,
and have the three kinds of stripes, " in the same degree as in Eng-
land.” 33 Mr. Swinhoe informs me that he examined two light-dun
ponies of two Chinese breeds, viz. those of Shanghai and Amoy ; both
had the spinal stripe, and the latter an indistinct shoulder-stripe.
We thus see that in all parts of the world breeds of the horse as
different as possible, when of a dun-colour (including under this
term a wide range of tint from cream to dusky black), and rarely
when of bay, grey, and chesnut shades, have the several above-
specified stripes. Horses which are of a yellow colour with white
mane and tail, and which are sometimes called duns, I have never
seen with stripes. 34
From reasons which will be apparent in the chapter on Reversion,
33 Mr. G. Clark, in Annal and Mag. of
Nat. History,' 2nd series, vol. ii., 1848, p.
363. Mr. Wallace informs me that he
saw in Java a dun and clay-coloured
horse with spinal and leg stripes.
34 See, also, on this point, The Field,'
July 27th, 1861, p. 91.

78
CHAP. II.
HORSES.
I have endeavoured, but with poor success, to discover whether duns,
which are so much oftener striped than other coloured horses, are
ever produced from the crossing of two horses, neither of which are
duns. Most persons to whom I have applied believe that one parent
must be a dun; and it is generally asserted, that, when this is the
case, the dun-colour and the stripes are strongly inherited.35 One
case has fallen under my own observation of a foal from a black
mare by a bay horse, which when fully grown was a dark fallow-
dun and had a narrow but plain spinal stripe. Hofacker 36 gives
two instances of mouse-duns (Mausrapp) being produced from two
parents of different colours and neither duns.
I have also endeavoured with little success to find out whether
the stripes are generally plainer or less plain in the foal than in the
adult horse. Colonel Poole informs me that, as he believes, "the
stripes are plainest when the colt is first foaled; they then become
less and less distinct till after the first coat is shed, when they come
out as strongly as before ; but certainly often fade away as the age
of the horse increases.” Two other accounts confirm this fading of
the stripes in old horses in India. One writer, on the other hand,
states that colts are often born without stripes, but that they appear
as the colt grows older. Three authorities affirm that in Norway
the stripes are less plain in the foal than in the adult. Perhaps
there is no fixed rule. In the case described by me of the young foal
which was narrowly striped over nearly all its body, there was no
doubt about the early and complete disappearance of the stripes.
Mr. W. W. Edwards examined for me twenty-two foals of race-
horses, and twelve had the spinal stripe more or less plain ; this
fact, and some other accounts which I have received, lead me to
believe that the spinal stripe often disappears in the English race-
horse when old. On the whole I infer that the stripes are generally
plainest in the foal, and tend to disappear in old age.
The stripes are variable in colour, but are always dark-
er than the rest of the body. They do not by any means
always coexist on the different parts of the body: the
legs may be striped without any shoulder-stripe, or the
converse case, which is rarer, may occur; but I have
never heard of either shoulder or leg-stripes without the
spinal stripe. The latter is by far the commonest of all
the stripes, as might have been expected, as it character-
35 "The Field,' 1861, pp. 431, 493, 545.
36 Ueber die Eigenschaften,' &c., 1828, s. 13, 14.

CHAP. II.
79
THEIR COLOURS AND STRIPES.
ises the other seven or eight species of the genus. It is
remarkable that so trifling a character as the shoulder-
stripe being double or triple should occur in such different
breeds as Welch and Devonshire ponies, the Shan pony,
heavy cart-horses, light South American horses, and the
lanky Kattywar breed. Colonel Hamilton Smith be-
lieves that one of his five supposed primitive stocks was
dun-coloured and striped; and that the stripes in all the
other breeds result from ancient crosses with this one
primitive dun; but it is extremely improbable that differ-
ent breeds living in such distant quarters of the worla
should all have been crossed with any one aboriginally
distinct stock. Nor have we any reason to believe that
the effects of a cross at a very remote period could be
propagated for so many generations as is implied on this
view.
With respect to the primitive colour of the horse hav-
ing been dun, Colonel Hamilton Smith " has collected a
large body of evidence showing that this tint was com-
mon in the East as far back as the time of Alexander,
and that the wild horses of Western Asia and Eastern
Europe now are, or recently were, of various shades of
dun. It seems that not very long ago a wild breed of
dun-coloured horses with a spinal stripe was preserved in
the royal parks in Prussia. I hear from Hungary that
the inhabitants of that country look at the duns with a
spinal stripe as the aboriginal stock, and so it is in Nor-
way. Dun-coloured ponies are not rare in the mountain-
ous parts of Devonshire, Wales, and Scotland, where the
aboriginal breed would have had the best chance of being
preserved. In South America in the time of Azara, when
the horse had been feral for about 250 years, 90 out of
37 Naturalist's Library,' vol. xii.
(1841), pp. 109, 156 to 163, 280, 281.
Cream-colour, passing into Isabella (i.e.
the colour of the dirty linen of Queen
Isabella), seems to have been common
in ancient times. See also Pallas's ac-
count of the wild horses of the East, who
speaks of dun and brown as the preva-
lent colours.

80
CHAP. II.
HORSES.
100 horses were “bai-châtains," and the remaining ten
were “zains,” and not more than one in 2000 black. Zain
is generally translated as dark without any white; but
as Azara speaks of mules being “zain-clair," I suspect
that zain must have meant dun-coloured. In some parts
of the world feral horses show a strong tendency to be-
38
come roans.
In the following chapters on the Pigeon we shall see
that in pure breeds of various colours, when a blue bird
is occasionally produced, certain black marks invariably
appear on the wings and tail; so again, when variously
coloured breeds are crossed, blue birds with the same
black marks are frequently produced. We shall further
see that these facts are explained by, and afford strong
evidence in favour of, the view that all the breeds are
descended from the rock-pigeon, or Columba livia, which
is thus coloured and marked. But the appearance of the
stripes on the various breeds of the horse, when of a dun-
colour, does not afford nearly such good evidence of their
descent from a single primitive stock as in the case of
the pigeon; because no certainly wild horse is known as
a standard of comparison; because the stripes when they
do appear are variable in character; because there is far
from sufficient evidence of the appearance of the stripes
from the crossing of distinct breeds; and lastly, because
all the species of the genus Equus have the spinal stripe,
and several have shoulder and leg stripes. Nevertheless
the similarity in the most distinct breeds in their general
range of colour, in their dappling, and in the occasional
38 Azara, 'Quadrupèdes du Paraguay,'
tom. ii. p. 307; for the colour of mules,
see p. 350. In North America, Catlin,
(vol. ii. p. 57) describes the wild horses,
believed to have descended from the
Spanish horses of Mexico, as of all col-
ours, ack, grey, roan, and roan pied
with sorrel. F. Michaux (Travels in
North America,' Eng translat., p. 235)
describes two wild horses from Mexico
as roan. In the Falkland Islands, where
the horse has been feral only between 60
and 70 years, I was told that roans and
iron-greys were the prevalent colours.
These several facts show that horses do
not generally revert to any uniform
colour.

CHAP. II.
81
ASSES.
appearance, especially in duns, of leg-stripes and of double
or triple shoulder-stripes, taken together, indicate the
probability of the descent of all the existing races from a
single, dun-coloured, more or less striped, primitive stock,
to which our horses still occasionally revert.
THE Ass.
39
Four species of Asses, besides three of zebras, have been
described by naturalists; but there can now be little
doubt that our domesticated animal is descended from
one alone, namely, the Asinus tæniopus of Abyssinia.
The Ass is sometimes advanced as an instance of an ani-
mal domesticated, as we know by the Old Testament, from
an ancient period, which has varied only in a very slight
degree. But this is by no means strictly true; for in
Syria alone there are four breeds;49 first, a light and
graceful animal, with an agreeable gait, used by ladies ;
secondly, an Arab breed reserved exclusively for the sad-
dle; thirdly, a stouter animal used for ploughing and va-
rious purposes ; and lastly, the large Damascus breed,
with a peculiarly long body and ears. In this country, and
generally in Central Europe, though the ass is by no
means uniform in appearance, it has not given rise to dis-
tinct breeds like those of the horse. This may probably
be accounted for by the animal being kept chiefly by poor
persons, who do not rear large numbers, nor carefully match
and select the
young For, as we shall see in a future
chapter, the ass can with ease be greatly improved in size
and strength by careful selection, combined no doubt
with good food; and we may infer that all its other cha-
racters would be equally amenable to selection. The
small size of the ass in England and Northern Europe is
apparently due far more to want of care in breeding than
89 Dr. Sclater, in 'Proc. Zoolog. Soc.,'
1862, p. 164.
40 W. C. Martin, History of the
Horse,' 1845, p. 207.

82
CHAP. II.
ASSES
> 41
to cold; for in Western India, where the ass is used as a
beast of burden by some of the lower castes, it is not
much larger than a Newfoundland dog," being generally
not more than from twenty to thirty inches high."
The ass varies greatly in colour; and its legs, especially
the fore-legs, both in England and other countries—for
instance, in China—are occasionally barred transversely
more plainly than those of dun-coloured horses. With
the horse the occasional appearance of leg-stripes was ac-
counted for, through the principle of reversion, by the
supposition that the primitive horse was thus striped ;
with the ass we may confidently advance this explanation,
for the parent-form, the A. toniopus, is known to be
barred, though only in a slight degree, across the legs.
The stripes are believed to occur most frequently and to
be plainest on the legs of the domestic ass during early
youth, as is apparently likewise the case with the horse.
The shoulder-stripe, which is so eminently characteristic
of the species, is nevertheless variable in breadth, length,
and manner of termination. I have measured a shoulder-
stripe four times as broad as another; and some more than
twice as long as others. In one light-grey ass the shoul-
der-stripe was only six inches in length, and as thin as a
piece of string; and in other animal of the same colour
there was only a dusky shade representing a stripe. I
have heard of three white asses, not albinoes, with no
trace of shoulder or spinal stripes ; and I have seen nine
other asses with no shoulder-stripe, and some of them
had no spinal stripe. Three of the nine were light-greys,
one a dark-grey, another grey passing into reddish-roan,
and the others were brown, two being tinted on parts of
their bodies with a reddish or bay-shade. Hence we may
41 Col. Sykes' Cat. of Mammalia,
Proc. Zoolog. Soc.,' July 12th, 1831.
Williamson, Oriental Field Sports,' vol,
ii., quoted by Martin, p. 206.
42 Blyth, in Charlesworth's Mag. of
Nat. Hist.,' vol. iv., 1840, p. 83. I have
also been assured by a breeder that this
is the case.
43 One case is given by Martin, The
Horse,' p. 205.

CHAP. II.
83
THEIR COLOURS AND STRIPES.
44
conclude that, if grey and reddish-brown asses had been
steadily selected and bred from the shoulder-stripe would
have been almost as generally and as completely lost as
in the case of the horse.
The shoulder-stripe on the ass is sometimes double, and
Mr. Blyth has seen even three or four parallel stripes.
I have observed in ten cases shoulder-stripes abruptly
truncated at the lower end, with the anterior angle pro-
duced into a tapering point, precisely as has been figured
in the dun Devonshire pony. I have seen three cases of
the terminal portion abruptly and angularly bent; and
two cases of a distinct though slight forking. In Syria,
Dr. Hooker and his party observed for me no less than
five instances of the shoulder-stripe being plainly forked
over the fore leg. In the common mule it is likewise
sometimes forked. When I first noticed the forking and
angular bending of the shoulder-stripe, I had seen enough
of the stripes in the various equine species to feel con-
vinced that even a character so unimportant as this had
a distinct meaning, and was thus led to attend to the
subject. I now find that in the Asinus Burchellii and
quagga, the stripe which corresponds with the shoulder-
stripe of the ass, as well as some of the stripes on the
neck, bifurcate, and that some of those near the shoulder
have their extremities angularly bent backwards. The
forking and angular bending of the stripes on the shoul-
ders apparently stand in relation with the changed direc-
tion of the nearly upright stripes on the sides of the body
and neck to the transverse bars on the legs. Finally we
see that the presence of shoulder, leg, and spinal stripes
in the horse,—their occasional absence in the ass,—the oc-
currence of double and triple shoulder-stripes in both ani-
mals, and the similar manner in which these stripes termi-
nate at their lower extremities,--are all cases of analogous
44 Journal As. Soc. of Bengal,' vol. xxviii. 1860, p. 231. Martin on the Horse,
p. 205.

84
CHAP. II.
ASSES.
variation in the horse and ass. These cases are probably
not due to similar conditions acting on similar constitu-
tions, but to a partial reversion in colour to the common
progenitor of these two species, as well as of the other
species of the genus. We shall hereafter have to return
to this subject, and discuss it more fully.

CHAP. III.
85
DOMESTIC PIGS.
CHAPTER III.
PIGS -- CATTLE - SHEEP - GOATS.
PIGS BELONG TO TWO DISTINCT TYPES, SUS SCRÓFA AND INDICA
— TORF-SCHWEIN - JAPAN PIG — FERTILITY OF CROSSED PIGS
CHANGES IN THE SKULL OF THE HIGHLY CULTIVATED RACES
CONVERGENCE OF CHARACTER GESTATION SOLID-HOOFED
SWINE - CURIOUS APPENDAGES TO THE JAWS-DECREASE IN
SIZE OF THE TUSKS-YOUNG PIGS LONGITUDINALLY STRIPED -
- FERAL PIGS - CROSSED BREEDS.
CATTLE. - ZEBU A DISTINCT SPECIES — EUROPEAN CATTLE PRO-
BABLY DESCENDED FROM THREE WILD FORMS - ALL THE RACES
NOW FERTILE TOGETHER — BRITISH PARK CATTLE — ON THE CO-
LOUR OF THE ABORIGINAL SPECIES — CONSTITUTIONAL DIFFER-
ENCES - 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 - GES-
TATION OF — CHANGES IN THE WOOL -- SEMI-MONSTROUS BREEDS.
GOATS. — REMARKABLE VARIATIONS OF.
The breeds of the pig have recently been more closely
studied, though much still remains to be done, than
those of almost any other domesticated animal. This
has been effected by Hermann von Nathusius in two
admirable works, especially in the later one on the Skulls
of the several races, and by Rütimeyer in his celebrated
Fauna of the ancient Swiss lake-dwellings.' Nathusius
has shown that all the known breeds may be divided in
two great groups: one resembling in all important re-
1 Hermann von Nathusius, 'Die Racen
des Schweines, 'Berlin, 1860; and 'Vor-
studien fur Geschichte,' &c., 'Schweine-
schädel,' Berlin, 1864. Rütimeyer, "Die
Fauna der Pfahlbauten,' Basel, 1861.

86
CHAP. III.
DOMESTIC PIGS.
spects and no doubt descended from the common wild
boar; so that this may be called the Sus scrofa group.
The other
group. differs in several important and con-
stant osteological characters; its wild parent-form is
unknown; the name given to it by Nathusius, accord-
ing to the law of priority, is Sus Indica of Pallas. This
name must now be followed, though an unfortunate one,
as the wild aboriginal does not inhabit India, and the best-
known domesticated breeds have been imported from Siam
and China.
Firstly, the Sus scrofa breeds, or those resembling the
common wild boar. These still exist, according to Na-
thusius (Schweineschädel, s. 75), in various parts of cen-
tral and northern Europe; formerly every kingdom, and
almost every province in Britain, possessed its own na-
tive breed; but these are now everywhere rapidly disap-
pearing, being replaced by improved breeds crossed with
the S. Indica form. The skull in the breeds of the S.
scrofa type resembles, in all important respects, that of
the European wild boar; but it has become (Schweine-
schädel, s. 63-68) higher and broader relatively to its
length; and the hinder part is more upright. The dif-
ferences, however, are all variable in degree. The breeds
which thus resemble S. scrofa in their essential skull-
characters differ conspicuously from each other in other
respects, as in the length of the ears and legs, curvature
of the ribs, colour, hairiness, size and proportions of the
body.
The wild Sus scrofa has a wide range, namely, Europe,
North Africa, as identified by osteological characters by
Rütimeyer, and Hindostan, as similarly identified by Na-
thusius. But the wild boars inhabiting these several coun-
tries differ so much from each other in external characters,
that they have been ranked by some naturalists as speci-
2 Nathusius, Die Racen des Schweines,
Berlin, 1860. An excellent appendix is
given with references to published and
trustworthy drawings of the breeds of
each country.

CHAP. III.
87
THEIR PARENTAGE.
fically distinct. Even within Hindostan these animals,
according to Mr. Blyth, form very distinct races in the
different districts; in the N. Western provinces, as I am
informed by the Rev. R. Everest, the boar never exceeds
36 inches in height, whilst in Bengal one has been mea-
sured 44 inches in height. In Europe, Northern Africa,
and Hindostan, domestic pigs have been known to cross
with the wild native species; and in Hindostan an ac-
curate observer, Sir Walter Elliot, after describing the
differences between wild Indian and wild German boars,
remarks that “the same differences are perceptible in the
domesticated individuals of the two countries." We may
therefore conclude that the breeds of the Sus scrofa type
have either descended from, or been modified by cross-
ing with, forms which may be ranked as geographical
races, but which are, according to some naturalists, dis-
tinct species.
Pigs of the Sus Indica type are best known to Eng-
lishmen under the form of the Chinese breed. The skull
of S. Indica, as described by Nathusius, differs from that
of S. scrofa in several minor respects, as in its greater
breadth and in some details in the teeth; but chiefly in
the shortness of the lachrymal bones, in the greater width
of the fore part of the palate-bones, and in the divergence
of the premolar teeth. It deserves especial notice that
these latter characters are not gained, even in the least
degree, by the domesticated forms of S. scrofa. After
reading the remarks and descriptions given by Nathu-
sius, it seems to me to be merely playing with words to
doubt whether S. Indica ought to be ranked as a spe-
cies; for the above-specified differences are more strongly
3 For Europe, see Bechstein, Natur-
gesch. Deutschlands,' 1801, b. i., s. 505.
Several accounts have been published
on the fertility of the offspring from
wild and tame swine. See Burdach's
Physiology,' and Godron, De l'Es-
pèce,' tom. i. p. 370. For Africa, Bull.
de la Soc. d'Acclimat.,' tom. iv. p. 389.
For India, see Nathusius, Schweine-
schädel,' s. 148.
4 Sir W. Elliot, Catalogue of Mamma-
lia, Madras Journal of Lit. and Sci-
ence,' vol. x. p. 219.

88.
CHAP. III.
DOMESTIC PIGS.
marked than any that can be pointed out between, for
instance, the fox and the wolf, or the ass and the horse.
As already stated, S. Indica is not known in a wild state;
but its domesticated forms, according to Nathusius, come
near to S. vittatus of Java and some allied species. A
pig found wild in the Aru islands (Schweineschädel, s.
169) is apparently identical with S. Indica ; but it is
doubtful whether this is a truly native animal. The do-
mesticated breeds of China, Cochin-China, and Siam be-
long to this type. The Roman or Neapolitan breed, the
Andalusian, the Hungarian, and the “Krause " swine of
Nathusius, inhabiting south-eastern Europe and Turkey,
and having fine curly hair, and the small Swiss "Bündt-
nerschwein” of Rütimeyer, all agree in their more impor-
tant skull characters with S. Indica, and, as is supposed,
have all been largely crossed with this form. Pigs of
this type have existed during a long period on the shores
of the Mediterranean, for a figure (Schweineschädel, s.
142) closely resembling the existing Neapolitan pig has
been found in the buried city of Herculaneum.
Rütimeyer has made the remarkable discovery that
there lived contemporaneously in Switzerland, during the
later Stone or Neolithic period, two domesticated forms,
the S. scrofa, and the S. scrofa palustris or Torfschwein.
Rütimeyer perceived that the latter approached the East-
ern breeds, and, according to Nathusius, it certainly be-
longs to the S. Indica group; but Rütimeyer has subse-
quently shown that it differs in some well-marked cha-
racters. This author was formerly convinced that his
Torfschwein existed as a wild animal during the first
part of the Stone period, and was domesticated during
a later part of the same period. Nathusius, whilst he
fully admits the curious fact first observed by Rüti-
meyer, that the bones of domesticated and wild animals
can be distinguished by their different aspect, yet, from
5 Pfahlbauten,' s. 163 et passim.

CHAP. III.
89
THEIR PARENTAGE.
special difficulties in the case of the bones of the pig
(Schweineschädel, s. 147), is not convinced of the truth
of this conclusion; and Rütimeyer himself seems now to
feel some doubt. As the Torfschwein was domesticated
at so early a period, and as its remains have been found
in several parts of Europe, belonging to various historic
and prehistoric ages, and as closely allied forms still
exist in Hungary and on the shores of the Mediterra-
nean, one is led to suspect that the wild s. Indica for-
merly ranged from Europe to China, in the same manner
as S. scrofa now ranges from Europe to Hindostan. Or,
as Rütimeyer apparently suspects, a third allied species
may formerly have lived in Europe and Eastern Asia.
Several breeds, differing in the proportions of the body,
in the length of the ears, in the nature of the hair, in co-
lour, &c., come under the S. Indica type. Nor is this
surprising, considering how ancient the domestication
of this form has been both in Europe and in China. In
this latter country the date is believed by an eminent
Chinese scholar' to go back at least 4900 years from the
present time. This same scholar alludes to the existence
of
many local varieties of the pig in China; and at the
present time the Chinese take extraordinary pains in feed-
ing and tending their pigs, not even allowing them to
walk from place to place. Hence the Chinese breed, as
Nathusius has remarked, displays in an eminent degree
the characters of a highly-cultivated race, and hence, no
doubt, its high value in the improvement of our Euro-
pean breeds. Nathusius makes a remarkable statement
(Schweineschädel, s. 138), that the infusion of the band,
or even of the cath, part of the blood of S. Indica into a
breed of S. scrofa, is sufficient plainly to modify the skull
of the latter species. This singular fact may perhaps be
8 See Rütimeyer's Neue Beitrage, ....
Torfschweine, Verh. Naturfor. Gesell. in
Basel, iv. i., 1865, s. 139.
7 Stan. Julien, quoted by De Blain-
ville, Ostéographie, p. 163.
8 Richardson, 'Pigs, their Origin,' &c.,
p. 26.
9. Die Racen des Schweines,' s. 47, 64.

90
CHAP. III.
DOMESTIC PIGS.
accounted for by several of the chief distinctive charac-
ters of S. Indica, such as the shortness of the lachrymal
bones, etc., being common to several of the species of the
genus; for in crosses the characters which are common
to many species apparently tend to be prepotent over
those appertaining to only a few species.
The Japan pig (S. pliciceps of Gray), which has been
recently exhibited in the Zoological Gardens, has an ex-
traordinary appearance from its short head, broad fore-
and rose, great fleshy ears, and deeply furrowed skin.
The following woodcut is copied from that given by Mr.
Fig. 2.-Head of Japan or Masked Pig. (Copied from Mr. Bartlett's paper in Proc.
Zoolog. Soc, 1861, p. 263.)
Bartlett. Not only is the face furrowed, but thick folds
10 Proc. Zoolog. Soc.,'1861, p. 263.

CHAP. III.
91
THEIR VARIATION.
of skin, which are harder than the other parts, almost
like the plates on the Indian rhinoceros, hang about the
shoulders and rump. It is coloured black, with white
feet, and breeds true. That it has long been domesticated
there can be little doubt; and this might have been in-
ferred even from the fact that its young are not longitu-
dinally striped; for this is a character common to all the
species included within the genus Sus and the allied gen-
era whilst in their natural state.Dr. Gray 12 has de-
scribed the skull of this animal, which he ranks not only
as a distinct species, but places it in a distinct section of
the genus. Nathusius, however, after his careful study
of the whole group, states positively (Schweineschädel, s.
153-158) that the skull in all essential characters closely
resembles that of the short-eared Chinese breed of the S.
Indica type. Hence Nathusius considers the Japan pig
as only a domesticated variety of S. Indica : if this really
be the case, it is a wonderful instance of the amount of
modification which can be effected under domestication.
Formerly there existed in the central islands of the Pa-
cific Ocean a singular breed of pigs. These are described
by the Rev. D. Tyerman and G. Bennett 13 as of small size,
hump-backed, with a disproportionately long head, with
short ears turned backwards, with a bushy tail not more
than two inches in length, placed as if it grew from the
back. Within half a century after the introduction into
these islands of European and Chinese pigs, the native
breed, according to the above authors became almost
completely lost by being repeatedly crossed with them.
Secluded islands, as might have been expected, seem fa-
vourable for the production or retention of peculiar breeds;
thus, in the Orkney Islands, the hogs have been described
as very small, with erect and sharp ears, and “with an
11 Sclater, in 'Proc. Zoolog. Soc.,' Feb.
26th, 1861.
12 Proc. Zoolog. Soc.,' 1962, p. 13.
13 Journal of Voyages and Travels
from 1921 to 1829,' vol. i. p. 300.

92
CHAP. III.
DOMESTIC PIGS.
appearance altogether different from the hogs brought
from the south.” 14
Seeing how different the Chinese pigs belonging to the
Sus Indica type, are in their osteological characters and
in external appearance from the pigs of the S. scrofa type,
so that they must be considered specifically distinct, it
is a fact well deserving attention, that Chinese and com-
mon pigs have been repeatedly crossed in various manners,
with unimpaired fertility. One great breeder who had
used pure Chinese pigs assured me that the fertility of the
half-breeds inter se and of their recrossed progeny was
actually increased; and this is the general belief of agri-
culturists. Again, the Japan pig or S. pliciceps of Gray
is so distinct in appearance from all common pigs, that it
stretches one's belief to the utmost to admit that it is
simply a domestic variety; yet this breed has been found
perfectly fertile with the Berkshire breed; and Mr. Eyton
informs me that he paired a half-bred brother and sister
and found them quite fertile together.
The modifications of the skull in the most highly cul-
tivated races are wonderful. To appreciate the amount
of change, Nathusius' work, with its excellent figures,
should be studied. The whole of the exterior of the skull
in all its parts has been altered; the hinder surface, in-
stead of sloping backwards, is directed forwards, entail-
ing many changes in other parts; the front of the head
is deeply concave; the orbits have a different shape; the
auditory meatus has a different direction and shape; the
incisors of the upper and lower jaws do not touch each
other, and they stand in both jaws above the plane of the
molars; the canines of the upper jaw stand in front of
those of the lower jaw, and this is a remarkable anomaly:
the articular surfaces of the occipital condyles are so
greatly changed in shape, that, as Nathusius remarks (s.
14 Rev. G. Low, 'Fauna Orcadensis,' p. 10. See also Dr. Hibbert's account of the
pig of the Shetland Islands.

CHAP. III.
93
THEIR VARIATION.
133), no naturalist, seeing this important part of the skull
by itself, would suppose that it belonged to the
genus
Sus. These and various other modifications, as Nathusius
observes, can hardly
be considered as
monstrosities, for
they are not injuri-
ous, and are strictly
inherited. The
whole head is much
shortened ;' thus,
whilst in common
breeds its length to
that of the body is
as 1 to 6, in the
cultur-races” the
proportion is as 1 to
9, and even recently
as 1 to 11.15 The fol-
lowing woodcut 16 of
the head of a wild
boar and of a sow
from a photograph
of the
the Yorkshire
Large Breed, may
aid in showing how
greatly the head in
a highly cultivated
race has been modi-
fied and shortened.
Nathusius has
Fig. 3.-Head of Wild Boar, and of “Golden Days,"
well discussed the
a pig of the Yorkshire Large Breed; the latter causes of the re-
from a photograph. (Copied from Sidney's edit.
markable changes
of The Pig,' by Youatt.)
15 Die Racen des Schweines,' s. 70.
16 These woodcuts are copied from en-
gravings given in Mr. S. Sidney's excel-
lent edition of The Pig,' by Youatt. See
pp. 1, 16, 19.

94
CHAP. II.
DOMESTIC PIGS.
17
in the skull and shape of the body which the highly cul-
tivated races have undergone. These modifications oc-
cur chiefly in the pure and crossed races of the S. Indica
type; but their commencement may be clearly detected
in the slightly improved breeds of the S. scrofa type."
Nathusius states positively (s. 99, 103), as the result of
common experience and of his experiments, that rich and
abundant food, given during youth, tends by some direct
action to make the head broader and shorter; and that
poor food works a contrary result. He lays much stress
on the fact that all wild and semi-domesticated pigs,
in ploughing up the ground with their muzzles, have,
whilst young, to exert the powerful muscles fixed to
the hinder part of the head. In highly cultivated races
this habit is no longer followed, and consequently the
back of the skull becomes modified in shape, entailing
other changes in other parts. There can hardly be a
doubt that so great a change in habits would affect
the skull; but it seems rather doubtful how far this
will account for the greatly reduced length of the skull
and for its concave front. It is well known (Nathu-
sius himself advancing many cases, s. 104) that there is
a strong tendency in many domestic animals—in bull-
and pug-dogs, in the niata cattle, in sheep, in Polish
fowls, short-faced tumbler pigeons, and in one variety of
the carp—for the bones of the face to become greatly
shortened. In the case of the dog, as H. Müller has
shown, this seems caused by an abnormal state of the pri-
mordial cartilage. We may, however, readily admit that
abundant and rich food supplied during many generations
would give an inherited tendency to increased size of
body, and that, from disuse, the limbs would become finer
and shorter, 18 We shall in a future chapter also see that
the skull and limbs are apparently in some manner cor-
17 Schweineschädel,' s. 74, 135.
18 Nathusius, ' Die Racen des Schweines,' s. 71.

CHAP. III.
95
THEIR VARIATION.
related, so that any change in the one tends to affect the
other.
Nathusius has remarked, and the observation is an
interesting one, that the peculiar form of the skull and
body in the most highly cultivated races is not charac-
teristic of any one race, but is common to all when im-
proved up to the same standard. Thus the large-bodied,
long-eared, English breeds with a convex back, and the
small-bodied, short-eared, Chinese breeds with a concave
back, when bred to the same state of perfection, nearly
resemble each other in the form of the head and body.
This result, it appears, is partly due to similar causes of
change acting on the several races, and partly to man
breeding the pig for one sole purpose, namely, for the
greatest amount of flesh and fat; so that selection has
always tended towards one and the same end. With
most domestic animals the result of selection has been
divergence of character, here it has been convergence.
The nature of the food supplied during many genera-
tions has apparently affected the length of the intestines;
for, according to Cuvier,° their length to that of the body
in the wild boar is as 9 to 1,-in the common domestic
boar as 13.5 to 1,--and in the Siam breed as 16 to 1. In
this latter breed the greater length may be due either to
descent from a distinct species or to more ancient domes-
tication. The number of mammæ vary, as does the pe-
riod of gestation. The latest authority says
21 that “the
period averages from 17 to 20 weeks,” but I think there
must be some error in this statement: in M. Tessier's
observations on 25 sows it varied from 109 to 123 days.
The Rev. W. D. Fox has given me ten carefully recorded
cases with well-bred pigs, in which the period varied from
101 to 116 days. According to Nathusius the period is
19
19 Die Racen des Schweines,' s. 47.
Schweineschädel,' s. 104. Compare,
also, the figures of the old Irisd and the
improved Irish breeds in Richardson on
The Pig,' 1847.
20 Quoted by Isid. Geoffroy, 'Hist.
Nat. Gén.,' tom. iii. p. 441.
21 S. Sidney, The Pig,' p. 61.

96
CHAP. III
DOMESTIC PIGS.
shortest in the races which come early to maturity; but
in these latter the course of development does not appear
to be actually shortened, for the young animal is born,
judging from the state of the skull, less fully developed,
or in a more embryonic condition,22 than in the case of
common swine, which arrive at maturity at a later age.
In the highly cultivated and early matured races, the
teeth, also, are developed earlier.
The difference in the number of the vertebræ and ribs
in different kinds of pigs, as observed by Mr. Eyton,"
and as given in the following table, has often been
quoted. The African sow probably belongs to the S.
scrofa type; and Mr. Eyton informs me that, since the
publication of his paper, cross-bred animals from the
African and English races were found by Lord Hill to
be perfectly fertile.
29
English
Long-legged
Male.
African
Female.
Chinese
Male.
French
Wild Boar, Domestic
from Cu- Boar, from
vier.
Cuvier.
14
15
6
13
6
15
4
14
5
Dorsal vertebræ..
Lumbar ..
Dorsal and lum-?
bar together ..
Sacral ..
21
5
19
5
19
4
19
4
19
4.
Total number)
of vertebræ ..
26
24
23
23
23
Some semi-monstrous breeds deserve notice. From the
time of Aristotle to the present time solid-hoofed swine
have occasionally been observed in various parts of the
world. Although this peculiarity is strongly inherited,
92 Schweineschädel,' s. 2, 20.
29 Proc. Zoolog. Soc.,' 1837, p. 23.
I have not given the caudal vertebræ,
as Mr. Eyton says some might possibly
have been lost. I have added together
the dorsal and lumbar vertebræ, owing
to Prof. Owen's remarks ('Journal Linn.
Soc.,' vol. ii. p. 28) on the difference
between dorsal and lumbar vertebræ
depending only on the development of
the ribs. Nevertheless the difference in
the number of the ribs in pigs de rves
notice.

CHAP. III.
97
THEIR VARIATION.
it is hardly probable that all the animals with solid hoofs
have descended from the same parents; it is more proba-
ble that the same peculiarity has reappeared at various
times and places. Dr. Struthers has lately described and
figured as the structure of the feet; in both front and hind
feet the distal phalanges of the two greater toes are re-
presented by a single, great, hoof-bearing phalanx ; and
in the front feet, the middle phalanges are represented by
a bone which is single towards the lower end, but bears
two separate articulations towards the upper end. From
other accounts it appears that an intermediate toe is
likewise sometimes superadded.
Another curious anomaly is offered by the appendages,
described by M. Eudes-Deslongchamps as often character-
izing the Normandy pigs. These appendages are always
attached to the same spot, to the corners of the jaw; they
are cylindrical, about three inches in length, covered with
bristles, and with a pencil of bristles rising out of a sinus
on one side: they have a cartilaginous centre, with two
Fig. 4.-Old Irish Pig, with jaw-appendages. (Copied from H. D. Richardson on Pigs.)
24 'Edinburgh New Philosoph. Journal,'
April, 1863. See also De Blainville's 'Os-
téographie,'p. 123, for various authorities
on this subject.
5

98
CHAP. III.
DOMESTIC PIGS.
small longitudinal muscles; they occur either symmetri-
cally on both sides of the face or on one side alone.
Richardson figures them on the gaunt old “ Irish Grey-
hound pig;” and Nathusius states that they occasionally
appear in all the long-eared races, but are not strictly in-
herited, for they occur or fail in animals of the same lit-
ter. 25 As no wild pigs are known to have analogous ap-
pendages, we have at present no reason to suppose that
their
appearance is due to reversion; and if this be so,
we are forced to admit that somewhat complex, though
apparently useless, structures may be suddenly developed
without the aid of selection. This case perhaps throws
some light on the manner of appearance of the hideous
fleshy protuberances, though of an essentially different
nature from the above-described appendages, on the
cheeks of the wart-hog or Phacocherus Africanus.
It is a remarkable fact that the boars of all domesti-
cated breeds have much shorter tusks than wild boars.
Many facts show that with all animals the state of the
hair is much affected by exposure to, or protection from,
climate; and as we see that the state of the hair and
teeth are correlated in Turkish dogs (other analogous
facts will be hereafter given), may we not venture to
surmise that the reduction of the tusks in the domestic
boar is related to his coat of bristles being diminished
from living under shelter ? On the other hand, as we
shall immediately see, the tusks and bristles reappear
with feral boars, which are no longer protected from the
weather. It is not surprising that the tusks should be
more affected than the other teeth ; as parts developed
to serve as secondary sexual characters are always liable
to much variation.
It is a well-known fact that the young of wild Euro-
25 Eudes-Deslongchamps, Mémoires
de la Soc. Linn. de Normandie,' vol. vii.,
1842, p. 41. Richardson, Pigs, their Ori-
gin, &c.,' 1547, p. 30. Nathusius, 'Die
Racen des Schweines,' 1860, s. 54.

CHAP. III.
99
THEIR VARIATION.
pean and Indian pigs 26 for the first six months, are longi-
tudinally banded with light-coloured stripes. This cha-
racter generally disappears under domestication. The
Turkish domestic pigs, however, have striped young, as
have those of Westphalia, “whatever may be their
hue;" 27 whether these latter pigs belong to the same curly-
haired race with the Turkish swine, I do not know. The
pigs which have run wild in Jamaica and the semi-feral
pigs of New Granada, both those which are black and
those which are black with a white band across the sto-
mach, often extending over the back, have resumed this
aboriginal character and produce longitudinally-striped
young. This is likewise the case, at least occasionally,
with the neglected pigs in the Zambesi settlement on the
coast of Africa.
The common belief that all domesticated animals, when
they run wild, revert completely to the character of their
parent-stock, is chiefly founded, as far as I can discover,
on feral pigs. But even in this case the belief is not
28
26 D. Johnson's Sketches of Indian
Field Sports,' p. 272. Mr. Crawfurd
informs me that the same fact holds
good with the wild pigs of the Malay
peninsula.
27 For Turkish pigs, see Desmarest,
Mammalogie,' 1820, p. 391. For those
of Westphalia, see Richardson's Pigs,
their Origin,' &c., 1817, p. 41.
28 With respect to the several fore-
going and following statements on feral
pigs, see Roulin, in "Mém. présentés par
divers Savans à l'Acad.,' &c., Paris, tom.
vi., 1835, p. 326. It should be observed
that his account does not apply to truly
feral pigs; but to pigs long introduced
into the country and living in a half-
wild state. For the truly feral pigs of
Jamaica, see Gosse's 'Sojourn in Ja-
maica,' 1851, p. 386; and Col. Hamilton
Smith, in "Nat. Library,' vol. ix. p. 93.
With respect to Africa, sce Livingstone's
"Expedition to the Zambesi,' 1865, p.
153. The most precise statement with
respect to the tusks of the West Indian
feral boars is by P. Labat (quoted by
Roulin); but this author attributes the
state of these pigs to descent from a do-
mestic stock which he saw in Spain.
Admiral Sulivan, R.N., had ample op-
portunities of observing the wild pigs
on Eagle Islet in the Falklands; and he
informs me that they resembled wild
boars with bristly ridged backs and large
tusks. The pigs which have run wild
in the province of Buenos Ayres (Reng-
ger, Säugethiere,' s. 331) have not re-
verted to the wild type. De Blainville
(Ostéographie,' p. 132) refers to two
skulls of domestic pigs sent from Pata-
gonia by Al. d'Orbigny, and he states
that they have the occipital elevation of
the wild European boar, but that the
head altogether is "plus courte et plus
ramassée." He refers, also, to the skin
of a feral pig from North America, and
says, “il ressemble tout à fait à un petit
sanglier, mais il est presque tout noir, et
peut-être un peu plus ramassé dans ses
formes."

100
CHAP. III.
DOMESTIC PIGS.
grounded on sufficient evidence; for the two main types
of S. scrofa and Indica have never been distinguished in
a feral state. The young, as we have just seen, reac-
quire their longitudinal stripes, and the boars invariably
reassume their tusks. They revert also in the general
shape of their bodies, and in the length of their legs and
muzzles, to the state of the wild animal, as might have
been expected from the amount of exercise which they
are compelled to take in search of food. In Jamaica the
feral pigs do not acquire the full size of the European
wild boar, “never attaining a greater height than 20
inches at the shoulder.” In various countries they reas-
sume their original bristly covering, but in different
degrees, dependent on the climate ; thus, according to
Roulin, the semi-feral pigs in the hot valleys of New
Granada are very scantily clothed; whereas, on the Pa-
ramos, at the height of 7000 to 8000 feet, they acquire a
thick covering of wool lying under the bristles, like that
on the truly wild pigs of France. These pigs on the
Paramos are small and stunted. The wild boar of India
is said to have the bristles at the end of its tail arranged
like the plumes of an arrow, whilst the European boar
has a simple tuft; and it is a curious fact that many, but
not all, of the feral pigs in Jamaica, derived from a Span-
ish stock, have a plumed tail. With respect to colour,
feral pigs generally revert to that of the wild boar; but
in certain parts of S. America, as we have seen, some of
the semi-feral pigs have a curious white band across their
stomachs; and in certain other hot places the pigs are
red, and this colour has likewise occasionally been ob-
served in the feral pigs of Jamaica. From these several
facts we see that with pigs when feral there is a strong
tendency to revert to the wild type; but that this ten-
29
29 Gosse's 'Jamaica,' p. 386, with a
quotation from Williamson's Oriental
Field Sports.' Also Col. Hamilton
Smith, in "Naturalist's Library,' vol.
ix. p. 94.

CHAP. III.
101
CATTLE: THEIR PARENTAGE.
dency is largely governed by the nature of the climate,
amount of exercise, and other causes of change to which
they have been subjected.
The last point worth notice is that we have unusually
good evidence of breeds of pigs now keeping perfectly
true, which have been formed by the crossing of several
distinct breeds. The Improved Essex pigs, for instance,
breed very true; but there is no doubt that they largely
owe their present excellent qualities to crosses originally
made by Lord Western with the Neapolitan race, and to
subsequent crosses with the Berkshire breed (this also
having been improved by Neapolitan crosses), and like-
wise, probably, with the Sussex breed. In breeds thus
formed by complex crosses, the most careful and unre-
mitting selection during many generations has been found
to be indispensable. Chiefly in consequence of so much
crossing, some well-known breeds have undergone rapid
changes; thus, according to Nathusius, *1 the Berkshire
breed of 1780 is quite different from that of 1810; and
since this latter period, at least two distinct forms have
borne the same name.
CATTLE.
DOMESTIC cattle are almost certainly the descendants of
more than one wild form, in the same manner as has been
shown to be the case with our dogs and pigs. Natural-
ists have generally made two main divisions of cattle; the
humped kinds inhabiting tropical countries, called in India
Zebus, to which the specific name of Bos Indicus has been
given; and the common non-humped cattle, generally in-
cluded under the name of Bos taurus. The humped cattle
were domesticated, as may be seen on the Egyptian monu-
ments, at least as early as the twelfth dynasty, that is
2100 B.C. They differ from common cattle in various
30 S. Sidney's edition of 'Youаtt on the Pig,'1860, pp. 7, 26, 27, 29, 30.
31 Schweineschädel,' s. 140.

102
CHAP. III.
CATTLE.
osteological characters, even in a greater degree, accord-
ing to Rütimeyer,2 than do the fossil species of Europe,
namely Bos primigenius, longifrons, and frontosus, from
each other. They differ, also, as Mr. Blyth," who has
particularly attended to this subject, remarks, in general
configuration, in the shape of their ears, in the point
where the dewlap commences, in the typical curvature
of their horns, in their manner of carrying their heads
when at rest, in their ordinary variations of colour, espe-
cially in the frequent presence of “nilgau-like markings
on their feet," and "in the one being born with teeth pro-
truding through the jaws, and the other not so." They
have different habits, and their voice is entirely different.
The humped cattle in India “seldom seek shade, and
never go into the water and there stand knee-deep, like
the cattle of Europe.” They have run wild in parts of
Oude and Rohilound, and can maintain themselves in a
region infested by tigers. They have given rise to many
races differing greatly in size, in the presence of one or
two humps, in length of horns, and other respects. Mr.
Blyth sums up emphatically that the humped and hump-
less cattle must be considered as distinct species. When
we consider the number of points in external structure
and habits, independently of their important osteological
differences, in which they differ from each other; and
that many of these points are not likely to have been
affected by domestication, there can hardly be a doubt,
notwithstanding the adverse opinion of some naturalists,
that the humped and non-humped cattle must be ranked
as specifically distinct.
32 Die Fauna der Pfahlbauten,'1861,
s. 109, 149, 222. See also Geoffroy Saint
Hilaire, in 'Mém. du Mus. d'Hist. Nat,'
tom. x. p. 172; and his son Isidore, in
Hist. Nat. Gén.,' tom. iii. p. 69. Vasey,
in his Delineations of the Ox Tribe,
1851, p. 127, says the zebu has four, and
the common ox five, sacral vertebræ.
Mr. Hodgson found the ribs either thir-
teen or fourteen in number; see a note in
Indian Field,' 1858, p. 62.
33 The Indian Field,' 1858, p. 74,
where Mr. Blyth gives his authorities
with respect to the feral humped cattle.
Pickering, also, in his 'Races of Man,'
1850, p. 274, notices the peculiar cha-
racter of the grunt-like voice of the hump-
ed cattle.

CHAP. III.
103
THEIR PARENTAGE.
The European breeds of humpless cattle are numerous.
Professor Low enumerates 19 British breeds, only a few
of which are identical with those on the Continent. Even
the small Channel islands of Guernsey, Jersey, and
Alderney, possess their own sub-breeds ; 34 and these
again differ from the cattle of the other British islands,
such as Anglesea, and the western isles of Scotland.
Desmarest, who paid attention to the subject, describes
15 French races, excluding sub-varieties and those im-
ported from other countries. In other parts of Europe
there are several distinct races, such as the pale-coloured
Hungarian cattle, with their light and free step, and their
enormous horns sometimes measuring above five feet from
tip to tip:85 the Podolian cattle are remarkable from the
height of their forequarters. In the most recent work on
Cattle,sº engravings are given of fifty-five European
breeds; it is, however, probable that several of these
differ very little from each other, or are merely syno-
nyms. It must not be supposed that numerous breeds
of cattle exist only in long-civilized countries, for we shall
presently see that several kinds are kept by the savages
of Southern Africa.
With respect to the parentage of the several European breeds, we
already know much from Nilsson's Memoir, 7 and more especially
from Rütimeyer's Pfahlbauten' and succeeding works. Two or
three specimens or forms of Bos, closely allied to still living domestic
races, have been found fossil in the more recent tertiary deposits of
Europe. Following Rütimeyer, we have :-
Bos primigenius.—This magnificent, well-known species was do-
mesticated in Switzerland during the Neolithic period; even at this
early period it varied a little, having apparently been crossed with
other races. Some of the larger races on the Continent, as the
34 Mr. H. E. Marquand, in The
Times,' June 23rd, 1856.
35 Vasey, “Delineations of the Ox
Tribe,' p. 124. Brace's 'Hungary,' 1851,
p. 94. The Hungarian cattle descend,
according to Rütimeyer ("Zahmen. Europ.
Rindes., 1866, s. 13), from Bos prime-
genius.
36 Moll and Gayot, ‘La Connaissance
Gén. du Boeuf, Paris, 1860. Fig. 82 is
that of the Podolian breed.
87 A translation appeared in three
parts in the 'Annals and Mag. of Nat.
Hist.,' 2nd series, vol. iv., 1849.

104
CHAP. III.
CATTLE.
39
Friesland, &c., and the Pembroke race in England, closely resemble
in essential structure B. primigenius, and no doubt are its descen-
dants. This is likewise the opinion of Nilsson. Bos primigenius
existed as a wild animal in Cæsar's time, and is now semi-wild,
though much degenerated in size, in the park of Chillingham; for
I am informed by Professor Rütimeyer, to whom Lord Tankerville
sent a skull, that the Chillingham cattle are less altered from the
true primigenius type than any other known breed. 38
Bos trochoceros.- This form is not included in the three species
above mentioned, for it is now considered by Rütimeyer to be the
female of an early domesticated form of B. primigenius, and as the
progenitor of his frontosus race. I may add that specific names
have been given to four other fossil oxen, now believed to be iden-
tical with B. primigenius.
Bos longifrons (or brachyceros) of Owen.—This very distinct spe.
cies was of small size, and had a short body with fine legs. It has
been found in England associated with the remains of the elephant
and rhinoceros.40 It was the commonest form in a domesticated
condition in Switzerland during the earliest part of the Neolithic
period. It was domesticated in England during the Roman period,
and supplied food to the Roman legionaries. 41 Some remains have
been found in Ireland in certain crannoges, of which the dates are
believed to be from 843–933 A.D.42 Professor Owen 43 thinks it pro-
bable that the Welsh and Highland cattle are descended from this
form ; as likewise is the case, according to Rütimeyer, with some of
the existing Swiss breeds. These latter are of different shades of
colour from light-grey to blackish-brown, with a lighter stripe
along the spine, but they have no pure white marks. The cattle of
North Wales and the Highlands, on the other hand, are generally
black or dark-coloured.
Bos frontosus of Nilsson.-This species is allied to B. longifrons,
but in the opinion of some good judges is distinct from it. Both co-
existed in Scania during the same late geological period,14 and both
38 See, also, Rütimeyer's Beitrage pal.
Gesch, der Wiederkauer,' Basel, 1865, s.
54.
39 Pictet's Paléontologie,' tom. i. p.
865 (2nd edit.) With respect to B. tro-
choceros, see Rütimeyer's 'Zahmen Eu-
rop. Rindes,' 1866, s. 26.
40 Owen, British Fossil Mammals,
1846, p. 510.
41 British Pleistocene Mammalia,' by
W. B. Dawkins and W. A. Sandford,
1866, p. xv.
42 W. R. Wilde, An Essay on the
Animal Remains, &c., Royal Irish Aca-
demy,' 1860, p. 29. Also Proc. of R.
Irish Academy,' 1858, p. 48.
43 Lecture: Royal Institution of G.
Britain,' May 2nd, 1856, p. 4. British
Fossil Mammals,' p. 513.
44 Nilsson, in Annals and Mag. of
Nat. Hist.,' 1849, vol. iv. p. 354.

CHAP. III.
105
THEIR PARENTAGE.
have been found in the Irish crannoges.45 Nilsson believes that his
B. frontosus may be the parent of the mountain cattle of Norway,
which have a high protuberance on the skull between the base of the
horns. As Professor Owen believes that the Scotch Highland cattle
are descended from his B. longifrons, it is worth notice that a capa-
ble judge 46 has remarked that he saw no cattle in Norway like the
Highland breed, but that they more nearly resembled the Devon-
shire breed.
Hence we see that three forms or species of Bos, ori-
ginally inhabitants of Europe, have been domesticated;
but there is no improbability in this fact, for the
genus Bos readily yields to domestication. Besides these
three species and the zebu, the yak, the gayal, and the
arni "7 (not to mention the buffalo or genus Bubalus) have
been domesticated; making altogether seven species of
Bos. The zebu and the three European species are now
extinct in a wild state, for the cattle of the B. primige-
nius type in the British parks can hardly be considered
as truly wild. Although certain races of cattle, domes-
ticated at a very ancient period in Europe, are the de-
scendants of the three above-named fossil species, yet it
does not follow that they were here first domesticated.
Those who place much reliance on philology argue that
our cattle were imported from the East.48
But as races
of men invading any country would probably give their
own names to the breeds of cattle which they might
there find domesticated, the argument seems inconclu-
sive. There is indirect evidence that our cattle are the
descendants of species which originally inhabited a tem-
perate or cold climate, but not a land long covered with
snow; for our cattle, as we have seen in the chapter on
Horses, apparently have not the instinct of scraping
away the snow to get at the herbage beneath. No one
45 See W. R. Wilde, ut supra; and Mr.
Blyth, in Proc. Irish Academy,' March
5th, 1864.
46 Laing's Tour in Norway,' p. 110.
47 Isid. Geoffroy St. Hilaire, Hist.
Nat. Gén.,' tom. iii. p. 96.
48 Idem, tom. iii. pp. 82, 91.

106
CHAP. III
CATTLE.
could behold the magnificent wild bulls on the bleak
Falkland Islands in the southern hemisphere, and doubt
about the climate being admirably suited to them.
Azara has remarked that in the temperate regions of La
Plata the cows conceive when two years old, whilst in
the much hotter country of Paraguay they do not con-
ceive till three years old; “from which fact," as he adds,
“one may conclude that cattle do not succeed so well in
warm countries."' 49
The above-named three fossil forms of Bos have been
ranked by nearly all palæontologists as distinct species;
and it would not be reasonable to change their denomina-
tion simply because they are now found to be the parents
of several domesticated races. But what is of most im-
portance for us, as showing that they deserve to be rank-
ed as species, is that they co-existed in different parts of
Europe during the same period, and yet kept distinct.
Their domesticated descendants, on the other hand, if not
separated, cross with the utmost freedom and become
commingled. The several European breeds have so often
been crossed, both intentionally and unintentionally, that,
if any sterility ensued from such unions, it would certain-
ly have been detected. As zebus inhabit a distant and
much hotter region, and as they differ in so many charac-
ters from our European cattle, I have taken pains to
ascertain whether the two forms are fertile when crossed.
The late Lord Powis imported some zebus and crossed
them with common cattle in Shropshire; and I was as-
sured by his steward that the cross-bred animals were
perfectly fertile with both parent-stocks. Mr. Blyth in-
forms me that in India hybrids, with various proportions
of either blood, are quite fertile; and this can hardly fail
to be known, for in some districts so the two species are
allowed to breed freely together. Most of the cattle
49 Quadrupèdes du Paraguay,' tom. ii. p. 360.
60 Walther, 'Das Rindvieb,' 1817, s. 80.

CHAP. IIL
107
CROSSED SPECIES FERTILE.
which were first introduced into Tasmania were humped,
so that at one time thousands of crossed animals existed
there; and Mr. B. O'Neile Wilson, M.A., writes to me
from Tasmania that he has never heard of any sterility
having been observed. He himself formerly possessed a
herd of such crossed cattle, and all were perfectly fertile;
so much so, that he cannot remember even a single cow
failing to calve. These several facts afford an important
confirmation of the Pallasian doctrine that the descen-
dants of species which when first domesticated would if
crossed probably have been in some degree sterile, become
perfectly fertile after a long course of domestication. In
a future chapter we shall see that this doctrine throws
much light on the difficult subject of Hybridism.
I have alluded to the cattle in Chillingham Park,
which, according to Rütimeyer, have been very little
changed from the Bos primigenius type. This park is so
ancient that it is referred to in a record of the year 1220.
The cattle in their instincts and habits are truly wild. They
are white, with the inside of the ears reddish brown, eyes
rimmed with black, muzzles brown, hoofs black, and horns
white tipped with black. Within a period of thirty-three
years about a dozen calves were born with “brown and
blue spots upon the cheeks or necks; but these, together
with
any defective animals, were always destroyed. “Ac-
cording to Bewick, about the year 1770 some calves ap-
peared with black ears; but these were also destroyed
by the keeper, and black ears have not since reappeared.
The wild white cattle in the Duke of Hamilton's park,
where I have heard of the birth of a black calf, are said
by Lord Tankerville to be inferior to those at Chilling-
ham. The cattle kept until the year 1780 by the Duke of
Queensberry, but now extinct, had their ears, muzzle, and
orbits of the eyes black. Those which have existed from
time immemorial at Chartley closely resemble the cattle
at Chillingham, but are larger, “ with some small differ-
ence in the colour of the ears.” “They frequently tend to

108
CHAP. III.
CATTLE.
become entirely black; and a singular superstition pre-
vails in the vicinity that, when a black calf is born, some
calamity impends over the noble house of Ferrers. All
the black calves are destroyed."
The cattle at Burton
Constable in Yorkshire, now extinct, had ears, muzzle,
and the tip of the tail black. Those at Gisburne, also in
Yorkshire, are said by Bewick to have been sometimes
without dark muzzles, with the inside alone of the ears
brown; and they are elsewhere said to have been low in
stature and hornless. 51
The several above-specified differences in the park-cat-
tle, slight though they be, are worth recording, as they
show that animals living nearly in a state of nature, and
exposed to nearly uniform conditions, if not allowed to
roam freely and to cross with other herds, do not keep as
uniform as truly wild animals. For the preservation of
a uniform character, even within the same park, a certain
degree of selection—that is, the destruction of the dark-
coloured calves—is apparently necessary.
The cattle in all the parks are white; but, from the oc-
casional appearance of dark-coloured calves, it is ex-
tremely doubtful whether the aboriginal Bos primigenius
was wbite.
The following facts, however, show that
there is a strong, though not invariable, tendency in wild
or escaped cattle, under widely different conditions of
life, to become white with coloured ears. If the old
writers Boethius and Leslie 52 can be trusted, the wild
cattle of Scotland were white and furnished with a great
mane; but the colour of their ears is not mentioned.
51 I am much indebted to the present
Earl of Tankerville for information about
his wild cattle; and for the skull which
was sent to Prof. Rütimeyer. The fullest
account of the Chillingham cattle is given
by Mr. Hindmarsh, together with a let-
ter by the late Lord Tankerville, in 'An-
nals and Mag. of Nat. Hist.,' vol. ii., 1839,
p. 274. See Bewick, 'Quadrupeds, 2nd
edit., 1791, p. 35, note. With respect to
those of the Duke of Queensberry, see
Pennant's Tour in Scotland,'p. 109. For
those of Chartley, see Low's Domesti-
cated Animals of Britain,' 1845, p. 238.
For those of Gisburne, see Bewick's
Quadrupeds, and Encyclop. of Rural
Sports,' p. 101.
52 Boethius was born in 1470; 'An-
nals and Mag. of Nat. Hist.,' vol. ii.,
1839, p. 281; and vol, iv., 1849, p. 424.

CHAP. III
109
PARK-CATTLE.
The primæval forest formerly extended across the whole
country from Chillingham to Hamilton, and Sir Walter
Scott used to maintain that the cattle still preserved in
these two parks, at the two extremities of the forest,
were remnants of its original inhabitants; and this view
certainly seems probable. In Wales,6during the tenth
century, some of the cattle are described as being white
with red ears.
Four hundred cattle thus coloured were
sent to King John; and an early record speaks of a hun-
dred cattle with red ears having been demanded as a
compensation for some offence, but, if the cattle were
of a dark or black colour, one hundred and fifty were to
be presented. The black cattle of North Wales appa-
rently belong, as we have seen, to the small longifrons
type: and as the alternative was offered of either 150
dark cattle, or 100 white cattle with red ears, we may
presume that the latter were the larger beasts, and pro-
bably belonged to the primigenius type. Youаtt has
remarked that at the present day, whenever cattle of the
short-horn breed are white, the extremities of their ears
are more or less tinged with red.
The cattle which have run wild on the Pampas, in
Texas, and in two parts of Africa, have become of nearly
uniform dark brownish-red.54 On the Ladrone Islands,
in the Pacific Ocean, immense herds of cattle, which
were wild in the year 1741, are described as “milk-white,
except their ears, which are generally black.” 65 The
Falkland Islands, situated far south, with all the con-
ditions of life as different as it is possible to conceive
from those of the Ladrones, offer a more interesting case.
Cattle have run wild there during eighty or ninety years;
53 Youatton Cattle, 1834, p. 48:
See also p. 242, on short-horn cattle.
Bell, in his British Quadrupeds,' p.
423, states that, after long attending to
the subject, he has found that white
cattle invariably have coloured ears.
54 Azara, 'Des Quadrupèdes du Para-
guay,' tom. ii. p. 361. Azara quotes Buf-
fon for the feral cattle of Africa. For
Texas, see 'Times,' Feb. 18th, 1846.
65 Anson's Voyage. See Kerr and
Porter's Collection,' vol. xii. p. 103.

110
CHAP. III.
CATTLE.
and in the southern districts the animals are mostly
white, with their feet, or whole heads, or only their ears
black; but my informant, Admiral Sulivan, who long
resided on these islands, does not believe that they are
ever purely white. So that in these two archipelagos
we see that the cattle tend to become white with coloured
ears. In other parts of the Falkland Islands other co-
lours prevail: near Port Pleasant brown is the common
tint'; round Mount Usborne, about half the animals in
some of the herds were lead or mouse-coloured, which
elsewhere is an unusual tint. These latter cattle, though
generally inhabiting high land, breed about a month
earlier than the other cattle; and this circumstance would
aid in keeping them distinct and in perpetuating this pe-
culiar colour. It is worth recalling to mind that blue or
lead-coloured marks have occasionally appeared on the
white cattle of Chillingham. So plainly different were
the colours of the wild herds in different parts of the
Falkland Islands, that in hunting them, as Admiral Suli-
van informs me, white spots in one district, and dark
spots in another district, were always looked out for on
the distant hills. In the intermediate districts interme-
diate colours prevailed. Whatever the cause may be,
this tendency in the wild cattle of the Falkland Islands,
which are all descended from a few brought from La
Plata, to break up into herds of three different colours,
is an interesting fact.
Returning to the several British breeds, the conspicuous
difference in general appearance between Short-horns,
Long-horns (now rarely seen), Herefords, Highland
cattle, Alderneys, &c., must be familiar to every one.
A large part of the difference, no doubt, may be due to
descent from primordially distinct species; but we may
feel sure that there has been in addition a considerable
amount of variation. Even during the Neolithic period,
66 See also Mr. Mackinnon's pamphlet on the Falkland Islands, p. 24.

CHAP. III.
111
THEIR VARIATION.
the domestic cattle were not actually identical with the
aboriginal species. Within recent times most of the
breeds have been modified by careful and methodical
selection. How strongly the characters thus acquired
are inherited, may be inferred from the prices realised
by the improved breeds; even at the first sale of Col-
ling's Short-horns, eleven bulls reached an average of
2141., and lately Short-horn bulls have been sold for a
thousand guineas, and have been exported to all quarters
of the world.
Some constitutional differences may be here noticed.
The Short-horns arrive at maturity far earlier than the
wilder breeds, such as those of Wales or the Highlands.
This fact has been shown in an interesting manner by Mr.
Simonds,67 who has given a table of the average period
of their dentition, which proves that there is a difference
of no less than six months in the appearance of the per-
manent incisors. The period of gestation, from observa-
tions made by Tessier on 1131 cows, varies to the extent
of eighty-one days; and what is more interesting, M.
Lefour affirms “ that the period of gestation is longer in
the large German cattle than in the smaller breeds.” 58
With respect to the period of conception, it seems certain
that Alderney and Zetland cows often become pregnant
earlier than other breeds. Lastly, as four fully-deve-
loped mammæ is a generic character in the genus Bos,
it is worth notice that with our domestic cows the two
rudimentary mammæ often become fairly well developed
and yield milk.
As numerous breeds are generally found only in long-
civilized countries, it may be well to show that in some
countries inhabited by barbarous races, who are frequently
60
57 The Age of the Ox, Sheep, Pig,'
&c., by Prof. James Simonds, published
by order of the Royal Agricult. Soc.
68 Ann. Agricult. France,' April,
1887, as quoted in The Veterinary,'
vol. xii. p. 725. I quote Tessier's obser-
vations from Youаtt on Cattle, p. 527.
59 The Veterinary,' vol. viii. p. 681,
and vol. x. p. 268. Low's Domest.
Animals of Great Britain,' p. 297.
60 Mr. Ogilby, in Proc. Zoolog. Soc.,'
1836, p. 138, and 1810, p. 4.

112
CHAP. III.
CATTLE.
at war with each other and therefore have little free com-
munication, several distinct breeds of cattle now exist or
formerly existed. At the Cape of Good Hope Leguat
observed, in the year 1720, three kinds. At the present
day various travellers have noticed the differences in the
breeds in Southern Africa. Sir Andrew Smith several
years ago remarked to me that the cattle possessed by
the different tribes of Caffres, though living near each
other under the same latitude and in the same kind of
country, yet differed, and he expressed much surprise at
the fact. Mr. Andersson has described 62 the Damara,
Bechuana, and Namaqua cattle; and he informs me in a
letter that the cattle north of Lake Ngami are likewise
different, as Mr. Galton has heard is the case with the
cattle of Benguela. The Namaqua cattle in size and
shape nearly resemble European cattle, and have short
stout horns and large hoofs. The Damara cattle are very
peculiar, being big-boned, with slender legs and small
hard feet; their tails are adorned with a tuft of long
bushy hair nearly touching the ground, and their horns
are extraordinarily large. The Bechuana cattle have even
larger horns, and there is now a skull in London with the
two horns 8 ft. 81 in. long, as measured in a straight line
from tip to tip, and no less than 13 ft. 5 in., as measured
along their curvature! Mr. Andersson in his letter to me
says that, though he will not venture to describe the
differences between the breeds belonging to the many
different sub-tribes, yet such certainly exist, as shown by
the wonderful facility with which the natives discrimi-
nate them.
That many breeds of cattle have originated through
variation, independently of descent from distinct species,
we may infer from what we see in South America, where
the genus Bos was not endemic, and where the cattle
61 Leguat's Voyage, quoted by Vasey
in his Delineations of the Ox-tribe,' p.
182.
62 Travels in South Africa,' pp. 317,
336.

CHAP. III.
113
THEIR VARIATION.
63
which now exist in such vast numbers are the descend-
ants of a few imported from Spain and Portugal. In
Columbia, Roulin describes two peculiar breeds,
namely, pelones, with extremely thin and fine hair, and
calongos, absolutely naked. According to Castelnau
there are two races in Brazil, one like European cattle,
the other different, with remarkable horns. In Paraguay,
Azara describes a breed which certainly originated in S.
America called chivos, “because they have straight ver-
tical horns, conical, and very large at the base.” He
likewise describes a dwarf race in Corrientes, with short
legs and a body larger than usual. Cattle without horns,
and others with reversed hair, have also originated in
Paraguay.
Another monstrous breed, called niatas or natas, of
which I saw two small herds on the northern bank of
the Plata, is so remarkable as to deserve a fuller descrip-
tion. This breed bears the same relation to other breeds,
as bull or pug dogs do to other dogs, or as improved
pigs, according to H. von Nathusius, do to common
pigs. Rütimeyer believes that these cattle belong to
the primigenius type. The forehead is very short and
broad, with the nasal end of the skull, together with the
whole plane of the upper molar-teeth, curved upwards.
The lower jaw projects beyond the upper, and has a
corresponding upward curvature. It is an interesting
fact that an almost similar conformation characterizes, as
I have been informed by Dr. Falconer, the extinct and
63 Mém. de l'Institut présent. par not form a distinct race. Prof. Wyman,
divers Savans,' tom. vi., 1835, p. 333. of Cambridge, United States, informs me
For Brazil, see Comptes Rendus,' June that the common cod-fish presents a
15th, 1846. See Azara, Quadrupèdes similar monstrosity, called by the fisher-
du Paraguay,' tom. ii. pp. 359, 361.
men the “bulldog cod." Prof. Wyman
64 Schweineschädel,' 1864, s. 104.. also concluded, after making numerous
Nathusius states that the form of skull inquiries in La Plata, that the niata
characteristic of the niata cattle occa- cattle transmit their peculiarities or form
sionally appears in European cattle ;
but he is mistaken, as we shall hereafter 65 Ueber Art des Zahmen Europ. Rin-
see, in supposing that these cattle do des, 1866. s. 28.
a race.

114
CHAP. III.
CATTLE.
gigantic Sivatherium of India, and is not known in any
other ruminant. The upper lip is much drawn back, the
nostrils are seated high up and are widely open, the
eyes
project outwards, and the horns are large. In walking
the head is carried low, and the neck is short. The hind
legs appear to be longer, compared with the front legs,
than is usual. The exposed incisor teeth, the short head
and upturned nostrils, give these cattle the most ludi-
crous, self-confident air of defiance. The skull which I
presented to the College of Surgeons has been thus de-
scribed by Professor Owen : 66“ It is remarkable from the
stunted development of the nasals, premaxillaries, and
fore-part of the lower jaw, which is unusually curved up-
wards to come into contact with the premaxillaries. The
nasal bones are about one-third the ordinary length, but
retain almost their normal breadth. The triangular vacui-
ty is left between them, the frontal and lachrymal, which
latter bone articulates with the premaxillary, and thus
excludes the maxillary from any junction with the nasal.”
So that even the connexion of some of the bones is chang-
ed. Other differences might be added: thus the plane of
the condyles is somewhat modified, and the terminal edge
of the premaxillaries forms an arch. In fact, on compari-
son with the skull of a common ox, scarcely a single bone
presents the same exact shape, and the whole skull has a
wonderfully different appearance.
The first brief published notice of this race was by
Azara, between the years 1783-96; but Don F. Muniz, of
Luxan, who has kindly collected information for me,
states that about 1760 these cattle were kept as curiosi-
ties near Buenos Ayres. Their origin is not positively
known, but they must have originated subsequently to
the year 1552, when cattle were first introduced. Signor
Muniz informs me that the breed is believed to have ori-
86 Descriptive Cat. of Ost. Collect. of
College of Surgeons,' 1553 p. 624. Vasey,
in his "Delineations of the Ox-tribe!
has given a figure of this skull; and I
sent a photograph of it to Prof. Rüti-
meyer.

CHAP. III.
115
CAUSES OF VARIATION.
ginated with the Indians southward of the Plata. Even
to this day those reared near the Plata show their less
civilized nature in being fiercer than common cattle, and
in the cow, if visited too often, easily deserting her first
calf. The breed is very true, and a niata bull and cow
invariably produce niata calves. The breed has already
lasted at least a century. A niata bull crossed with a
common cow, and the reverse cross, yield offspring hav-
ing an intermediate character, but with the niata cha-
racter strongly displayed. According to Signor Muniz,
there is the clearest evidence, contrary to the common
belief of agriculturists in analogous cases, that the niata
cow when crossed with a common bull transmits her pe-
culiarities more strongly than does the niata bull when
crossed with a common cow. When the pasture is toler-
ably long, these cattle feed as well as common cattle with
their tongue and palate; but during the great droughts,
when so many animals perish on the Pampas, the niata
breed lies under a great disadvantage, and would, if not
attended to become extinct; for the common cattle, like
horses, are able just to keep alive by browsing on the
twigs of trees and on reeds with their lips: this the nia-
tas cannot so well do, as their lips do not join, and hence
they are found to perish before the common cattle. This
strikes me as a good illustration of how little we are able
to judge from the ordinary habits of an animal, on what
circumstances, occurring only at long intervals of time,
its rarity or extinction may depend. It shows us, also,
how natural selection would have determined the rejec-
tion of the niata modification had it arisen in a state of
nature.
Having described the semi-monstrous niata breed, I
may allude to a white bull, said to have been brought
from Africa, which was exhibited in London in 1829, and
which has been well figured by Mr. Harvey. It had a
67 Loudon's Magazine of Nat. Hist.,'
vol. i. 1929, p. 113. Separate figures are
given of the animal, its hoofs, eye, and
dewlap.

116
CEAP. III.
CATTLE.
hump, and was furnished with a mane. The dewlap was
peculiar, being divided between its forelegs into parallel
divisions. Its lateral hoofs were annually shed, and
grew to the length of five or six inches. The eye was
very peculiar, being remarkably prominent, and “resem-
bled a cup and ball, thus enabling the animal to see on
all sides with equal ease; the pupil was small and oval,
or rather a parallelogram with the ends cut off, and ly-
ing transversely across the ball.” A new and strange
breed might probably have been formed by careful breed-
ing and selection from this animal.
I have often speculated on the probable causes through
which each separate district in Great Britain came to pos-
sess in former times its own peculiar breed of cattle; and
the question is, perhaps, even more perplexing in the case
of Southern Africa. We now know that the differences
may be in part attributed to descent from distinct spe-
cies; but this will not suffice. Have the slight differ-
ences in climate and in the nature of the pasture, in the
different districts of Britain, directly induced correspond-
ing differences in the cattle? We have seen that the
semi-wild cattle in the several British parks are not iden-
tical in colouring or size, and that some degree of selection
has been requisite to keep them true. It is almost cer-
tain that abundant food given during many generations
directly affects the size of a breed. That climate di-
rectly affects the thickness of the skin and the hair is
likewise certain; thus Roulin asserts 69 that the hides of
the feral cattle on the hot Llanos "are always much less
heavy than those of the cattle raised on the high plat-
form of Bogota; and that these hides yield in weight
and in thickness of hair to those of the cattle which have
run wild on the lofty Paramos." The same difference has
been observed in the hides of the cattle reared on the
68
88 Low, 'Domesticated Animals of the
British Isles,' p. 264.
69 Mém. de l'Institut présent. par di-
vers Savans,' tom. vi., 1835, p. 382.

CHAP. III.
117
CAUSES OF VARIATION. .
bleak Falkland Islands and on the temperate Pampas.
Low has remarked 10 that the cattle which inhabit the
more humid parts of Britain have longer hair and thicker
skins than other British cattle; and the hair and horns
are so closely related to each other, that, as we shall see
in a future chapter, they are apt to vary together; thus
climate might indirectly affect, through the skin, the
form and size of the horns. When we compare highly
improved stall-fed cattle with the wilder breeds, or com-
pare mountain and lowland breeds, we cannot doubt that
an active life, leading to the free use of the limbs and
lungs, affects the shape and proportions of the whole
body. It is probable that some breeds, such as the semi-
monstrous niata cattle, and some peculiarities, such as
being hornless, &c., have appeared suddenly from what
we may call a spontaneous variation ; but even in this
case a rude kind of selection is necessary, and the ani-
mals thus characterized must be at least partially sepa-
rated from others. This degree of care, however, has
sometimes been taken even in little-civilized districts,
where we should least have expected it, as in the case of
the niata, chivo, and hornless cattle in S. America.
That methodical selection has done wonders within a
recent period in modifying our cattle, no one doubts.
During the process of methodical selection it has occa-
sionally happened that deviations of structure, more
strongly pronounced than mere individual differences,
yet by no means deserving to be called monstrosities,
have been taken advantage of: thus the famous Long-
horn Bull, Shakespeare, though of the pure Canley stock,
"scarcely inherited a single point of the long-horned
breed, his horns excepted ; 7 yet in the hands of Mr. Fow-
ler, this bull greatly improved his race. We have also
reason to believe that selection, carried on so far uncon-
70 Idem, pp. 304, 368, &c.
71 Youаtt on Cattle, p. 193. A full ac-
count of this bull is taken from Mar-
shall.

118
CHAP. III.
CATTLE.
sciously that there was at no one time any distinct intention
to improve or change the breed, has in the course of time
modified most of our cattle; for by this process, aided by
more abundant food, all the lowland British breeds have
increased greatly in size and in early maturity since the
reign of Henry VII." It should never be forgotten that
many animals have to be annually slaughtered; so that
each owner must determine which shall be killed and
which preserved for breeding. In every district, as You-
att has remarked, there is a prejudice in favor of the na-
tive breed; so that animals possessing qualities, whatever
they may be, which are most valued in each district, will
be oftenest preserved ; and this unmethodical selection
assuredly will in the long run affect the character of the
whole breed. But it may be asked, can this rude kind of
selection have been practised by barbarians such as those
of southern Africa ? In a future chapter on Selection we
shall see that this has certainly occurred to some extent.
Therefore, looking to the origin of the many breeds of
cattle which formerly inhabited the several districts of
Britain, I conclude that, although slight differences in the
nature of the climate, food, &c., as well as changed habits
of life, aided by correlation of growth, and the occasional
appearance from unknown causes of considerable devia-
tions of structure, have all probably played their parts; yet
that the occasional preservation in each district of those
individual animals which were most valued by each own-
er has perhaps been even more effective in the production
of the several British breeds. As soon as two or more
breeds had once been formed in any district, or when
new breeds descended from distinct species were intro-
duced, their crossing, especially if aided by some selec-
tion, will have multiplied the number and modified the
characters of the older breeds.
72 Youаtt on Cattle, p. 116. Lord Spencer has written on this same subject.

CHAP. III.
119
SHEEP. THEIR VARIATION.
SHEEP.
I SHALL treat this subject briefly. Most authors look at
our domestic sheep as descended from several distinct
species; but how many still exist is doubtful. Mr. Blyth
believes that there are in the whole world fourteen spe-
cies, one of which, the Corsican moufflon, he concludes
(as I am informed by him) to be the parent of the small-
er, short-tailed breeds, with crescent-shaped horns, such
as the old Highland sheep. The larger, long-tailed breeds,
having horns with a double flexure, such as the Dorsets,
merinos, &c., he believes to be descended from an un-
known and extinct species. M. Gervais makes six species
of Ovis ;* but concludes that our domestic sheep form a
distinct genus, now completely extinct. A German na-
turalist74 believes that our sheep descend from ten abo-
riginally distinct species, of which only one is still living
in a wild state! Another ingenious observer, though
not a naturalist, with a bold defiance of everything
known on geographical distribution, infers that the sheep
of Great Britain alone are the descendants of eleven en-
demic British forms! Under such a hopeless state of
doubt it would be useless for my purpose to give a de-
tailed account of the several breeds; but a few remarks
may be added.
Sheep have been domesticated from a very ancient pe-
riod. Rütimeyer 78 found in the Swiss lake-dwellings the
remains of a small breed, with thin and tall legs, and with
horns like those of a goat: this race differs somewhat
from any one now known. Almost every country has its
73 Blyth on the genus Ovis, in An- 74 Dr. L. Fitzinger, 'Ueber die Racen
nals and Mag. of Nat. History,' vol. vii., des Zahmen Schafes,' 1860, s. 86.
1841, p. 261 : with respect to the parent- 75 J. Anderson, Recreations in Agri-
age of the breeds, see Mr. Blyth's excel- culture and Natural History,' vol. ii. p.
lent articles in 'Land and Water,' 1867, 164.
pp. 134, 156. Gervais, Hist. Nat. des 78 Pfahlbauten,' s. 127, 193.
Mammifères,' 1855, tom. ii, p. 191.

120
CHAP. III.
SHEEP.
78
own peculiar breed; and many countries have many
breeds differing greatly from each other. One of the
most strongly marked races is an Eastern one with a long
tail, including, according to Pallas, twenty vertebræ, and
so loaded with fat, that, from being esteemed a delicacy,
it is sometimes placed on a truck which is dragged about
by the living animal. These sheep, though ranked by
Fitzinger as a distinct aboriginal form, seem to bear in
their drooping ears the stamp of long domestication.
This is likewise the case with those sheep which have two
great masses of fat on the rump, with the tail in a rudi-
mentary condition. The Angola variety of the long-tailed
race has curious masses of fat on the back of the head and
beneath the jaws." Mr. Hodgson in an admirable paper
on the sheep of the Himalaya infers from the distribution
of the several races, “ that this caudal augmentation in
most of its phases is an instance of degeneracy in these
pre-eminently Alpine animals.” The horns present an
endless diversity in character; being, especially in the
female sex, not rarely absent, or, on the other hand,
amounting to four or even eight in number. The horns,
when numerous, arise from a crest on the frontal bone,
which is elevated in a peculiar manner. It is remarkable
that multiplicity of horns “is generally accompanied by
great length and coarseness of the fleece.” 79
This corre-
lation, however, is not invariable; for I am informed by
Mr. D. Forbes, that the Spanish sheep in Chile resemble,
in fleece and in all other characters, their parent merino-
race, except that instead of a pair they generally bear four
horns. The existence of a pair of mamma is a generic
character in the genus Ovis as well as in several allied
forms; nevertheless, as Mr. Hodgson has remarked, “this
character is not absolutely constant even among the true
and proper sheep: for I have more than once met with
77 Youаtt on Sheep, p. 120.
78 Journal of the Asiatic Soc. of Ben.
gal,' vol. xvi. pp. 1007, 1016.
79 Youаtt on Sheep, pp. 142-169.

CHAP. III.
121
THEIR VARIATION.
Cágias (a sub-Himalayan domestic race) possessed of four
teats.” 80 This case is the more remarkable as, when any
part or organ is present in reduced number in comparison
with the same part in allied groups, it usually is subject
to little variation. The presence of interdigital pits has
likewise been considered as a generic distinction in sheep;
but Isidore Geoffroy 81 has shown that these pits or pouch-
es are absent in some breeds.
In sheep there is a strong tendency for characters,
which have apparently been acquired under domestica-
tion, to become attached either exclusively to the male
sex, or to be more highly developed in this than in the
other sex.
Thus in many breeds the horns are deficient
in the ewe, though this likewise occurs occasionally with
the female of the wild musmon. In the rams of the
Wallachian breed “the horns spring almost perpendicu-
larly from the frontal bone, and then take a beautiful
spiral form; in the ewes they protrude nearly at right
angles from the head, and then become twisted in a
singular manner. Mr. Hodgson states that the extra-
ordinarily arched nose or chaffron, which is so highly
developed in several foreign breeds, is characteristic of
the ram alone, and apparently is the result of domestica-
I hear from Mr. Blyth that the accumulation of
fat in the fat-tailed sheep of the plains of India is greater
in the male than in the female; and Fitzinger 84 remarks
that the mane in the African maned race is far more
developed in the ram than in the ewe.
Different races of sheep, like cattle, present constitu-
tional differences. Thus the improved breeds arrive at
maturity at an early age, as has been well shown by Mr.
Simonds through their early average period of dentition.
The several races have become adapted to different kinds
tion.83
80 Journal Asiat. Soc. of Bengal,' vol.
xvi., 1847, p. 1015.
81 Hist. Nat. Gén.,' tom. iii. p. 435.
82 Youаtt on Sheep, p. 138.
83 Journal Asiat. Soc. of Bengal.' vol.
xvi., 1847, pp. 1015, 1016.
84 Racen des Zahmen Schafes,' s. 77.
3
6

122
CHAP. III.
SHEEP.
of pasture and climate: for instance, no one can rear
Leicester sheep on mountainous regions, where Cheviots
flourish. As Youаtt has remarked, “in all the different
districts of Great Britain we find various breeds of sheep
beautifully adapted to the locality which they occupy.
No one knows their origin; they are indigenous to the
soil, climate, pasturage, and the locality on which they
graze; they seem to have been formed for it and by it." 85
Marshall relates 86 that a flock of heavy Lincolnshire and
light Norfolk sheep which had been bred together in a
large sheep-walk, part of which was low, rich, and moist,
and another part high and dry, with benty grass, when
turned out, regularly separated from each other; the
heavy sheep drawing off to the rich soil, and the lighter
sheep to their own soil; so that “whilst there was plenty
of grass the two breeds kept themselves as distinct as
rooks and pigeons.” Numerous sheep from various parts
of the world have been brought during a long course of
years to the Zoological Gardens of London; but as
Youatt, who attended the animals as a veterinary surgeon,
remarks, “few or none die of the rot, but they are
phthisical; not one of them from a torrid climate lasts
out the second year, and when they die their lungs are
tuberculated.” B7 Even in certain parts of England it has
been found impossible to keep certain breeds of sheep;
thus on a farm on the banks of the Ouse, the Leicester
sheep were so rapidly destroyed by pleuritis 88. that the
owner could not keep them ; the coarser-skinned sheep
never being affected.
The period of gestation was formerly thought to be so
analterable à character, that a supposed difference be-
tween the wolf and the dog in this respect was esteemed
85 Rural Economy of Norfolk,' vol.
ii. p. 136.
86 Youаtt on Sheep, p. 312. On same
subject, see excellent remarks in 'Gar-
dner's Chronicle,' 1858, p. 368. For ex-
periments in crossing Cheviot sheep with
Leicesters, see Youatt, p. 325.
87 Youаtt on Sheep, note, p. 491.
88 The Veterinary,' vol. X., p. 217.

CHAP. III.
123
THEIR VARIATION.
a sure sign of specific distinction; but we have seen that
the period is shorter in the improved breeds of the pig,
and in the larger breeds of the ox, than in other breeds
of these two animals. And now we know, on the excel-
Jent authority of Hermann von Nathusius,eº that Merino
and Southdown sheep, when both have long been kept
under exactly the same conditions, differ in their average
period of gestation, as is seen in the following Table :
2
Merinos
Southdowns
Half-bred Merinos and Southdowns
& blood of Southdown
} 용
​150-3 days.
144.2
146.3
145.5
144.2
>
In this graduated difference, in these cross-bred animals
having different proportions of Southdown blood, we see
how strictly the two periods of gestation have been trans-
mitted. Nathusius remarks that, as Southdowns grow
with remarkable rapidity after birth, it is not surprising
that their fætal development should have been shortened.
It is of course possible that the difference in these two
breeds may be due to their descent from distinct parent-
species; but as the early maturity of the Southdowns has
long been carefully attended to by breeders, the differ-
ence is more probably the result of such attention. Lastly,
the fecundity of the several breeds differs much; some
generally producing twins or even triplets at a birth, of
which fact the curious Shangai sheep (with their trunca-
ted and rudimentary ears, and great Roman noses), lately
exhibited in the Zoological Gardens, offer a remarkable
instance.
Sheep are perhaps more readily affected by the direct
action of the conditions of life to which they have been
exposed than almost any other domestic animal. AC-
cording to Pallas, and more recently according to Erman,
89 A translation of his paper is given in 'Bull. Soc. Imp. d'Acclimat.,' tom. ix., 1862,
p. 723.

124
CHAP. III.
SHEEP.
the fat-tailed Kirghisian sheep, when bred for a few gen-
erations in Russia, degenerate, and the mass of fat dwin-
dles away,“ the scanty and bitter herbage of the steppes
seems so essential to their development.” Pallas makes
an analogous statement with respect to one of the Crimean
breeds. Burnes states that the Karakool breed, which
produces a fine, curled, black, and valuable fleece, when
removed from its own canton near Bokhara to Persia, or to
other quarters, loses its peculiar fleece. In all such cases,
however, it may be that a change of any kind in the con-
ditions of life causes variability and consequent loss of
character, and not that certain conditions are necessary
for the development of certain characters.
Great heat, however, seems to act directly on the fleece:
several accounts have been published of the change which
sheep imported from Europe undergo in the West Indies.
Dr. Nicholson of Antigua informs me that, after the third
generation, the wool disappears from the whole body,
except over the loins; and the animal then appears like a
goat with a dirty door-mat on its back. A similar change
is said to take place on the west coast of Africa.°1 On
the other hand, many wool-bearing sheep live on the hot
plains of India. Roulin asserts that in the lower and
heated valleys of the Cordillera, if the lambs are sheared
as soon as the wool has grown to a certain thickness, all
goes on afterwards as usual; but if not sheared, the wool
detaches itself in flakes, and short shining hair like that
on a goat is produced ever afterwards. This curious re-
sult seems merely to be an exaggerated tendency natural
90 Erman's Travels in Siberia' (Eng.
trans.), vol. i. p. 228. For Pallas on the
fat-tailed sheep, I quote from Anderson's
account of the Sheep of Russia,' 1794,
p. 34. With respect to the Crimean sheep,
see Pallas' Travels' (Eng. trans.), vol.
ii. p. 454. For the Karakool sheep, see
Burnes' Travels in Bokhara,' vol. iii. p.
151.
91 See Report of the Directors of the
Sierra Leone Company, as quoted in
White's Gradation of Man,' p. 95. With
respect to the change which sheep un-
dergo in the West Indies, see also Dr.
Davy, in 'Edin. New. Phil. Journal,'Jan.
1852. For the statement made by Roulin,
see Mém. de l'Institut présent. par di-
vers Savans,' tom. vi., 1835, p. 347.

CHAP III.
125
CAUSES OF VARIATION.
to the Merino breed, for as a great authority, namely,
Lord Somerville, remarks," the wool of our Merino sheep
after shear-time is hard and coarse to such a degree as to
render it almost impossible to suppose that the same ani-
mal could bear wool so opposite in quality, compared to
that which has been clipped from it: as the cold weather
advances, the fleeces recover their soft quality.” As in
sheep of all breeds the fleece naturally consists of longer
and coarser hair covering shorter and softer wool, the
change which it often undergoes in hot climates is proba-
bly merely a case of unequal development; for even with
those sheep which like goats are covered with hair, a
small quantity of underlying wool may always be found.º2
In the wild mountain-sheep (Ovis montana) of North
America there is an annual analogous change of coat;
“the wool begins to drop out in early spring, leaving in
its place a coat of hair resembling that of the elk, a .
change of pelage quite different in character from the or-
dinary thickening of the coat or hair, common to all furred
animals in winter,-for instance, in the horse, the cow,
&c., which shed their winter coat in the spring."
A slight difference in climate or pasture sometimes
slightly affects the fleece, as has been observed even in
different districts in England, and as is well shown by
the great softness of the wool brought from Southern
Australia. But it should be observed, as Youatt repeat-
edly insists, that the tendency to change may generally
be counteracted by careful selection. M. Lasterye, after
discussing this subject, sums up as follows:
follows: “The
preser-
vation of the Merino race in its utmost purity at the Cape
of Good Hope, in the marshes of Holland, and under the
rigorous climate of Sweden, furnishes an additional sup-
7 93
92 Youаtt on Sheep, p. 69, where Lord
Somerville is quoted. See p. 117, on the
presence of wool under the hair. With
respect to the fleeces of Australian sheep,
p. 185. On selection counteracting any
tendency to change, see pp. 70, 117, 120,
168.
93 Audubon and Bachman,' The Quad-
rupeds of North America,' 1846, vol. v.
p. 365.

126
CHAP. III.
SHEEP.
94
port of this my unalterable principle, that fine-wooled
sheep may be kept wherever industrious men and intelli-
gent breeders exist.'
That methodical selection has effected great changes
in several breeds of sheep no one, who knows anything
on the subject, entertains a doubt. The case of the South-
downs, as improved by Ellman, offers perhaps the most
striking instance. Unconscious or occasional selection
has likewise slowly produced a great effect, as we shall
see in the chapters on Selection. That crossing has large-
ly modified some breeds, no one who will study what has
been written on this subject-for instance, Mr. Spooner's
paper-will dispute; but to produce uniformity in a
crossed breed, careful selection and “rigorous weeding,"
as this author expresses it, are indispensable.'
In some few instances new breeds have suddenly ori-
ginated; thus, in 1791, a ram-lamb was born in Massachu-
setts, having short crooked legs and a long back, like a
turnspit-dog. From this one lamb the otter or ancon
semi-monstrous breed was raised; as these sheep could
not leap over the fences, it was thought that they would
be valuable; but they have been supplanted by merinos,
and thus exterminated. These sheep are remarkable from
transmitting their character so truly that Colonel Hum-
phreys never heard of “but one questionable case of
an ancon ram and ewe not producing ancon offspring.
When they are crossed with other breeds the offspring,
with rare exceptions, instead of being intermediate in
character, perfectly resemble either parent; and this has
occurred even in the case of twins. Lastly, “the ancons
have been observed to keep together, separating them-
selves from the rest of the flock when put into enclosures
with other sheep.”
A more interesting case has been recorded in the Re-
94 Journal of R. Agricult. Soc of Eng-
land,' vol. xx., part ii. W. C. Spooner on
Cross-Breeding.
95 Philosoph. Transactions,' London,
1813, p. 88.

CHAP. III.
127
GOATS.
port of the Juries for the Great Exhibition (1851), namely,
the production of a merino ram-lamb on the Mauchamp
farm, in 1828, which was remarkable for its long, smooth,
straight, and silky wool. By the year 1833 M. Graux
had raised rams enough to serve his whole flock, and after
a few more years he was able to sell stock of his new
breed. So peculiar and valuable is the wool, that it sells
at 25 per cent. above the best merino wool: even the
fleeces of half-bred animals are valuable, and are known
in France as the "Mauchamp-merino." It is interesting,
as showing how generally any marked deviation of struc-
ture is accompanied by other deviations, that the first
ram and his immediate offspring were of small size, with
large heads, long necks, narrow chests, and long flanks;
but these blemishes were removed by judicious crosses
and selection. The long smooth wool was also correlated
with smooth horns; and as horns and hair are homolo-
gous structures, we can understand the meaning of this
correlation. If the Mauchamp and ancon breeds had
originated a century or two ago, we should have had no
record of their birth; and many a naturalist would no
doubt have insisted, especially in the case of the Mau-
champ race, that they had each descended from, or been
crossed with, some unknown aboriginal form.
GOATS.
FROM the recent researches of M. Brandt, most natural-
ists now believe that all our goats are descended from
the Capra ægagrus of the mountains of Asia, possibly
mingled with the allied Indian species C. Falconeri of
India.98 In Switzerland, during the early Stone period,
the domestic goat was commoner than the sheep; and
96 Isidore Geoffroy St. Hilaire, Hist.
Nat. Générale,' tom. iii. p. 87. Mr.
Blyth ('Land and Water, 1867, p. 37)
has arrived at a similar conclusion, but
he thinks that certain Eastern races
may perhaps be in part descended from
the Asiatic markhor.

128
CHAP. III.
GOATS.
this very ancient race differed in no respect from that
now common in Switzerland. At the present time, the
many races found in several parts of the world differ
greatly from each other; nevertheless, as far as they
have been tried, they are all quite fertile when crossed.
So numerous are the breeds, that Mr. G. Clark a has
described eight distinct kinds imported into the one
island of Mauritius. The ears of one kind were enor-
mously developed, being, as measured by Mr. Clark, no
less than 19 inches in length and 4 inches in breadth.
As with cattle, the mamme of those breeds which are
regularly milked become greatly developed; and, as Mr.
Clark remarks, “it is not rare to see their teats touching
the ground." The following cases are worth notice as
presenting unusual points of variation. According to
Godron,100 the mamme differ greatly in shape in different
breeds, being elongated in the common goat, hemisphe-
rical in the Angora race, and bilobed and divergent in
the goats of Syria and Nubia. According to this same
author, the males of certain breeds have lost their usual
offensive odour. In one of the Indian breeds the males
and females have horns of widely-different shapes ;
in some breeds the females are destitute of horns. 102 The
presence of interdigital pits or glands on all four feet
has been thought to characterise the genus Ovis, and
their absence to be characteristic of the genus Capra ;
but Mr. Hodgson has found that they exist in the front
101 and
97 Rütimeyer, Pfahlbauten,' s. 127.
98 Godron, De l'Espèce,' tom. i. p.
402.
99 Annals and Mag. of Nat. History,
vol. ii. (2nd series), 1848, p. 363.
100 De l'Espèce,' tom. i. p. 406. Mr.
Clark also refers to differences in the
shape of the mammæ. Godron states
that in the Nubian race the scrotum is
divided into two lobes; and Mr. Clark
gives a ludicrous proof of this fact, for
he saw in the Mauritius a male goat of
the Muscat breed purchased at a high
price for a female in full milk. These
differences in the scrotum are probably
not due to descent from distinct species;
for Mr. Clark states that this part varies
much in form.
101 Mr. Clark, 'Annals and Mag. of
Nat. Hist.,' vol. ii. (2nd series), 1848, P.
361.
102 Desmarest, 'Encyclop. Méthod.
Mammalogie,' p. 450.

CHAP. III.
129
GOATS.
103
feet of the majority of Himalayan goats."
Mr. Hodg-
son measured the intestines in two goats of the Dúgú
race, and he found that the proportional length of the
great and small intestines differed considerably. In one
of these goats the cæcum was thirteen inches, and in the
other no less than thirty-six inches in length!
103 Journal of Asiatic Soc. of Bengal,' vol. xvi., 1847, pp. 1020, 1025.

130
CHAP. IV.
DOMESTIC RABBITS.
CHAPTER IV.
DOMESTIC RABBITS.
DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RAB-
BIT — ANCIENT DOMESTICATION - ANCIENT SELECTION - LARGE
LOP-EARED RABBITS — VARIOUS BREEDS - FLUCTUATING CIA-
RACTERS ORIGIN OF THE HIMALAYAN BREED — CURIOUS CASE
OF INHERITANCE — FERAL RABBITS IN JAMAICA AND THE FALK-
LAND ISLANDS — PORTO SANTO FERAL RABBITS — OSTEOLOGICAL
CHARACTERS - - SKULL- SKULL OF HALF-LOP RABBITS - VARIA-
TIONS IN THE SKULL ANALOGOUS TO DIFFERENCES IN DIFFER-
ENT SPECIES OF HARES — VERTEBRÆSTERNUM — SCAPULA
EFFECTS OF USE AND DISUSE ON THE PROPORTIONS OF THE
LIMBS AND BODY - CAPACITY OF THE SKULL AND REDUCED
SIZE OF THE BRAIN SUMMARY ON THE MODIFICATIONS OF
DOMESTICATED RABBITS.
All naturalists, with, as far as I know, a single exception,
believe that the several domestic breeds of the rabbit are
descended from the common wild species; I shall there-
fore describe them more carefully than in the previous
cases. Professor Gervais ? states that the true wild rab-
bit is smaller than the domestic; its proportions are not
absolutely the same; its tail is smaller; its ears are shorter
and more thickly clothed with hair; and these characters,
without speaking of colour, are so many
indications
OP-
posed to the opinion which unites these animals under the
same specific denomination." Few naturalists will agree
with this author that such slight differences are sufficient
to separate as distinct species the wild and domestic rab-
bit. How extraordinary it would be, if close confinement,
1 M. P. Gervais, Hist. Nat. des Mammifères,' tom. i., 1834, p. 288.

CHAP. IV.
131
THEIR PARENTAGE.
perfect tameness, unnatural food, and careful breeding, all
prolonged during many generations, had not produced at
least some effect! The tame rabbit has been domesticated
from an ancient period. Confucius ranges
rabbits
among
animals worthy to be sacrificed to the gods, and, as he
prescribes their multiplication, they were probably at this
early period domesticated in China. They are mentioned
by several of the classical writers. In 1631 Gervaise
Markham writes, “You shall not, as in other cattell, looke
to their shape, but to their richnesse, onely elect your
buckes, the largest and goodliest conies you can get; and
for the richnesse of the skin, that is accounted the richest
which hath the equallest mixture of blacke and white
haire together, yet the blacke rather shadowing the white;
the furre should be thicke, deepe, smooth, and shining;
they are of body much fatter and larger, and, when
another skin is worth two or three pence, they are worth
two shillings. From this full description we see that
silver-grey rabbits existed in England at this period; and,
what is far more important, we see that the breeding or
selection of rabbits was then carefully attended to. Al-
drovandi, in 1637, describes, on the authority of several
old writers (as Scaliger, in 1557), rabbits of various co-
lours, some “like a hare," and he adds that P. Valerianus
(who died a very old man in 1558) saw at Verona rabbits
four times bigger than ours.?
From this fact of the rabbit having been domesticated
at an ancient period, we must look to the northern hemi-
sphere of the Old World, and to the warmer temperate re-
gions alone, for the aboriginal parent-form; for the rabbit
cannot live without protection in countries as cold as Swe-
den, and, though it has run wild in the tropical island of
Jamaica, it has never greatly multiplied there. It now ex-
2 U. Aldrovandi, 'De Quadrupedibus studied the subject in Cottage Garden-
digitatis,' 1637, p. 383. For Confucius er,' Jan. 22nd, 1861, p. 250.
and G. Markham, see a writer who has

132
CHAP. IV.
DOMESTIC RABBITS.
ists, and has long existed, in the warmer temperate parts
of Europe, for fossil remains have been found in several
countries. The domestic rabbit readily becomes feral in
these same countries, and when variously coloured kinds
are turned out they generally revert to the ordinary grey
colour. The wild rabbits, if taken young, can be domes-
ticated, though the process is generally very troublesome.”
The various domestic races are often crossed, and are be-
lieved to be perfectly fertile together, and a perfect gra-
dation can be shown to exist from the largest domestic
kinds, having enormously developed ears, to the common
wild kind. The parent-form must have been a burrowing
animal, a habit not common, as far as I can discover, to
any other species in the large genus Lepus. Only one
wild species is known with certainty to exist in Europe;
but the rabbit (if it be a true rabbit) from Mount Sinai,
and likewise that from Algeria, present slight differences,
and these forms have been considered by some authors as
specifically distinct. But such slight differences would aid
us little in explaining the more considerable differences
characteristic of the several domestic races. If the latter
are the descendants of two or more closely allied species,
all, excepting the common rabbit, have been exterminated
in a wild state; and this is very improbable, seeing with
what pertinacity this animal holds its ground. From these
several reasons we may infer with safety that all the domes-
tic breeds are the descendants of the common wild spe-
cies. But from what we hear of the late marvellous suc-
cess in rearing hybrids between the hare and rabbit," it is
3 Owen, 'British Fossil Mammals,' p.
212.
4 Bechstein, Naturgesch. Deutsch-
lands,' 1801, b. i. p. 1183. I have re-
ceived similar accounts with respect to
England and Scotland.
5 Pigeons and Rabbits,' by E. S. De-
lamer, 1854, p. 133. Sir J. Sebright (Ob-
servations on Instinct, 1836, p. 10)
speaks most strongly on the difficulty.
But this difficulty is not invariable, as
I have received two accounts of perfect
success in taming and breeding from the
wild rabbit. See also Dr. P. Broca, in
Journal de la Physiologie,' tom. ii. p.
368.
6 Gervais, Hist. Nat. des Mammi-
fères,' tom. i. p. 292.
7 See Dr. P. Broca's interesting me-
moir on this subject in Brown-Sequard's
Journ. de Phys.,' vol. ii. p. 367.

CHAP. IV.
133
THEIR VARIATION.
possible, though not probable, from the great difficulty
in making the first cross, that some of the larger races,
which are coloured like the hare, may have been modified
by crosses with this animal. Nevertheless, the chief dif-
ferences in the skeletons of the several domestic breeds
cannot, as we shall presently see, have been derived from
a cross with the hare.
There are many breeds which transmit their characters
more or less truly. Every one has seen the enormous
lop-eared rabbits exhibited at our shows; various allied
sub-breeds are reared on the Continent, such as the so-
called Andalusian, which is said to have a large head
with a round forehead, and to attain a greater size than
any other kind; another large Paris breed is named the
Rouennais, and has a square head; the so-called Pa-
tagonian rabbit has remarkably short ears and a large
round head. Although I have not seen all these breeds,
I feel some doubt about there being any marked difference
in the shape of their skulls. English lop-eared rabbits
often weigh 8 lbs. or 10 lbs., and one has been exhibited
weighing 18 lbs. ; whereas a full-sized wild rabbit weighs
only about 31 lbs. The head or skull in all the large lop-
eared rabbits examined by me is much longer relatively
to its breadth than in the wild rabbit. Many of them
have loose transverse folds of skin or dewlaps beneath the
throat, which can be pulled out so as to reach nearly to
the ends of the jaws. Their ears are prodigiously develop-
ed, and hang down on each side of their faces. A rabbit
has been exhibited with its two ears, measured from the
tip of one to the tip of the other, 22 inches in length, and
each ear was 53 inches in breadth. In a common wild
rabbit I found that the length of the two ears, from tip to
tip, was 75 inches, and the breadth only 13 inch. The
great weight of the body in the larger rabbits, and the
8 They are briefly described in the 'Journal of Horticulture,' May 7th, 1861, p. 108.

134
CHAP. IV.
DOMESTIC RABBITS.
immense development of their ears, are the qualities
which win prizes, and have been carefully selected.
The hare-coloured, or, as it is sometimes called, the Bel-
gian rabbit, differs in nothing except colour from the
other large breeds; but Mr. J. Young, of Southampton,
a great breeder of this kind, informs me that the females,
in all the specimens examined by him, had only six mam-
mæ; and this certainly was the case with two females
which came into my possession. Mr. B. P. Brent, how-
ever, assures me that the number is variable with other
domestic rabbits. The common wild rabbit always has
ten mammæ. The Angora rabbit is remarkable from the
length and fineness of its fur, which even on the soles of
the feet is of considerable length. This breed is the only
one which differs in its mental qualities, for it is said to
be much more sociable than other rabbits, and the male
shows no wish to destroy its young' Two live rabbits
were brought to me from Moscow, of about the size of
the wild species, but with long soft fur, different from
that of the Angora. These Moscow rabbits had pink
eyes and were snow-white, excepting the ears, two spots
near the nose, the upper and under surface of the tail,
and the hinder tarsi, which were blackish-brown. In
short, they were coloured nearly like the so-called Hima-
layan rabbits, presently to be described, and differed
from them only in the character of their fur. There are
two other breeds which come true to colour, but differ
in no other respect, namely silver-greys and chinchillas.
Lastly, the Nicard or Dutch rabbit may be mentioned,
which varies in colour, and is remarkable from its small
size, some specimens weighing only 14 lb. ; rabbits of
this breed make excellent nurses for other and more
delicate kinds.":10
Certain characters are remarkably fluctuating, or are
10
9 Journal of Horticulture,' 1861, p.
880.
Journal of Horticulture,' May
28th, 1861, p. 169.

CHAP. IV.
135
THEIR VARIATION.
وو
very feebly transmitted by domestic rabbits: thus, one
breeder tells me that with the smaller kinds he has hardly
ever raised a whole litter of the same colour: with the
large lop-eared breeds “it is impossible,” says a great
judge, 11 “ to breed true to colour, but by judicious cross-
ing a great deal may be done towards it. The fancier
should know how his does are bred, that is, the colour of
their parents.” Nevertheless, certain colours, as we shall
presently see, are transmitted truly. The dewlap is not
strictly inherited. Lop-eared rabbits, with their ears
hanging flat down on each side of the face, do not trans-
mit this character at all truly. Mr. Delamer remarks
that, “ with fancy rabbits, when both the parents are per-
fectly formed, have model ears, and are handsomely
marked, their progeny do not invariably turn out the
same.” When one parent, or even both, are oar-laps, that
is, have their ears sticking out at right angles, or when
one parent or both are half-lops, that is, have only one ear
dependent, there is nearly as good a chance of the pro-
geny having both ears full-lop, as if both parents had
been thus characterized. But I am informed, if both pa-
rents have upright ears, there is hardly a chance of a full-
lop. In some half-lops the ear that hangs down is broader
and longer than the upright ear; so that we have the
unusual case of a want of symmetry on the two sides.
This difference in the position and size of the two ears
probably indicates that the lopping of the ear results
from its great length and weight, favored no doubt by
the weakness of the muscles consequent on disuse. An-
derson 1 mentions a breed having only a single ear; and
Professor Gervais another breed which is destitute of
ears.
12
11 Journal of Horticulture, 1861, p.
327.
With respect to the ears, 802
Delamer on 'Pigeons and Rabbits,'1854,
p. 141 ; also "Poultry Chronicle,' vol. ii.
p. 499, and ditto for 1854, p. 586.
12 Delamer, 'Pigeons and Rabbits,' p.
136. See also Journal of Horticulture,'
1861, p. 375.
13 An Account of the different kinds
of Sheep in the Russian Dominions,
1794, p. 39.

136
CHAP. IV.
DOMESTIC RABBITS.
Fig. 5.-Half-lop Rabbit. (Copied from E. S. Delamer's work.)
The origin of the Himalayan breed (sometimes called
Chinese, or Polish, or Russian) is so curious, both in itself,
and as throwing some light on the complex laws of in-
heritance, that it is worth giving in detail. These pretty
rabbits are white, except their ears, nose, all four feet, and
the upper side of tail, which are all brownish-black; but
as they have red eyes, they may be considered as albinoes.
I have received several accounts of their breeding per-
fectly true. From their symmetrical marks, they were
at first ranked as specifically distinct, and were provision-
ally named L. nigripes.14 Some good observers thought
that they could detect a difference in their habits, and
stoutly maintained that they formed a new species. Their
origin is now well known. A writer, in 1857, stated that
he had produced Himalayan rabbits in the following man-
ner. But it is first necessary briefly to describe two other
breeds: silver-greys or silver-sprigs generally have black
14 Proc. Zoolog. Soc.,' June 23rd, 1857, p. 159.
15 Cottage Gardener,' 1857, p. 141.

CHAP. IV.
137
THE HIMALAYAN BREED.
heads and legs, and their fine grey fur is interspersed with
numerous black and white long hairs. They breed perfect-
ly true, and have long been kept in warrens. When they
escape and cross with common rabbits, the product, as I
hear from Mr. Wyrley Birch, of Wretham Hall, is not a
mixture of the two colours, but about half take after the
one parent, and the other half after the other parent. Sec-
ondly, chinchillas or tame silver-greys (I will use the for-
mer name) have short, paler, mouse or slate-coloured fur,
interspersed with long, blackish, slate-coloured, and white
hairs. 16
These rabbits breed perfectly true. Now, the
writer above referred to had a breed of chinchillas which
had been crossed with the common black rabbit, and their
offspring were either blacks or chinchillas. These latter
were again crossed with other chinchillas (which had also
been crossed with silver-greys), and from this complicated
cross Himalayan rabbits were raised. From these and
other similar statements, Mr. Bartlett was led to make
a careful trial in the Zoological Gardens, and he found
that by simply crossing silver-greys with chinchillas he
could always produce some few Himalayans; and the lat-
ter, notwithstanding their sudden origin, if kept separate,
bred perfectly true.
The Himalayans, when first born, are quite white, and
are then true albinoes; but in the course of a few months
they gradually assume their dark ears, nose, feet, and tail.
Occasionally, however, as I am informed by Mr. W. A.
Wooler and the Rev. W. D. Fox, the young are born of
a very pale grey colour, and specimens of such fur were
sent me by the former gentleman. The grey tint, how-
ever, disappears as the animal comes to maturity. So
that with these Himalayans there is a tendency, strictly
confined to early youth, to revert to the colour of the
adult silver-grey parent-stock. Silver-greys and chin-
16 Journal of Horticulture,' April
9th, 1861, p. 35.
17 Mr. Bartlett, in Proc. Zoolog. Soc.,'
1861, p. 40.

138
CHAP. IV.
DOMESTIC RABBITS.
chillas, on the other hand, present a remarkable contrast
in their colour whilst quite young, for they are born
perfectly black, but soon assume their characteristic
grey or silver tints. The same thing occurs with grey
horses, which, as long as they are foals, are generally of
a nearly black colour, but soon become grey, and get
whiter and whiter as they grow older. Hence the usual
rule is that Himalayans are born white and afterwards
become in certain parts of their bodies dark-coloured ;
whilst silver-greys are born black and afterwards become
sprinkled with white. Exceptions, however, and of a
directly opposite nature, occasionally occur in both cases.
For young silver-greys are sometimes born in warrens,
as I hear from Mr. W. Birch, of a cream-colour, but
these young animals ultimately become black. The
Himalayans, on the other hand, sometimes produce, as is
stated by an experienced amateur,18 a single black young
one in a litter; but such, before two months elapse,
become perfectly white.
To sum up the whole curious 'case : wild silver-greys
may be considered as black rabbits which become grey
at an early period of life. When they are crossed with
common rabbits, the offspring are said not to have
blended colours, but to take after either parent; and in
this respect they resemble black and albino varieties of
most quadrupeds, which often transmit their colours in
this same manner. When they are crossed with chin-
chillas, that is, with a paler sub-variety, the young are
at first pure albinoes, but soon become dark-coloured in
certain parts of their bodies, and are then called Hima-
layans. The young Himalayans, however, are sometimes
at first either pale grey or completely black, in either
case changing after to white. In a future chapter I
shall advance a large body of facts showing that, when
18 Phenomenon in Himalayan Rabbits,' in Journal of Horticulture, ' 1865, Jan.
27th, p. 102.

CHAP. IV.
139
FERAL RABBITS.
two varieties are crossed both of which differ in colour
from their parent-stock, there is a strong tendency in the
young to revert to the aboriginal colour; and what is
very remarkable, this reversion occasionally supervenes,
not before birth, but during the growth of the animal.
Hence, if it could be shown that silver-greys and chin-
chillas were the offspring of a cross between a black and
albino variety with the colours intimately blended-a
supposition in itself not improbable, and supported by
the circumstance of silver-greys in warrens sometimes
producing creamy-white young, which ultimately become
black—then all the above-given paradoxical facts on the
changes of colour in silver-greys and in their descendants
the Himalayans would come under the law of reversion,
supervening at different periods of growth and in differ-
ent degrees, either to the original black or to the ori-
ginal albino parent-variety.
It is, also, remarkable that Himalayans, though produced
so suddenly, breed true. But as, whilst young, they are
albinoes, the case falls under a very general rule; for
albinism is well known to be strongly inherited, as with
white mice and many other quadrupeds, and even with
white flowers. But why, it may be asked, do the ears,
tail, nose, and feet, and no other part of the body, revert
to a black colour? This apparently depends on a law,
which generally holds good, namely, that characters com-
mon to many species of a genus-and this, in fact, implies
long inheritance in common from the ancient progenitor of
the genus—are found to resist variation, or to reappear if
lost, more persistently than the characters which are con-
fined to the separate species. Now, in the genus Lepus,
a large majority of the species have their ears and the
upper surface of the tail tinted black; but the persistence
of these marks is best seen in those species which in win-
ter become white: thus, in Scotland the L. variabilis 19
19 G. R. Waterhouse, Natural History of Mammalia: Rodents,' 1846, pp. 52, 60,
105.

140
CHAP. IV.
DOMESTIC RABBITS.
in its winter dress has a shade of colour on its nose, and
the tips of its ears are black: in the L. tibetanus the ears
are black, the upper surface of the tail greyish black, and
the soles of the feet brown: in L. glacialis the winter fur
is
pure white, except the soles of the feet and the points
of the ears. Even in the variously-coloured fancy rabbits
we may often observe a tendency in these same parts to
be more darkly tinted than the rest of the body. Thus,
as it seems to me, the appearance of the several coloured
marks on the Himalayan rabbit, as it grows old, is ren-
dered intelligible. I may add a nearly analogous case:
fancy rabbits very often have a white star on their fore-
heads; and the common English hare, whilst young, gen-
erally has, as I have myself observed, a similar white star
on its forehead.
When variously coloured rabbits are set free in Europe,
and are thus placed under their natural conditions, they
generally revert to the aboriginal grey colour; this may
be in part due to the tendency in all crossed animals, as
lately observed, to revert to their primordial state. But
this tendency does not always prevail; thus silver-grey
rabbits are kept in warrens, and remain true though liv-
ing almost in a state of nature; but a warren must not
be stocked with both silver-greys and common rabbits;
otherwise “in a few years there will be none but common
greys surviving." When rabbits run wild in foreign
countries, under different conditions of life, they by no
means always revert to their aboriginal colour. In Ja-
maica the feral rabbits are described as “slate-coloured,
deeply tinted with sprinklings of white on the neck, on
the shoulders, and on the back; softening off to blue-
white under the breast and belly.”21 But in this tropi-
9 20
20 Delamer on Pigeons and Rabbits,'
p.
114.
21 Gosse's Sojourn in Jamaica,' 1851,
p. 441, as described by an excellent ob-
server, Mr. R. Hill. This is the only
known case in which rabbits have be-
come feral in a hot country. They can
be kept, however, at Loanda (see Living-
stone's Travels,' p. 407). In parts c:
India, as I am informed by Mr. Blyth,
they breed well.

CHAP. IV,
141
FERAL RABBITS.
cal island the conditions were not favourable to their
increase, and they never spread widely; and, as I hear
from Mr. R. Hill, owing to a great fire which occurred
in the woods, they have now become extinct. Rabbits
during many years have run wild in the Falkland Islands;
they are abundant in certain parts, but do not spread ex-
tensively. Most of them are of the common grey colour;
a few, as I am informed by Admiral Sulivan, are hare-
coloured, and many are black, often with nearly symme-
trical white marks on their faces. Hence, M. Lesson
described the black variety as a distinct species, under the
name of Lepus magellanicus, but this, as I have elsewhere
shown, is an error. 22 Within recent times the sealers
have stocked some of the small outlying islets in the
Falkland group with rabbits; and on Pebble Islet, as I
hear from Admiral Sulivan, a large proportion are bare-
coloured, whereas on Rabbit Islet a large proportion are
of a bluish colour which is not elsewhere seen. How the
rabbits were coloured which were turned out on these
islets is not known.
The rabbits which have become feral on the island of
Porto Santo, near Madeira, deserve a fuller account. In
1418 or 1419, J. Gonzales Zarco happened to have
a female rabbit on board which had produced young
during the voyage, and he turned them all out on the
island. These animals soon increased so rapidly, that
they became a nuisance, and actually caused the aban-
donment of the settlement. Thirty-seven years subse-
quently, Cada Mosto describes them as innumerable ; nor
is this surprising, as the island was not inhabited by any
beast of prey or by any terrestrial mammal. We do not
know the character of the mother-rabbit; but we have
22 Darwin's Journal of Researches,
p. 193; and 'Zoology of the Voyage of
the Beagle : Mammalia,' p. 92.
23 Kerr's Collection of Voyages,' vol.
ii. p. 177; p. 205 for Cada Mosto. Ac-
cording to a work published in Lisbon
in 1717, entitled Historia Insulana,'
written by a Jesuit, the rabbits were
turned out in 1420. Some authors be-
lieve that the island was discovered in
1413.

142
CHAP. IV.
DOMESTIC RABBITS.
every reason to believe that it was the common domesti-
cated kind. The Spanish peninsula, whence Zarco sailed,
is known to have abounded with the common wild spe-
cies at the most remote historical period. As these rab-
bits were taken on board for food, it is improbable that
they should have been of any peculiar breed. That the
breed was well domesticated is shown by the doe having
littered during the voyage. Mr. Wollaston, at my re-
quest, brought home two of these feral rabbits in spirits
of wine; and, subsequently, Mr. W. Haywood sent to
me three more specimens in brine, and two alive. These
seven specimens, though caught at different periods,
closely resembled each other. They were full grown, as
shown by the state of their bonés. Although the condi-
tions of life in Porto Santo are evidently highly favour-
able to rabbits, as proved by their extraordinarily rapid
increase, yet they differ conspicuously in their small size
from the wild English rabbit. Four English rabbits,
measured from the incisors to the anus, varied between
17 and 17 inches in length; whilst two of the Porto
Santo rabbits were only 141 and 15 inches in length.
But the decrease in size is best shown by weight; four
wild English rabbits averaged 3 lb. 5 oz., whilst one of
the Porto Santo rabbits, which had lived for four years
in the Zoological Gardens, but had become thin, weighed
only 1 lb. 9 oz. A fairer test is afforded by the compari-
son of the well-cleaned limb-bones of a P. Santo rabbit
killed on the island with the same bones of a wild English
rabbit of average size, and they differed in the propor-
tion of rather less than five to nine. So that the Porto
Santo rabbits have decreased nearly three inches in length,
and almost half in weight of body. The head has not
24
24 Something of the same kind has oc-
curred on the island of Lipari, where,
according to Spallanzani (Voyage dans
les deux Siciles,' quoted by Godron sur
l'Espèce, p. 364), a countryman turned
out some rabbits which multiplied pro-
digiously, but, says Spallanzani, "les
lapins de l'ile de Lipari sont plus petits
que ceux qu'on élève en domesticité."

CHAP. IV.
143
FERAL RABBITS.
decreased in length proportionally with the body; and
the capacity of the brain-case is, as we shall hereafter see,
singularly variable. I prepared four skulls, and these
resembled each other more closely than do generally the
skulls of wild English rabbits; but the only difference in
structure which they presented was that the supra-orbital
processes of the frontal bones were narrower.
In colour the Porto Santo rabbit differs considerably
from the common rabbit; the upper surface is redder, and
is rarely interspersed with any black or black-tipped hairs.
The throat and certain parts of the under surface, instead
of being pure white, are generally pale grey or leaden
colour. But the most remarkable difference is in the ears
and tail; I have examined many fresh English rabbits,
and the large collection of skins in the British Museum
from various countries, and all have the upper surface of
the tail and the tips of the ears clothed with blackish-
grey fur; and this is given in most works as one of the
specific characters of the rabbit. Now in the seven Porto
Santo rabbits the upper surface of the tail was reddish-
brown, and the tips of the ears had no trace of the black
edging. But here we meet with a singular circumstance:
in June, 1861, I examined two of these rabbits recently
sent to the Zoological Gardens, and their tails and ears
were coloured as just described; but when one of their
dead bodies was sent to me in February, 1865, the ears
were plainly edged, and the upper surface of the tail was
covered, with blackish-grey fur, and the whole body was
much less red; so that under the English climate this
individual rabbit had recovered the proper colour of its
fur in rather less than four years!
The two little Porto Santo rabbits, whilst alive in the
Zoological Gardens, had a remarkably different appear-
ance from the common kind. They were extraordinarily
wild and active, so that many persons exclaimed on seeing
them that they were more like large rats than rabbits.
They were nocturnal to an unusual degree in their habits,

144
CHAP. IV.
DOMESTIC RABBITS.
and their wildness was never in the least subdued ; so
• that the superintendent, Mr. Bartlett, assured me that he
had never had a wilder animal under his charge. This
is a singular fact, considering that they are descended
from a domesticated breed; I was so much surprised at
it, that I requested Mr. Haywood to make inquiries on
the spot, whether they were much hunted by the inhabi.
tants, or persecuted by hawks, or cats, or other animals;
but this is not the case, and no cause can be assigned for
their wildness. They live on the central, higher rocky
land and near the sea-cliffs, and, being exceedingly shy
and timid, seldom appear in the lower and cultivated
districts. They are said to produce from four to six
young at a birth, and their breeding season is in July
and August. Lastly, and this is a highly remarkable
fact, Mr. Bartlett could never succeed in getting these
two rabbits, which were both males, to associate or breed
with the females of several breeds which were repeatedly
placed with them.
If the history of these Porto Santo rabbits had not been
known, most naturalists, on observing their much reduced
size, their reddish colour above and grey beneath, with
neither tail nor ears tipped with black, would have ranked
them as a distinct species. They would have been strongly
confirmed in this view by seeing them alive in the Zoo-
logical Gardens, and hearing that they refused to couple
with other rabbits. Yet this rabbit, which there can be
little doubt would thus have been ranked as a distinct
species, has certainly originated since the year 1420. Fi-
nally, from the three cases of the rabbits which have run
wild in Porto Santo, Jamaica, and the Falkland Islands,
we see that these animals do not, under new conditions of
life, revert to or retain their aboriginal character, as is so
generally asserted to be the case by most authors.

Chap. IV. DIFFERENCES IN THEIR SKELETONS. 145
Osteological Characters.
When we remember, on the one hand, how frequently
it is stated that important parts of the structure never
vary; and, on the other hand, on what small differences
in the skeleton fossil species have often been founded,
the variability of the skull and of some other bones in the
domesticated rabbit well deserves attention. It must not
be supposed that the more important differences imme-
diately to be described strictly characterise any one
breed; all that can be said is, that they are generally pre-
sent in certain breeds. We should bear in mind that selec-
tion has not been applied to fix any character in the skel-
eton, and that the animals have not had to support them-
selves under uniform habits of life. We cannot account
for most of the differences in the skeleton; but we shall
see that the increased size of the body, due to careful
nurture and continued selection, has affected the head in
a particular manner. Even the elongation and lopping
of the ears have influenced in a small degree the form of
the whole skull. The want of exercise has apparently
modified the proportional length of the limbs in compari-
son with the body.
As a standard of comparison, I prepared skeletons of two wild rab-
bits from Kent, one from the Shetland Islands, and one from Antrim
in Ireland. As all the bones in these four specimens from such dis-
tant localities closely resembled each other, presenting scarcely any
appreciable difference, it may be concluded that the bones of the
wild rabbit are generally uniform in character.
Skull.--I have carefully examined skulls of ten large lop-eared fan-
cy rabbits, and of five common domestic rabbits, which latter differ
from the lop-eared only in not having such large bodies or ears, yet
both larger than in the wild rabbit. First for the ten lop-eared rab-
bits: in all these the skull is remarkably elongated in comparison
with its breadth. In a wild rabbit the length was 3.15 inches, in a
large fancy rabbit 4:30; whilst the breadth of the cranium enclosing
the brain was in both almost exactly the same. Even by taking as the
standard of comparison the widest part of the zygomatic arch, the
7

146
CHAP. IV
DOMESTIC RABBITS.
skulls of the lop-eared are proportionally to their breadth three-
quarters of an inch too long. The depth of the head has increased
almost in the same proportion with the length; it is the breadth
alone which has not increased. The parietal and occipital bones en-
closing the brain are less arched, both in a longitudinal and trans-
verse line, than in the wild rabbit, so that the shape of the cranium
is somewhat different. The surface is rougher, less cleanly sculp-
tured, and the lines of sutures are more prominent.
Although the skulls of the large lop-eared rabbits in comparison
with those of the wild rabbit are much elongated relatively to their
breadth, yet, relatively to the size of body, they are far from elon-
gated. The lop-eared rabbits which I examined were, though not fat,
more than twice as heavy as the wild specimens; but the skull was
very far from being twice as long. Even if we take the fairer stand-
ard of the length of body, from the nose to the anus, the skull is
not on an average as long as it ought to be by a third of an inch.
In the small feral P. Santo rabbit, on the other hand, the head rela-
tively to the length of the body is about a quarter of an inch too
long
This elongation of the skull relatively to its breadth, I find a uni-
versal character, not only with the large lop-eared rabbits, but in all
the artificial breeds; as is well seen in the skull of the Angora. I
was at first much surprised at the fact, and could not imagine why
domestication should produce this uniform result; but the explana-
tion seems to lie in the circumstance that during a number of gen-
erations the artificial races have been closely confined, and have had
little occasion to exert either their senses, or intellect, or voluntary
muscles; consequently the brain, as we shall presently more fully
see, has not increased relatively with the size of body. As the brain
has not increased, the bony case enclosing it has not increased, and
this has evidently affected through correlation the breadth of the
entire skull from end to end.
In all the skulls of the large lop-eared rabbits, the supra-orbital
plates or processes of the frontal bones are much broader than in
the wild rabbit, and they generally project more upwards. In the
zygomatic arch the posterior or projecting point of the malar-bone
is broader and blunter; and in the specimen, fig. 8; it is so in a
remarkable degree. This point approaches nearer to the auditory
meatus than in the wild rabbit, as may be best seen in fig. 8; but
this circumstance mainly depends on the changed direction of the
meatus. The inter-parietal bone (see fig. 9) differs much in shape
in the several skulls; generally it is more oval, or has a greater
width in the line of the longitudinal axis of the skull, than in the

Chap. IV. DIFFERENCES IN THEIR SKELETONS. 147
( ERSTIN LASSEN
DUNDW
Fig. 6.-Skull of Wild Rabbit, of mataral
size.
Fig. 7.—Skull of large Lop-eared Rabbit, of natu-
ral size.
wild rabbit. The posterior margin of "the square raised plat-
form" 25 of the occiput, instead of being truncated, or projecting
slightly as in the wild rabbit, is in most lop-eared rabbits pointed,
25 Waterhouse, Nat. Hist. Mammalia,' pol. ii. p. 36.

148
CHAP. IV.
DOMESTIC RABBITS.
as in fig. 9, C. The paramas-
toids relatively to the size of
the skull are generally much
thicker than in the wild rab-
bit.
The occipital foramen (fig.
10) presents some remarkable
differences: in the wild rabbit,
the lower edge between the
condyles is considerably and
almost angularly hollowed out,
and the upper edge is deeply
and squarely notched; hence
the longitudinal axis exceeds
the transverse axis. In the
Fig. 8.- Part of Zygomatic Arch, showing skulls of the lop-eared rabbits
the projecting end of the malar bone, and
the transverse axis exceeds the
the auditory meatus : of natural size.
Upper figure, Wild Rabbit. Lower figure, longitudinal; for in none of
Lop-eared, hare-coloured Rabbit.
these skulls was the lower edge
between the condyles so deeply
hollowed out; in five of them there was no upper square notch, in
three there was a trace of the notch, and in two alone it was well
A
B
C developed. These differ-
ences in the shape of the
foramen are remarkable,
considering that it gives
passage to so important
a structure as the spinal
marrow, though appa-
Fig. 9.-Posterior end of Skull, of natural size, rently the outline of the
showing the inter-parietal bone. A. Wild Rabbit.
latter is not affected by
B. Feral Rabbit from island of P. Santo, near
the shape of the passage.
Madeira. C. Large Lop-eared Rabbit.
In all the skulls of the
large lop-eared rabbits, the bony auditory meatus is conspicuously
larger than in the wild rabbit. In a skull 4.3 inches in length, and
which barely exceeded
А
in breadth the skull of
a wild rabbit (which was
3.15 inches in length),
the longer diameter of
the meatus was exactly
twice as great. The ori-
Fig. 10.-Occipital Foramen of natural size, in-
fice is more compressed,
A. Wild Rabbit; B. Large Lop-eared Rabbit. and its margin on the
B

CHAP. IV. DIFFERENCES IN THEIR SKELETONS. 149
side nearest the skull stands up higher than the outer side. The
whole meatus is directed more forwards. As in breeding lop-eared
rabbits the length of the ears, and their consequent lopping and ly-
ing flat on the face, are the
chief points of excellence,
there can hardly be a doubt
that the great change in
the size, form, and direction
of the bony meatus, rela-
tively to this same part in
the wild rabbit, is due to
the continued selection of
individuals having larger
and larger ears. The influ-
ence of the external ear on
the bony meatus is well
shown in the skulls (I have
examined three) of half-lops
(see fig. 5), in which one ear
stands upright, and the
other and longer ear hang's
down; for in these skulls
there was a plain difference
in the form and direction of
the bony meatus on the two
sides. But it is a much
more interesting fact, that
the changed direction and
increased size of the bony
meatus have slightly affect-
ed on the same side the
structure of the whole skull.
I here give a drawing of the
skull of a half-lop; and it
Fig. 11.-Skull of natural size of Half-lop Rab- may be observed that the
bit, showing the different direction of the au- suture between the parietal
ditory meatus on the two sides, and the con- and frontal bones does not
sequent general distortion of the skull. The
run strictly at right angles
left ear of the animal (or right side of the fig-
ure) lopped forwards.
to the longitudinal axis of
the skull; the left frontal
bone projects beyond the right one; both the posterior and anterior
margins of the left zygomatic arch on the side of the lopping ear
stand a little in advance of the corresponding bones on the opposite
side. Even the lower jaw is affected, and the condyles are not quite

150
CHAP. IV.
DOMESTIC RABBITS.
symmetrical, that on the left standing a little in advance of that
on the right. This seems to me a remarkable case of correlation
of growth. Who would have surmised that by keeping an animal
during many generations under confinement, and so leading to the
disuse of the muscles of the ears, and by continually selecting indi-
viduals with the longest and largest ears, he would thus indirectly
have affected almost every suture in the skull and the form of the
lower jaw !
In the large lop-eared rabbits the only difference in the lower jaw,
in comparison with that of the wild rabbit, is that the posterior margin
of the ascending ramus is broader and more infected. The teeth in
neither jaw present any difference, except that the small incisors,
beneath the large ones, are proportionally a little longer. The mo-
lar teeth have increased in size proportionally with the increased
width of the skull, measured across the zygomatic arch, and not
proportionally with its increased length. The inner line of the
sockets of the molar teeth in the upper jaw of the wild rabbit forms
a perfectly straight line; but in some of the largest skulls of the
lop-eared this line was plainly bowed inwards. In one specimen
there was an additional molar tooth on each side of the upper jaw,
between the molars and premolars; but these two teeth did not
correspond in size; and as no rodent has seven molars, this is merely
a monstrosity, though a curious one.
The five other skulls of common domestic rabbits, some of which
approach in size the above-described largest skulls, whilst the others
exceed but little those of the wild rabbit, are only worth notice as
presenting a perfect gradation in all the above-specified differences
between the skulls of the largest lop-eared and wild rabbits. In all,
however, the supra-orbital plates are rather larger, and in all the
auditory meatus is larger, in conformity with the increased size of the
external ears, than in the wild rabbit. The lower notch in the oc-
cipital foramen in some was not so deep as in the wild, but in all
five skulls the upper notch was well developed.
The skull of the Angora rabbit, like the latter five skulls, is inter-
mediate in general proportions, and in most other characters, between
those of the largest lop-eared and wild rabbits. It presents only one
singular character: though considerably longer than the skull of the
wild, the breadth measured within the posterior supra-orbital fissures
is nearly a third less than in the wild. The skulls of the silver-grey
and chinchilla and Himalayan rabbits are more elongated than in
the wild, with broader supra-orbital plates, but differ little in any
other respect, excepting that the upper and lower notches of the
occipital foramen are not so deep or so well developed. The skull
of the Moscow rabbit scarcely differs in any respect from that of the

Chap. IV. DIFFERENCES IN THEIR SKELETONS. 151
wild rabbit. In the Porto Santo feral rabits the supra-orbital plates
are generally narrower and more pointed than in our wild rabbits.
As some of the largest lop-eared rabbits of which I prepared skele-
tons were coloured almost like hares, and as these latter animals
and rabbits have, as it is affirmed, been recently crossed in France,
it might be thought that some of the above-described characters
had been derived from a cross at a remote period with the hare.
Consequently I examined skulls of the hare, but no light could thus
be thrown on the peculiarities of the skulls of the larger rabbits. It
is, however, an interesting fact, as illustrating the law that varieties
of one species often assume the characters of other species of the
same genus, that I found, on comparing the skulls of ten species of
hares in the British Museum, that they differed from each other
chiefly in the very same points in which domestic rabbits vary,-
namely, in general proportions, in the form and size of the supra-
orbital plates, in the form of the free end of the malar bone, and in
the line of suture separating the occipital and frontal bones. More-
over two eminently variable characters in the domestic rabbit, namely,
the outline of the occipital foramen and the shape of the "raised
platform ” of the occiput, were likewise variable in two instances in
the same species of hare.
Vertebrae.-The number is uni-
a
form in all the skeletons which I
have examined, with two excep-
tions, namely, in one of the small
feral Porto Santo rabbits and in one
of the largest lop-eared kinds; both
of these had as usual seven cervical,
twelve dorsal with ribs, but, in-
stead of seven lumbar, both had
eight lumbar vertebre. This is re-
markable, as Gervais gives seven
as the number for the whole genus
Lepus. The caudal vertebræ appa-
rently differ by two or three, but I
7
did not attend to them, and they Fig. 12. — Atlas Vertebræ, of natural
are difficult to count with certainty.
size; inferior surface viewed ob-
In the first cervical vertebra, liquely. Upper figure, Wild Rabbit.
or atlas, the anterior margin of Lower figure, Hare-coloured, large
the neural arch varies a little in
Lop-eared Rabbit. a, supra-median,
atlantoid process; 6, infra-median
wild specimens, being either nearly
process.
smooth, or furnished with a small
supra-median atlantoid process; I have figured a specimen with the
largest process (a) which I have seen; but it will be observed how

152
CHAP. IV.
DOMESTIC RABBITS.
inferior this is in size and different in shape to that in a large lop-
eared rabbit. In the latter, the infra-median process (6) is also pro-
portionally much thicker and longer. The alæ are a little squarer
in outline.
Third cervical vertebra.-In the wild rabbit (fig. 13, A a) this
vertebra, viewed on the inferior surface, has a transverse process,
which is directed obliquely backwards, and consists of a single
pointed bar; in the fourth vertebra this process is slightly forked
in the middle. In the large lop-eared rabbits this process (B a)
is forked in the third vertebra, as in the fourth of the wild
rabbit. But the third cervical vertebræ of the wild and lop-
eared (A , B b) rabbits
А.
B
differ more conspicuous-
ly when their anterior
articular surfaces are
compared ; for the ex-
tremities of the antero-
dorsal processes in the
wild rabbit are simply
rounded, whilst in the
lop-eared they are trifid, a
a
with a deep central pit.
The canal for the spinal
marrow in the lop-eared
(B b is more elongated
Fig. 13.-Third Cervical Vertebra, of natural
in a transverse direction
size, of-A. Wild Rabbit; B. Hare-coloured,
large, Lop-eared Rabbit. a, a, inferior sur-
than in the wild rabbit;
face; 6, 6, anterior articular surfaces.
and the passages for the
arteries are of a slightly different shape. These several differences
in this vertebra seem to me well deserving attention.
First dorsal vertebra.--Its neural spine varies in length in the
wild rabbit ; being sometimes very short, but generally more than
half as long as that of the second dorsal; but I have seen it in two
large lop-eared rabbits three-fourths of the length of that of the
second dorsal vertebra.
Ninth and tenth dorsal vertebra.- In the wild rabbit the neural
spine of the ninth vertebra is just perceptibly thicker than that of
the eighth; and the neural spine of the tenth is plainly thicker and
shorter than those of all the anterior vertebræ. In the large lop-
eared rabbits the neural spines of the tenth, ninth, eighth, and even
in a slight degree that of the seventh vertebra, are very much thick-
er, and of somewhat different shape, in comparison with those of the
wild rabbit. So that this part of the vertebral column differs con-
siderably in appearance from the same part in the wild rabbit, and

CHAP. IV. DIFFERENCES IN THEIR SKELETONS. 153
А
closely resembles in an interesting manner these same vertebræ in
some species of hares. In the Angora, Chinchilla, and Himalayan
rabbits, the
neural spines
of the eighth
and ninth ver-
tebræ are in a
slight degree
thicker than in
the wild.
On
the other hand,
in one of the
feral Porto San-
to
rabbits,
B
which in most
of its characters
deviates in an
exactly орро-
site manner to
Fig. 14.- Dorsal Vertebræ, from sixth to tenth inclusive, of
what the large
natural size, viewed laterally. A. Wild Rabbit. B. Large,
Hare-coloured, so called Spanish Rabbit.
lop-eared rab-
bits do from
the common wild rabbit, the neural spines of the ninth and tenth
vertebræ were not at all larger than those of the several anterior
vertebræ. In this same Porto Santo specimen there was no trace in
the ninth vertebra of the anterior lateral processes (see woodcut 14),
which are plainly developed in all British wild rabbits, and still
more plainly developed in the large lop-eared rabbits. In a half-
wild rabbit from Sandon Park,26 a hæmal spine was moderately
well developed on the under side of the twelfth dorsal vertebra, and
I have seen this in no other specimen.
Lumbar vertebra.--I have stated that in two cases there were
eight instead of seven lumbar vertebræ. The third lumbar ver-
tebra in one skeleton of a wild British rabbit, and in one of the
Porto Santo feral rabbits, had a hæmal spine; whilst in four skele-
tons of large lop-eared rabbits, and in the Himalayan rabbit, this
same vertebra had a well-developed hæmal spine.
26 These rabbits have run wild for a
considerable time in Sandon Park, and
in other places in Staffordshire and
Shropshire. They originated, as I have
been informed by the gamekeeper, from
variously-coloured domestic rabbits
which had been turned out. They vary
in colour; but many are symmetrically
coloured, being white with a streak
along the spine, and with the ears and
certain marks about the head of a
blackish-grey tint. They have rather
longer bodies than common rabbits.

154
CHAP. IV.
DOMESTIC RABBITS.
Pelvis.-In four wild specimens this bone was almost absolutely
identical in shape; but in several domesticated breeds shades of
differences could be distinguished. In the large lop-eared rabbits
the whole upper part of the ilium is straighter, or less splayed out-
wards, than in the wild rabbit ; and the tuberosity on the inner lip
of the anterior and upper part of the ilium is proportionally more
prominent.
Sternum.-The posterior end of the posterior sternal bone in the
wild rabbit (fig. 15, A) is thin and slightly enlarged ; in some of the
large lop-eared rabbits (B) it is much more enlarged towards the
extremity; whilst in other specimens (C) it keeps nearly of the
same breadth from end to end, but is much-thicker at the extremity.
A
B
C
A
B
Fig. 15.- Terminal bone of
Sternum, of natural size.
A. Wild Rabbit. B. Hare-
coloured, Lop-eared Rabbit.
C. Hare-coloured, Spanish
Rabbit. (N.B.-The left-
hand angle of the upper
articular extremity of B
was broken, and has been
accidentally thus repre-
sented.)
0
D
Fig. 16.- Acromion of Scapula, of natural size.
A. Wild Rabbit. B, C, D. Large, Lop-eared
Rabbits.
Scapula.—The acromion sends out a rectangular bar, ending in
an oblique knob, which latter in the wild rabbit (fig. 16, A) varies a
little in shape and size, as does the apex of the acromion in sharp-
ness, and the part just below the rectangular bar in breadth. But
the variations in these respects in the wild rabbit are very slight;
whilst in the large lop-eared rabbits they are considerable. Thus
in some specimens (B) the oblique terminal knob is developed into a
short bar, forming an obtuse angle with the rectangular bar. In
another specimen (C) these two unequal bars form nearly a straight
line. The apex of the acromion varies much in breadth and sharp-
ness, as may be seen by comparing figs. B, C, and D.

CHAP. IV. EFFECTS OF USE AND DISUSE. .
155
Limbs.In these I could detect no variation; but the bones of
the feet were too troublesome to compare with much care,
I have now described all the differences in the skele-
tons which I have observed. It is impossible not to be
struck with the high degree of variability or plasticity
of many of the bones. We see how erroneous the often-
repeated statement is, that only the crests of the bones
which give attachment to muscles vary in shape, and
that only parts of slight importance become modified
under domestication. No one will say, for instance, that
the occipital foramen, or the atlas, or the third cervical
vetebra is a part of slight importance. If the several
vertebræ of the wild and lop-eared rabbits, of which
figures have been given, had been found fossil, palæonto-
logists would have declared without hesitation that they
had belonged to distinct species.
The effects of the use and disuse of parts.-In the large lop-
eared rabbits the relative proportional lengths of the bones of the
same leg, and of the front and hind legs compared with each other,
have remained nearly the same as in the wild rabbit; but in
weight, the bones of the hind legs apparently have not increased
in due proportion with the front legs. The weight of the whole
ody in the large rabbits examined by me was from twice to twice
and a half as great as that of the wild rabbit; and the weight of
the bones of the front and hind limbs taken together (excluding the
feet, on account of the difficulty of perfectly cleaning so many small
bones) has increased in the large lop-eared rabbits in nearly the
same proportion ; consequently in due proportion to the weight of
body which they have to support. If we take the length of the
body as the standard of comparison, the limbs of the large rabbits
have not increased in length in due proportion by one inch, or by
one inch and a half. Again, if we take as the standard of compari-
son the length of the skull, which, as we have before seen, has not
increased in length in due proportion to the length of the body, the
limbs will be found to be, proportionally with those of the wild rab-
bit, from half to three-quarters of an inch too short. Hence, what-
ever standard of comparison be taken, the limb bones of the large
lop-eared rabbits have not increased in length, though they have in
weight, in full proportion to the other parts of the frame; and this,

156
CHAP. IV.
DOMESTIC RABBITS.
I presume, may be accounted for by the inactive life which during
many generations they have spent. Nor has the scapula increased
in length in due proportion to the increased length of the body.
The capacity of the osseous case of the brain is a more interesting
point, to which I was led to attend by finding, as previously stated,
that with all domesticated rabbits the length of the skull relatively
to its breadth has greatly increased in comparison with that of the
wild rabbit. If we had possessed a large number of domesticated
rabbits of nearly the same size with the wild rabbit, it would have
been a simple task to have measured and compared the capacities
of their skulls. But this is not the case; almost all the domestic
breeds have larger bodies than wild rabbits, and the lop-eared kinds
are more than double their weight. As a small animal has to
exert its senses, intellect, and instincts equally with a large animal,
we ought not by any means to expect an animal twice or thrice as
large as another to have a brain of double or treble the size.27 Now,
after weighing the bodies of four wild rabbits, and of four large but
not fattened lop-eared rabbits, I find that on an average the wild
are to the lop-eared in weight as 1 to 2:17; in average length of
body as 1 to 1.41; whilst in capacity of skull (measured as hereafter
to be described) they are only as 1 to 1'15. Hence we see that the
capacity of the skull, and consequently the size of the brain, has in-
creased but little relatively to the increased size of the body; and
this fact explains the narrowness of the skull relatively to its length
in all domestic rabbits.
In the upper half of the following table I have given the mea-
surements of the skulls of ten wild rabbits; and in the lower half
of eleven thoroughly domesticated kinds. As these rabbits differ
so greatly in size, it is necessary to have some standard by which to
compare the capacities of their skulls. I have selected the length
of skull as the best standard, for in the larger rabbits it has not, as
already stated, increased in length so much as the body; but as the
skull, like every other part, varies in length, neither it nor any
other part affords a perfect standard.
In the first column of figures the extreme length of the skull is
given in inches and decimals. I am aware that these measurements
pretend to greater accuracy than is possible; but I have found it
the least trouble to record the exact length which the compass gave.
The second and third columns give the length and weight of body,
whenever these measurements have been made. The fourth column
27 See Prof. Owen's remarks on this &c.,' read before Brit. Association, 1862;
subject in his paper on the 'Zoological with respect to Birds, see 'Proc. Zoolog.
Significance of the Brain, &c., of Man, Soc. Jan. 11th, 1848, p. 8.

CHAP. IV.
157
EFFECTS OF USE AND DISUSE.
gives the capacity of the skull by the weight of small shot with
which the skulls had been filled; but it is not pretended that these
weights are accurate within a few grains. In the fifth column the
capacity is given which the skull ought to have had by calculation,
according to the length of skull, in comparison with that of the
wild rabbit No. 1 ; in the sixth column the difference between the
actual and calculated capacities, and in the seventh the percentage
of increase or decrease, are given. For instance, as the wild rabbit
No. 5 has a shorter and lighter body than the wild rabbit No. 1, we
might have expected that its skull would have had less capacity;
the actual capacity, as expressed by the weight of shot, is 875
grains, which is 97 grains less than that of the first rabbit. But
comparing these two rabbits by the length of their skulls, we see
that in No. 1 the skull is 3.15 inches in length, and in No. 5 2.96
inches in length ; according to this ratio, the brain of No.5 ought to
have had a capacity of 913 grains of shot, which is above the actual
capacity, but only by 38 grains. Or, to put the case in another way
(as in column VII), the brain of this small rabbit, No. 5, for every
100 grains of weight is only 4 per cent. too light that is, it ought,
according to the standard rabbit No. 1, to have been 4 per cent.
heavier. I have taken the rabbit No. 1 as the standard of compari-
son because, of the skulls having a full average length, this has
the least capacity; so that it is the least favourable to the result
which I wish to show, namely, that the brain in all long-domesti-
cated rabbits has decreased in size, either actually, or relatively to
the length of the head and body, in comparison with the brain of
the wild rabbit. Had I taken the Irish rabbit, No. 3, as the stan-
dard, the following results would have been somewhat more
striking
Turning to the Table: the first four wild rabbits have skulls of
the same length, and these differ but little in capacity. The Sandon
rabbit (No. 4) is interesting, as, though now wild, it is known to be
descended from a domesticated breed, as is still shown by its pecu-
liar colouring and longer body; nevertheless the skull has recovered
its normal length and full capacity. The next three rabbits are
wild, but of small size, and they all have skulls with slightly les-
sened capacities. The three Porto Santo feral rabbits (Nos. 8 to 10)
offer a perplexing case; their bodies are greatly reduced in size, as
in a lesser degree are their skulls in length and in actual capacity,
in comparison with the skulls of wild English rabbits. But when
we compare the capacities of the skull in the three Porto Santo
rabbits, we observe a surprising difference, which does not stand in
any relation to the slight difference in the length of their skulls,
nor, as I believe, to any difference in the size of their bodies; but I

158
CHAP. IV
DOMESTIC RABBITS.
neglected to weigh separately their bodies. I can hardly suppose
that the medullary matter of the brain in these three rabbits, living
under similar conditions, can differ as much as is indicated by the
proportional difference of capacity in their skulls ; nor do I know
whether it is possible that one brain may contain considerably more
fluid than another. Hence I can throw no light on this case.
Looking to the lower half of the Table, which gives the measure-
ments of domesticated rabbits, we see that in all the capacity of the
skull is less, but in very various degrees, than might have been
anticipated according to the length of their skulls, relatively to
that of the wild rabbit No. 1. In line 22 the average measurements
of seven large lop-eared rabbits are given. Now the question
arises, has the average capacity of the skull in these seven large
rabbits increased as much as might have been expected from their
greatly increased size of body. We may endeavour to answer this
question in two ways: in the upper half of the Table we have
measurements of the skulls of six small wild rabbits (Nos. 5 to 10),
and we find that on an average the skulls are in length 18 of an
inch shorter, and in capacity 91 grains less, than the average length
and capacity of the three first wild rabbits on the list. The seven
large lop-eared rabbits, on an average, have skulls 4:11 inches in
length, and 1136 grains in capacity; so that these skulls have in-
creased in length more than five times as much as the skulls of the
six small wild rabbits have decreased in length; hence we might
have expected that the skulls of the large lop-eared rabbits would
have increased in capacity five times as much as the skulls of the
six small rabbits have decreased in capacity; and this would have
given an average increased capacity of 455 grains, whilst the real
average increase is only 155 grains. Again, the large lop-eared
rabbits have bodies of nearly the same weight and size as the com-
mon hare, but their heads are longer; consequently, if the lop-
eared rabbits had been wild, it might have been expected that their
skulls would have had nearly the same capacity as that of the skull
of the hare. But this is far from being the case ; for the average
capacity of the two hare-skulls (Nos. 23, 24) is so much larger than
the average capacity of the seven lop-eared skulls, that the latter
would have to be increased 21 per cent. to come up to the standard
of the hare 28
28 This standard is apparently consi-
derably too low, for Dr. Crisp. Proc.
Zoolog. Soc.,'1861, p. 56) gives 210 grains
as the actual weight of the brain of a
hare which weighed 7 lbs., and 125
grains as the weight of the brain of a
rabbit which weighed 3 lbs. 5 oz., that
is, the same weight as the rabbit No. 1
in my list. Now the contents of the
skull of rabbit No. 1 in shot is in my

CHAP. IV.
159
EFFECTS OF USE AND DISUSE.
I have previously remarked that, if we had possessed many do-
mestic rabbits of the same average size with the wild rabbit, it
would have been easy to compare the capacity of their skulls.
Now the Himalayan, Moscow, and Angora rabbits (Nos. 11, 12, 13
of Table) are only a little larger in body, and have skulls only a
little longer, than the wild animal, and we see that the actual ca-
pacity of their skulls is less than in the wild animal, and considera-
bly less by calculation (column 7), according to the difference in the
length of their skulls. The narrowness of the brain-case in these
three rabbits could be plainly seen and proved by external measure-
ment. The Chinchilla rabbit (No. 14) is a considerably larger ani-
mal than the wild rabbit, yet the capacity of its skull only slightly
exceeds that of the wild rabbit. The Angora rabbit, No. 13, offers
the most remarkable case; this animal in its pure white colour and
length of silky fur bears the stamp of long domesticity. It has a
considerably longer head and body than the wild rabbit, but the
actual capacity of its skull is less than that of even the little wild
Porto Santo rabbits. By the standard of the length of skull the
capacity (see column 7) is only half of what it ought to have been!
I kept this individual animal alive, and it was not unhealthy nor
idiotic. This case of the Angora rabbit so much surprised me, that
I repeated all the measurements and found them correct. I have
also compared the capacity of the skull of the Angora with that of
the wild rabbit by other standards, namely, by the length and
weight of the body, and by the weight of the limb-bones; but by
all these standards the brain appears to be much too small, though
in a less degree when the standard of the limb-bones was used ;
and this latter circumstance may probably be accounted for by the
limbs of this anciently domesticated breed having become much
reduced in weight, from its long-continued inactive life. Hence I
infer that in the Angora breed, which is said to differ from other
breeds in being quieter and more social, the capacity of the skull
has really undergone a remarkable amount of reduction.
From the several facts above given,-namely, firstly,
that the actual capacity of the skull in the Himalayan,
Moscow, and Angora breeds, is less than in the wild
rabbit, though they are in all their dimensions rather
larger animals; secondly, that the capacity of the skull
table 972 grains; and according to Dr.
Crisp's ratio of 125 to 210, the skull of
the hare ought to have contained 1682
grains of shot, instead of only in the
largest hare in my table) 1455 grains.

160
CHAP. IV.
DOMESTIC RABBITS.
I.
II.
Name of Breed.
WILD AND SEMI-WILD RABBITS
Length
Length of Body
of
from
Skull. Incisors
III.
IV.
Capacity
of
Weight
Skull
of
mea-
whole sured by
Small
Shot.
V.
VI.
Capacity,
calculated
Difference
according to
between
Length of
actual and
Skull rela-
calculated
tively to that capacities
of No.1.
of Skulls.
VII.
Showing how much per
cent. the Brain, by calcu-
lation, according to the
length of the Skull, is too
light or too heavy, rela-
tively to the Brain of the
Wild Rabbit No. 1.
to Anus. Body.
grains.
grains.
inches.
17.4
18.5
17.0
lbs. ozs. grains.
3 5 972
979
992
977
2 14 875
918
2 11 938
893
756
835
828
73
15.5
913
950
910
873
879
1910
888
38
32
28
20
123
75
60
S[2 per cent. too heavy in
comparison with No. 1.]
4 per cent. too light.
3
3
too heavy.
2
16
too light.
9
7
.
97
inches.
1. Wild rabbit, Kent
3.15
2.
Shetland Islands
3.15
3.
Ireland
3.15
4. Domestic rabbit, run wild, Sandon
3.15
5. Wild, common variety, small specimen, Kent 2.96
6. Wild, fawn-coloured variety, Scotland
3:1
7. Silver-grey, small specimen, Thetford warren
2.95
8. Feral rabbit, Porto Santo
2.83
9
2.85
10.
2.95
Average of the three Porto Santo rabbits 2.88
DOMESTIC RABBITS.
11. Himalayan
3.5
12. Moscow
3.25
13. Angora
3.5
14. Chinchilla
3.65
15. Large lop-eared
4.1
16.
4:1
17.
4.07
18.
19.
20.
4.25
21. Large hare-coloured
3.86
22. Average of above seven large lop-eared rabbits 4:11
23. Hare (L. timidus) English specimen
3.61
24.
German specimen
3.82
12
3 8
3 1
12
24
9
20.5
17.0
19.5
22.0
24.5
25.0
54
13
18
9
7 0
7 13
963
803
697
995
1065
1153
1037
1208
1232
1124
1131
1136
1080
1002
1080
1126
1265
1265
1255
1265
1326
1311
1191
1268
117
199
383
131
200
112
218
57
94
187
60
132
21
73
4:1
4.3
25.0
7 4
4
7
16
5
11
24.0
24.62
6 14
7 4
7
7
0
0
1315
1455
19

CHAP. IV.
161
EFFECTS OF USE AND DISUSE.
of the large lop-eared rabbits has not been increased in
nearly the same ratio as the capacity of the skull of
the smaller wild rabbits has been decreased; and third-
ly, that the capacity of the skull in these same large lop-
eared rabbits is very inferior to that of the hare, an ani-
mal of nearly the same size, I conclude, notwithstand-
ing the remarkable differences in capacity in the skulls
of the small P. Santo rabbits, and likewise in the large
lop-eared kinds, that in all long-domesticated rabbits the
brain has either by no means increased in due propor-
tion with the increased length of the head and increased
size of the body, or that it has actually decreased in
size, relatively to what would have occurred had these
animals lived in a state of nature. When we remember
that rabbits, from having been domesticated and closely
confined during many generations, cannot have exerted
their intellect, instincts, senses, and voluntary move-
ments, either in escaping from various dangers or in
searching for food, we may conclude that their brains
will have been feebly exercised, and consequently have
suffered in development. We thus see that the most im-
portant and complicated organ in the whole organization
is subject to the law of decrease in size from disuse.
Finally, let us sum up the more important modifica-
tions which domestic rabbits have undergone, together
with their causes as far as we can obscurely see them.
By the supply of abundant and nutritious food, together
with little exercise, and by the continued selection of
the heaviest individuals, the weight of the larger breeds
has been more than doubled. The bones of the limbs
have increased in weight (but the hind legs less than the
front legs), in due proportion with the increased weight
of body; but in length they have not increased in due
proportion, and this may have been caused by the want
of
proper exercise. With the increased size of the body
the third cervical vertebra has assumed characters proper
to the fourth cervical; and the eighth and ninth dorsal

162
CHAP. IV.
DOMESTIC RABBITS.
vertebræ have similarly assumed characters proper to
the tenth and posterior vertebræ. The skull in the
larger breeds has increased in length, but not in due
proportion with the increased length of body; the brain
has not duly increased in dimensions, or has even ac-
tually decreased, and consequently the bony case for the
brain has remained narrow, and by correlation has
affected the bones of the face and the entire length of
the skull. The skull has thus acquired its characteristic
narrowness. From unknown causes the supra-orbital pro-
cesses of the frontal bones and the free end of the malar
bones have increased in breadth; and in the larger breeds
the occipital foramen is generally much less deeply
notched than in wild rabbits. Certain parts of the sca-
pula and the terminal sternal bones have become highly
variable in shape. The ears have been increased enormous-
ly in length and breadth through continued selection;
their weight, conjoined probably with the disuse of their
muscles, has caused them to lop downwards; and this
has affected the position and form of the bony auditory
meatus; and this again, by correlation, the position in a
slight degree of almost every bone in the upper part of
the skull, and even the position of the condyles of the
lower jaw.

CHAP. V. PIGEONS: DESCRIPTION OF BREEDS. 163
CHAPTER V.
DOMESTIC PIGEONS.
ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS IN-
DIVIDUAL VARIABILITY VARIATIONS OF A REMARKABLE NA-
TURE OSTEOLOGICAL CHARACTERS: SKULL, LOWER JAW, NUM-
BER OF VERTEBRÆ- CORRELATION OF GROWTH : TONGUE WITH
BEAK, EYELIDS AND NOSTRILS WITH WATTLED SKIN — NUMBER
OF WING-FEATHERS, AND LENGTH OF WING- - COLOUR AND DOWN
WEBBED AND FEATHERED FEET - ON THE EFFECTS OF DISUSE
-LENGTH OF FEET IN CORRELATION WITH LENGTH OF BEAK -
LENGTH OF STERNUM, SCAPULA, AND FERCULA — LENGTH OF
WINGS — SUMMARY ON THE POINTS OF DIFFERENCE IN THE
SEVERAL BREEDS.
I HAVE been led to study domestic pigeons with particu-
lar care, because the evidence that all the domestic races
have descended from one known source is far clearer
than with any other anciently domesticated animal.
Secondly, because many treatises in several languages,
some of them old, have been written on the pigeon, so
that we
are enabled to trace the history of several
breeds. And lastly, because, from causes which we can
partly understand, the amount of variation has been ex-
traordinarily great. The details will often be tediously
minute; but no one who really wants to understand the
progress of change in domestic animals will regret this;
and no one who has kept pigeons and has marked the
great difference between the breeds and the trueness with
which most of them propagate their kind, will think this
care superfluous. Notwithstanding the clear evidence

164
CHAP. V.
DOMESTIC PIGEONS.
that all the breeds are the descendants of a single species,
I could not persuade myself until some years had passed
that the whole amount of difference between them had
arisen since man first domesticated the wild rock-pigeon.
I have kept alive all the most distinct breeds, which I
could procure in England or from the Continent; and have
prepared skeletons of all. I have received skins from
Persia, and a large number from India and other quarters
of the world. Since my admission into two of the Lon-
don pigeon-clubs, I have received the kindest assistance
from many of the most eminent amateurs.?
The races of the Pigeon which can be distinguished,
and which breed true, are very numerous. MM. Boitard
and Corbié º describe in detail 122 kinds; and I could add
several European kinds not known to them. In India,
judging from the skins sent me, there are many breeds
unknown here; and Sir W. Elliot informs me that a col-
lection imported by an Indian merchant into Madras
1 The Hon. C. Murray has sent me
some very valuable specimens from Per-
sia; and H.M. Consul, Mr. Keith Ab-
bott, has given me information on the
pigeons of the same country. I am
deeply indebted to Sir Walter Elliot for
an immense collection of skins from
Madras, with much information regard-
ing them. Mr. Blyth has freely com-
municated to me his stores of knowledge
on this and all other related subjects.
The Rajah Sir James Brooke sent me
specimens from Borneo, as has H.M.
Consul, Mr. Swinhoe, from Amoy in
China, and Dr. Daniell from the west
coast of Africa.
2 Mr. B. P. Brent, well known for his
various contributions to poultry litera-
ture, has aided me in every way during
several years; so has Mr. Tegetmeier,
with unwearied kindness. This latter
gentleman, who is well known for his
works on poultry, and who has largely
bred pigeons, has looked over this and
the following chapters. Mr. Bult for-
merly showed me his unrivalled collec-
tion of Pouters, and gave me specimens.
I had access to Mr. Wicking's collection,
which contained a greater assortment of
many kinds than could anywhere else
be seen; and he has always aided me
with specimens and information given
in the freest manner. Mr. Haynes and
Mr. Corker have given me specimens of
their magnificent Carriers. To Mr. Har-
rison Weir I am likewise indebted. Nor
must I by any means pass over the as-
sistance received from Mr. J. M. Eaton,
Mr. Baker, Mr. Evans, and Mr. J. Baily,
jun., of Mount-street-to the latter gen-
tlemen I have been indebted for some
valuable specimens. To all these gen-
tlemen I beg permission to return my
sincere and cordial thanks.
3 'Les Pigeons de Volière et de Co-
lombier,' Paris, 1824. During forty-five
years the sole occupation of M. Corbié
was the care of the pigeons belonging
to the Duchess of Berry.

CHAP. V.
165
DESCRIPTION OF BREEDS.
from Cairo and Constantinople included several kinds
unknown in India. I have no doubt that there exist
considerably above 150 kinds which breed true and have
been separately named. But of these the far greater
number differ from each other only in unimportant cha-
racters. Such differences will be here entirely passed
over, and I shall confine myself to the more important
points of structure. That many important differences
exist we shall presently see. I have looked through the
magnificent collection of the Columbidæ in the British
Museum, and, with the exception of a few forms (such as
the Didunculus, Calænas, Goura, &c.), I do not hesitate
to affirm that some domestic races of the rock-pigeon
differ fully as much from each other in external charac-
ters as do the most distinct natural genera. We may
look in vain through the 288 known species * for a beak
so small and conical as that of the short-faced tumbler;
for one so broad and short as that of the barb; for one
so long, straight, and narrow, with its enormous wattles,
as that of the English carrier; for an expanded upraised
tail like that of the fantail; or for an esophagus like that
of the pouter. I do not for a moment pretend that the
domestic races differ from each other in their whole or-
ganisation as much as the more distinct natural genera.
I refer only to external characters, on which, however,
it must be confessed that most genera of birds have been
founded. When, in a future chapter, we discuss the prin-
ciple of selection as followed by man, we shall clearly
see why the differences between the domestic races are
almost always confined to external, or at least to exter-
nally visible, characters.
Owing to the amount and gradations of difference be-
tween the several breeds, I have found it indispensable in
the following classification to rank them under Groups,
4 Coup d'oeil sur l'Ordre des Pi-
geons,' par Prince C. L. Bonaparte, Pa-
ris, 1855. This author makes 288 spe-
cies, ranked under 85 genera,

166
CHAP. V.
DOMESTIC PIGEONS. .
Races, and Sub-races; to which varieties and sub-varie-
ties, all strictly inheriting their proper characters, must
often be added. Even with the individuals of the same
sub-variety, when long kept by different fanciers, differ-
ent strains can sometimes be recognised. There can be
no doubt that, if well-characterized forms of the several
Races had been found wild, all would have been ranked
as distinct species, and several of them would certainly
have been placed by ornithologists in distinct genera.
A
good classification of the various domestic breeds is ex-
tremely difficult, owing to the manner in which many of
the forms graduate into each other; but it is curious how
exactly the same difficulties are encountered, and the
same rules have to be followed, as in the classification of
any natural but difficult group of organic beings. An
“artificial classification” might be followed which would
present fewer difficulties than a “natural classification;"
but then it would interrupt many plain affinities. Ex-
treme forms can readily be defined ; but intermediate
and troublesome forms often destroy our definitions.
Forms which may be called “aberrant” must sometimes
be included within groups to which they do not accurate-
ly belong. Characters of all kinds must be used; but as
with birds in a state of nature, those afforded by the beak
are the best and most readily appreciated. It is not pos-
sible to weigh the importance of all the characters which
have to be used so as to make the groups and sub-groups
of equal value. Lastly, a group may contain only one
race, and another and less distinctly defined group may
contain several races and sub-races, and in this case it is
difficult, as in the classification of natural species, to avoid
placing too high a value on characters which are common
to a large number of forms.
In my measurements I have never trusted to the eye;
and when speaking of a part being large or small, I
always refer to the wild rock-pigeon (Columba livia)

CHAP. V.
167
DESCRIPTION OF BREEDS.
as the standard of comparison. The measurements are
given in decimals of an inch."
I will now give a brief description of all the principal
breeds. The following diagram may aid the reader in
learning their names and seeing their affinities. The
rock-pigeon, or Columba livia (including under this name
two or three closely allied sub-species or geographical
races, hereafter to be described), may be confidently
viewed, as we shall see in the next chapter, as the com-
mon parent-form. The names in italics on the right-hand
side of the table show us the most distinct breeds, or those
which have undergone the greatest amount of modifica-
tion. The lengths of the dotted lines rudely represent
the degree of distinctness of each breed from the parent-
stock, and the names placed under each other in the col-
umns show the more or less closely connecting links. The
distances of the dotted lines from each other approximate-
ly represent the amount of difference between the seve-
ral breeds.
23
5 As I so often refer to the size of the measurements of two wild birds, kindly
C. livia, or rock-pigeon, it may be con- sent me by Dr. Edmondstone from the
venient to give the mean between the Shetland Islands :-
Inches.
Length from feathered base of beak to end of tail
14.25
to oil-gland
9:5
from tip of beak to end of tail
15.02
of tail feathers
4.62
from tip to tip of wing
26.75
of folded wing...
9.25
Beak.-Length from tip of beak to feathered base
-77
Thickness, measured vertically at further end of nostrils
-23
Breadth, measured at same place ..
-16
Feet.-Length from end of middle toe (without claw) to distal end of tibia 2.77
Length from end of middle toe to end of hind toe (without claws).. .. 2:02
Weight 1474 ounces.

168
CHAP. V
DOMESTIC PIGEONS.
Fig. 17.- The Rock-pigeon, or Columba livia..
Pigeons.
The parent-form of all domesticated
6 This drawing was made from a dead
bird. The six following figures were
drawn with great care by Mr. Luke Wells
from living birds selected by Mr. Teget-
meier. It may be confidently asserted
that the characters of the six breeds
which have been figured are not in the
least exaggerated.

CHAP. V.
169
DESCRIPTION OF BREEDS.
COLUMBA LIVIA OR ROCK-PIGEON.
GROUP I.
GROUP II.
GROUP III.
GROUP IV.
1.
2.
3.
4.
5.
6.
7.
SUB-
GROUPS.
8.
10.
11.
9. SUB-
GROUPS.
.........
Kali-Par
Persian
Tumbler
..Murassa
Lotan
Tumbler
Bussorah
Common
Tumbler
Bagadotten
Scanderoon
Tronfo
German P.
Lille P.
Dragon
Dutch P.
Java
Fantail
Pigeon
Cygne
Turbit.
Nun.
Laugher.
Spot.
Trumpeter.
Swallow.
English Frill-back.
Dove-cot pigeon.
English
Pouter.
Runt.
English
Carrier
Jacobin.
Barb. Fantail. African Short Indian
Orol.
faced
Frill-
Tumbler. back.
00

170
CAAP. V.
DOMESTIC PIGEONS.
en
Fig. 18.-English Pouter.
GROUP I.
This
group includes a single race, that of the Pouters.
If the most strongly marked sub-race be taken, namely,
the Improved English Pouter, this is perhaps the most
distinct of all domesticated pigeons.

CHAP. V.
171
DESCRIPTION OF BREEDS.
RACE I. -POUTER PIGEONS. (Kropf-tauben, German,
Grosses-gorges, or boulans, French.)
Esophagus of great size, barely separated from the
crop, often inflated. Body and legs elongated. Beak of
moderate dimensions.
Sub-race I.-The improved English Pouter, when its crop is fully
inflated, presents a truly astonishing appearance. The habit of
slightly inflating the crop is common to all domestic pigeons, but is
carried to an extreme in the Pouter. The crop does not differ, ex-
cept in size, from that of other pigeons; but is less plainly separated
by an oblique construction from the cesophagus. The diameter of
the upper part of the oesophagus is immense, even close up to the
head. The beak in one bird which I possessed was almost com-
pletely buried when the oesophagus was fully expanded. The males,
especially when excited, pout more than the females, and they glory
in exercising this power. If a bird will not, to use the technical ex-
pression, "play,” the fancier, as I have witnessed, by taking the beak
into his mouth, blows him up like a balloon; and the bird, then
puffed up with wind and pride, struts about, retaining his magnifi-
cent size as long as he can. Pouters often take flight with their crops
inflated ; and after one of my birds had swallowed a good meal of
peas and water, as he flew up in order to disgorge them and thus
feed his nearly fledged young, I have heard the peas rattling in his
inflated crop as if in a bladder. When flying they often strike the
backs of their wings together, and thus make a clapping noise.
Pouters stand remarkably upright, and their bodies are thin and
elongated. In connexion with this form of body, the ribs are gene-
rally broader and the vertebræ more numerous than in other breeds.
From their manner of standing their legs appear longer than they
really are, though in proportion with those of C. livia, the legs and
feet are actually longer. The wings appear much elongated, but by
measurement, in relation to the length of body, this is not the case.
The beak likewise appears longer, but it is in fact a little shorter
(about .03 of an inch), proportionally with the size of the body, and
relatively to the beak of the rock-pigeon. The Pouter, though not
bulky, is a large bird ; I measured one which was 341 inches from
tip to tip of wing, and 19 inches from tip of beak to end of tail. In
a wild rock-pigeon from the Shetland Islands the same measurements
gave only 283 and 141. There are many sub-varieties of the Pouter
of different colours, but these I pass over.

172
CHAP. V.
DOMESTIC PIGEONS.
Sub-race II. Dutch Pouter. This seems to be the parent-form
of our improved English Pouters. I kept a pair, but I suspect that
they were not pure birds. They are smaller than English Pouters,
and less well developed in all their characters. Neumeister" says
that the wings are crossed over the tail, and do not reach to its ex-
tremity.
Sub-race III. The Lille Pouter.—I know this breed only from
description. It approaches in general form the Dutch Pouter, but
the inflated oesophagus assumes a spherical form, as if the pigeon
had swallowed a large orange, which had stuck close under the
beak. This inflated ball is represented as rising to a level with the
crown of the head. The middle toe alone is feathered. A variety
of this sub-race, called the claquant, is described by MM. Boitard
and Corbié ; it pouts but little, and is characterised by the habit of
violently hitting its wings together over its back,-a habit which
the English Pouter has in a slight degree.
Sub-race IV. Common German Pouter.--I know this bird only
from the figures and description given by the accurate Neumeister,
one of the few writers on pigeons who, as I have found, may be al-
ways trusted. This sub-race seems considerably different. The up-
per part of the oesophagus is much less distended. The bird stands
less upright. The feet are not feathered, and the legs and beak are
shorter. In these respects there is an approach in form to the com-
mon rock-pigeon. The tail-feathers are very long, yet the tips of
the closed wings extend beyond the end of the tail; and the length
of the wings, from tip to tip, and of the body, is greater than in the
English Pouter.
GROUP II.
This group includes three Races, namely, Carriers,
Runts, and Barbs, which are manifestly allied to each
other Indeed, certain carriers and runts pass into each
other by such insensible gradations that an arbitrary line
has to be drawn between them. Carriers also graduate
through foreign breeds into the rock-pigeon. Yet, if well-
characterised Carriers and Barbs (see figs. 19 and 20) had
existed as wild species, no ornithologist would have placed
them in the same genus with each other or with the rock-
7. Das Ganze der Taubenzucht:' Wei-
mar, 1887, pl. 11 and 12.
8 Boitard and Corbié, 'Les Pigeons,'
&c., 177, pl. 6.

CHAP. V.
173
DESCRIPTION OF BREEDS.
pigeon. This group may, as a general rule, be recognised
by the beak being long, with the skin over the nostrils
swollen and often carunculated or wattled, and with that
round the eyes bare and likewise carunculated. The
mouth is very wide, and the feet are large. Nevertheless
the Barb, which must be classed in this same group, has a
very short beak, and some runts have very little bare skin
round their eyes.
RACE II. —CARRIERS. (Türkische Taube: Pigeons Turcs :
Dragons.)
Beak elongated, narrow, pointed ; eyes surrounded by
much naked, generally carunculated skin; neck and body
elongated.
Sub-race I. The English Carrier.—This is a fine bird, of large
size, close feathered, generally dark-coloured, with an elongated
neck. The beak is attenuated and of wonderful length: in one
specimen it was 14 inch in length from the feathered base to the
tip; therefore nearly twice as long as that of the rock-pigeon, which
measured only 77. Whenever I compare proportionally any part in
the carrier and rock-pigeon, I take the length of the body from the
base of the beak to the end of the tail as the standard of compari-
son; and according to this standard, the beak in one Carrier was
nearly half an inch longer than in the rock-pigeon. The upper
mandible is often slightly arched. The tongue is very long. The
development of the carunculated skin or wattle round the eyes, over
the nostrils, and on the lower mandible is prodigious. The eyelids,
measured longitudinally, were in some specimens exactly twice as
long as in the rock-pigeon. The external orifice or furrow of the
nostrils was also twice as long. The open mouth in its widest part
was in one case 75 of an inch in width, whereas in the rock-pigeon
it is only about 4 of an inch. This great width of mouth is shown
in the skeleton by the reflexed edges of the ramus of the lower jaw.
The head is flat on the summit and narrow between the orbits. The
feet are large and coarse; the length, as measured from end of hind
toe to end of middle toe (without the claws), was in two specimens
2.6 inches; and this, proportionally with the rock-pigeon, is an ex-
cess of nearly a quarter of an inch. One very fine Carrier measured
314 inches from tip to tip of wing. Birds of this sub-race are too
valuable to be flown as carriers.

174
CHAP. V.
DOMESTIC PIGEONS.
Sub-race II. Dragons ; Persian Carriers.—The English Dragon
differs from the improved English Carrier in being smaller in all
its dimensions, and in having less wattle round the eyes and over
Fig. 19.- English Carrier.
the nostrils, and none on the lower mandible. Sir W. Elliot sent
me from Madras a Bagdad Carrier (sometimes called khandési), the
name of which shows its Persian origin; it would be considered

CHAP. V.
175
DESCRIPTION OF BREEDS.
here a very poor Dragon; the body was of the size of the rock-
pigeon, with the beak a little longer, namely, 1 inch from the tip to
the feathered base. The skin round the eyes was only slightly
wattled, whilst that over the nostrils was fairly wattled. The Hon.
C. Murray, also, sent me two Carriers direct from Persia ; these had
nearly the same character as the Madras bird, being about as large
as the rock-pigeon, but the beak in one specimen was as much as
1.15 in length; the skin over the nostrils was only moderately, and
that round the eyes scarcely at all wattled.
Sub-race III. Bagadotten-Tauben of Neumeister (Pavdotten or
Hocker-Tauben).--I owe to the kindness of Mr. Baily, jun., a dead
specimen of this singular breed imported from Germany. It is
certainly allied to the Runts; nevertheless, from its close affinity
with Carriers, it will be convenient here to describe it. The beak
is long, and is hooked or bowed downwards in a highly remarkable
manner, as will be seen in the woodcut to be hereafter given when
I treat of the skeleton. The eyes are surrounded by a wide space
of bright red skin, which, as well as that over the nostrils, is mode-
rately wattled. The breast-bone is remarkably protuberant, being
abruptly bowed outwards. The feet and tarsi are of great length,
larger than in first-rate English Carriers. The whole bird is of
large size, but in proportion to the size of the body the feathers of
the wing and tail are short; a wild-rock pigeon of considerably less
size, had tail-feathers 4:6 inches in length, whereas in the large
Bagadotten these feathers were scarcely over 41 inches in length.
Riedel º remarks that it is a very silent bird.
Sub-race IV. Bussorah Carrier.-Two specimens were sent me
by Sir W. Elliot from Madras, one in spirits and the other skinned.
The name shows its Persian origin. It is much valued in India,
and is considered as a distinct breed from the Bagdad Carrier,
which forms my second sub-race. At first I suspected that these
two sub-races might have been recently formed by crosses with
other breeds, though the estimation in which they are held renders
this improbable; but in a Persian treatise, 10 believed to have been
written about 100 years ago, the Bagdad and Bussorah breeds are
described as distinct. The Bussorah Carrier is of about the same
size with the wild roek-pigeon. The shape of the beak, with some
little carunculated skin over the nostrils,-the much elongated
eyelids,—the broad mouth measured internally,—the narrow head, -
the feet proportionally a little longer than in the rock-pigeon,--and
9 Die Taubenzucht,' Ulm, 1924, s. 42.
10 This treatise was written by Sayzid
Mohammed Musari, who died in 1770: I
owe to the great kindness of Sir W. Elliot
a translation of this curious treatise.

176
CHAP. V.
DOMESTIC PIGEONS.
the general appearance, all show that this bird is an undoubted
Carrier; yet in one specimen the beak was of exactly the same
length as in the rock-pigeon. In the other specimen the beak (as
well as the opening of the nostrils) was only a very little longer,
viz. by .08 of an inch. Although there was a considerable space of
bare and slightly carunculated skin round the eyes, that over the
nostrils was only in a slight degree rugose. Sir W. Elliot informs
me that in the living bird the eye seems remarkably large and
prominent, and the same fact is noticed in the Persian treatise; but
the bony orbit is barely larger than that in the rock-pigeon.
Amongst the several breeds sent to me from Madras by Sir W.
Elliot there is a pair of the Kala Par, black birds with the beak
slightly elongated, with the skin over the nostrils rather full, and
with a little naked skin round the eyes. This breed seems more
closely allied to the Carrier than to any other breed, being nearly
intermediate between the Bussorah Carrier and the rock-pigeon.
The names applied in different parts of Europe and in India to
the several kinds of Carriers all point to Persia or the surrounding
countries as the source of this Race. And it deserves especial notice
that, even if we neglect the Kala Par as of doubtful origin, we get
a series broken by very small steps, from the rock-pigeon, through
the Bussorah, which sometimes has a beak not at all longer than
that of the rock-pigeon and with the naked skin round the eyes and
over the nostrils very slightly swollen and carunculated, through
the Bagdad sub-race and Dragons, to our improved English Carriers,
which present so marvellous a difference from the rock-pigeon or
Columba livia.
RACE III.—RUNTS. (Scanderoons: Die Florentiner-Taube
and Hinkel-Taube of Neumeister: Pigeon Bagadais,
Pigeon Romain.)
Beak long, massive ; body of great size.
Inextricable confusion reigns in the classification, affinities, and
naming of Runts. Several characters which are generally pretty
constant in other pigeons, such as the length of the wings, tail,
legs, and neck, and the amount of naked skin round the eyes, are
excessively variable in Runts. When the naked skin over the
nostrils and round the eyes is considerably developed and wattled,
and when the size of body is not very great, Runts graduate in so
insensible a manner into Carriers, that the distinction is quite arbi-
trary. This fact is likewise shown by the names given to them in
different parts of Europe. Nevertheless, taking the most distinct

CHAP. V.
177
DESCRIPTION OF BREEDS.
forms, at least five sub-races (some of them including well-marked
varieties) can be distinguished, which differ in such important points
of structure, that they would be considered as good species in a state
of nature.
Sub-race I. Scanderoon of English writers (Die Florentiner and
Hinkel-Taube of Neumeister). -Birds of this sub-race, of which I
kept one alive and have since seen two others, differ from the Baga-
dotten of Neumeister only in not having the beak nearly so much
curved downwards, and in the naked skin round the eyes and over
the nostrils being hardly at all wattled. Nevertheless I have felt
myself compelled to place the Bagadotten in Race II., or that of the
Carriers, and the present bird in Race III., or that of the Runts.
The Scanderoon has a very short, narrow, and elevated tail ; wings
extremely short, so that the first primary feathers were not longer
than those of a small tumbler pigeon! Neck long, much bowed ;
breast-bone prominent. Beak long, being 1.15 inch from tip to
feathered base; vertically thick; slightly curved downwards. The
skin over the nostrils swollen, not wattled; naked skin round the
eyes, broad, slightly carunculated. Legs long, feet very large. .
Skin of neck bright red, often showing a naked medial line, with a
naked red patch at the distant end of the radius of the wing. My
bird, as measured from the base of the beak to the root of the tail,
was fully 2 inches longer than the rock-pigeon ; yet the tail itself
was only four inches in length, whereas in the rock-pigeon, which
is a much smaller bird, the tail is 45 inches in length.
The Hinkel or Florentiner-Taube of Neumeister (Table XIII., fig.
1) agrees with the above description in all the specified characters
(for the beak is not mentioned), except that Neumeister expressly
says that the neck is short, whereas in my Scanderoon it was re-
markably long and bowed ; so that the Hinkel forms a well-marked
variety
Sub-race II. Pigeon Cygne and Pigeon Bagadais of Boitard and
Corbié (Scanderoon of French writers).—I kept two of these birds
alive, imported from France. They differed from the first sub-race
or true Scanderoon in the much greater length of the wing and tail,
in the beak not being so long, and in the skin about the head being
more carunculated. The skin of the neck is red; but the naked
patches on the wings are absent. One of my birds measured 381
inches from tip to tip of wing. By taking the length of the body as
the standard of comparison, the two wings were no less than 5
inches longer than those of the rock-pigeon! The tail was 61 inches
in length, and therefore 21 inches longer than that of the Scande-
roon,-a bird of nearly the same size. The beak is longer, thicker,
and broader than in the rock-pigeon, proportionally with the size

178
CHAP. V.
DOMESTIC PIGEONS.
of body. The eyelids, nostrils, and internal gape of mouth are all
proportionally very large, as in Carriers. The foot, from the end of
the middle to end of hind toe, was actually 2.85 inches in length,
which is an excess of 32 of an inch over the foot of the rock-pigeon,
relatively to the size of the two birds.
Sub-race III. Spanish and Roman Runts.—I am not sure that I
am right in placing these Runts in a distinct sub-race; yet, if we
take well-characterized birds, there can be no doubt of the propriety
of the separation. They are heavy, massive birds, with shorter
necks, legs, and beaks than in the foregoing races.
The skin over
the nostrils is swollen, but not carunculated; the naked skin round
the eyes is not very wide, and only slightly carunculated ; and I
have seen a fine so-called Spanish Runt with hardly any naked skin
round the eyes. Of the two varieties to be seen in England, one,
which is the rarer, has very long wings and tail, and agrees pretty
chosely with the last sub-race; the other, with shorter wings and
tail, is apparently the Pigeon Romain ordinaire of Boitard and
Corbie. These Runts are apt to tremble like Fantails. They are
bad flyers. A few years ago Mr. Gulliver 11 exhibited a Runt which
weighed 1 lb. 14 oz.; and, as I am informed by Mr. Tegetmeier,
two Runts from the south of France were lately exhibited at the
Crystal Palace, each of which weighed 2 lbs. 24 oz. A very fine
rock-pigeon from the Shetland Islands weighed only 141 oz.
Sub-race IV. Tronfo of Aldrovandi (Leghorn Runt ?).—In Aldro-
vandi's work published in 1600 there is a coarse woodcut of a great
Italian pigeon, with an elevated tail, short legs, massive body, and
with the beak short and thick. I had imagined that this latter
character, so abnormal in the group, was merely a false representa-
tion from bad drawing; but Moore, in his work published in 1735,
says that he possessed a Leghorn Runt of which "the beak was
very short for so large a bird." In other respects Moore's bird re-
sembled the first sub-race of Scanderoon, for it had a long bowed
neck, long legs, short beak, and elevated tail, and not much wattle
about the head. So that Aldrovandi's and Moore's birds must have
formed distinct varieties, both of which seem to be now extinct in
Europe. Sir W. Elliot, however, informs me that he has seen in
Madras a short-beaked Runt imported from Cairo.
Sub-race V. Murassa (adorned Pigeon) of Madras.-Skins of these
handsome chequered birds were sent me from Madras by Sir W.
Elliot. They are rather larger than the largest rock-pigeon, with
longer and more massive beaks. The skin over the nostrils is
11 Poultry Chronicle,' vol. ii. p. 573.

CHAP. V.
179
DESCRIPTION OF BREEDS.
rather full and very slightly carunculated, and they have some
naked skin round the eyes : feet large. This breed is intermediate
between the rock-pigeon and a very poor variety of Runt or Car-
rier.
From these several descriptions we see that with Runts, as with
Carriers, we have a fine gradation from the rock-pigeon (with the
Tronfo diverging as a distinct branch) to our largest and most mas-
sive Runts. But the chain of affinities, and many points of resem-
blance, between Runts and Carriers, make me believe that these
two races have not descended by independent lines from the rock-
pigeon, but from some common parent, as represented in the Table,
which had already acquired a moderately long beak, with slightly
swollen skin over the nostrils, and with some slightly carunculated
naked skin round the eyes.
RACE IV.—BARBS. (Indische-Taube: Pigeons Polonais.)
Beak short, broad, deep; naked skin round, the eyes,
broad and carunculated ; skin over nostrils slightly
swollen.
Misled by the extraordinary shortness and form of the beak, I did
not at first perceive the near affinity of this Race to that of Carriers
until the fact was pointed out to me by Mr. Brent. Subsequently,
after examining the Bussorah Carrier, I saw that no very great
amount of modification would be requisite to convert it into a Barb.
This view of the affinity of Barbs to Carriers is supported by the
analogical difference between the short and long-beaked Runts;
and still more strongly by the fact that young Barbs and Dragons,
within 24 hours after being hatched, resemble each other more
closely than do young pigeons of other and equally distinct breeds.
At this early age, the length of beak, the swollen skin over the ra-
ther open nostrils, the gape of the mouth, and the size of the feet,
are the same in both; although these parts afterwards become
widely different. We thus see that embryology (as the comparison
of very young animals may perhaps be called) comes into play in
the classification of domestic varieties, as with species in a state of
nature.
Fanciers, with some truth, compare the head and beak of the
Barb to that of a bullfinch. The Barb, if found in a state of nature,
would certainly have been placed in a new genus formed for its re-
ception. The body is a little larger than that of the rock-pigeon,
but the beak is more than 2 of an inch shorter ; although shorter,

180
CHAP. V.
DOMESTIC PIGEONS.
FTY. 20.-English Barb.
it is both vertically and horizontally thicker. From the outward
flexure of the rami of the lower jaw, the mouth internally is very

CHAP. V.
181
DESCRIPTION OF BREEDS.
broad, in the proportion of .6 to 4 to that of the rock-pigeon. The
whole head is broad. The skin over the nostrils is swollen, but not
carunculated, except slightly in first-rate birds when old; whilst the
naked skin round the eye is broad and much carunculated. It is
sometimes so much developed, that a bird belonging to Mr. Harri-
son Weir could hardly see to pick up food from the ground. The
eyelids in one specimen were nearly twice as long as those of the
rock-pigeon. The feet are coarse and strong, but proportionally
rather shorter than in the rock-pigeon. The plumage is generally
dark and uniform. Barbs, in short, may be called short-beaked Car-
riers, bearing the same relation to Carriers that the Tronfo of Aldro-
vandi does to the common Runt.
GROUP III.
This group is artificial, and includes a heterogeneous
collection of distinct forms. It may be defined by the
beak in well-characterised specimens of the several races,
being shorter than in the rock-pigeon, and by the skin
round the eyes not being much developed.
RACE V.-FANTAILS.
Sub-race I. European Fantails (Pfauen-Taube ; Trembleurs). Tail
expanded, directed upwards, formed of many feathers; oil-gland
aborted; body and beak rather short.
The normal number of tail-feathers in the genus Columba is 12;
but Fantails have from only 12 (as has been asserted) up to, ac-
cording to MM. Boitard and Corbié, 42. I have counted in one of
my own birds 33, and at Calcutta Mr. Blyth 12 has counted in an im-
perfect tail 34 feathers. In Madras, as I am informed by Sir W.
Elliot, 32 is the standard number; but in England number is much
less valued than the position and expansion of the tail. The feath-
ers are arranged in an irregular double row; their permanent ex-
pansion, like a fan, and their upward direction, are more remarkable
characters than their increased number. The tail is capable of the
same movements as in other pigeons, and can be depressed so as to
sweep the ground. It arises from a more expanded basis than in
other pigeons; and in three skeletons there were one or two extra
coccygeal vertebræ. I have examined many specimens of various
12 Annals and Mag. of Nat. History,' vol. xix., 1847, p. 105.

182
CHAP. V.
DOMESTIC PIGEONS.
I'ig. 21.-English Fantail.
colours from different countries, and there was no trace of the oil-
gland; this is a curious case of abortion.13 The neck is thin and
13 This gland occurs in most birds;
but Nitzsch (in his 'Pterylographie,'
1810, p. 55) states that it is absent in two
species of Columba, in several species of

CHAP. V.
183
DESCRIPTION OF BREEDS.
bowed backwards. The breast is broad and protuberant. The feet
are small. The carriage of the bird is very different from that of
other pigeons; in good birds the head touches the tail-feathers,
which consequently often become crumpled. They habitually trem-
ble much; and their necks have an extraordinary, apparently con-
vulsive, backward and forward movement. Good birds walk in a
singular manner, as if their small feet were stiff. Owing to their
large tails, they fly badly on a windy day. The dark-coloured vari-
eties are generally larger than white Fantails.
Although between the best and common Fantails, now existing
in England, there is a vast difference in the position and size of the
tail, in the carriage of the head and neck, in the convulsive move-
ments of the neck, in the manner of walking, and in the breadth
of the breast, the differences so graduate away, that it is impossi-
ble to make more than one sub-race. Moore, however, an excellent
old authority, 14 says, that in 1735 there were two sorts of broad-
tailed shakers (i.e. fantails), "one having a neck much longer and
more slender than the other;" and I am informed by Mr. B. P. Brent
that there is an existing German Fantail with a thicker and shorter
beak.
Sub-race II. Java Fantail.-Mr. Swinhoe sent me from Amoy, in
China, the skin of a Fantail belonging to a breed known to have
been imported from Java. It was coloured in a peculiar manner,
unlike any European Fantail, and, for a fantail, had a remarkably
short beak. Athough a good bird of the kind, it had only 14 tail-
feathers; but Mr. Swinhoe has counted in other birds of this breed
from 18 to 24 tail-feathers. From a rough sketch sent to me, it is
evident that the tail is not so much expanded or so much upraised
as in even second-rate European Fantails. The bird shakes its neck
like our Fantails. It had a well-developed oil-gland. Fantails
were known in India, as we shall hereafter see, before the year
1600 ; and we may suspect that in the Java Fantail we see the
breed in its earlier and less improved condition.
Psittacus, in some species of Otis, and in
most or all birds of the Ostrich family.
It can hardly be an accidental coinci-
dence that the two species of Columba,
which are destitute of an oil-gland, have
an unusual number of tail-feathers,
namely 16, and in this respect resemble
Fantails.
14 See the two excellent editions pub-
lished by Mr.J. M. Eaton in 1852 and
1859, entitled 'A Treatise on Fancy
Pigeons.

184
CHAP. V.
DOMESTIC PIGEONS.
RACE VI.-TURBIT AND OWL. (Möven-Taube: Pigeons
à cravate.)
Feathers divergent along the front of the neck and
breast; beak very short, vertically rather thick ; oesopha
gus somewhat enlarged.
Turbits and Owls differ from each other slightly in the shape of
the head, in the former having a crest, and in the curvature of the
beak, but they may be here conveniently grouped together. These
pretty birds, some of which are very small, can be recognised at
once by the feathers irregularly diverging, like a frill, along the
front of the neck, in the same manner, but in a less degree, as along
the back of the neck in the Jacobin. This bird has the remarkable
habit of continually and momentarily inflating the upper part of
the oesophagus, which causes a movement in the frill. When the
oesophagus of a dead bird was inflated, it was seen to be larger
than in other breeds, and not so distinctly separated from the crop.
The Pouter inflates both its true crop and oesophagus; the Turbit
inflates in a much less degree the oesophagus alone. The beak of
the Turbit is very short, being 28 of an inch shorter than that of
the rock-pigeon, proportionally with the size of their bodies; and in
some owls brought by Mr. E. Vernon Harcourt from Tunis, it was
even shorter. The beak is vertically thicker, and perhaps a little
broader, in proportion to that of the rock-pigeon.
RACE VII.-TUMBLERS. (Tümmler, or Burzel-Tauben:
Culbutants.)
During flight, tumble backwards ; body generally
small ; beak generally short, sometimes excessively short
and conical.
This Race may be divided into four sub-races, namely, Persian,
Lotan, Common, and Short-faced Tumblers. These sub-races in-
clude many varieties which breed true. I have examined eight
skeletons of various kinds of Tumblers: excepting in one imperfect
and doubtful specimen, the ribs are only seven in number, whereas
the rock-pigeon has eight ribs.
Sub-race I. Persian Tumblers.--I have received a pair direct
from Persia, from the Hon. C. Murray. They were rather smaller
birds than the wild rock-pigeon, being about the size of the common

CHAP. V.
185
DESCRIPTION OF BREEDS.
dovecot-pigeon, white and mottled, slightly feathered on the feet,
with the beak just perceptibly shorter than in the rock-pigeon. H.M.
Fig. 22.--African Owl.
1. fiue
Consul, Mr. Keith Abbott, informs me that the difference in the
length of beak is so slight, that only practised Persian fanciers can

186
CHAP. V.
DOMESTIC PIGEONS.
distinguish these Tumblers from the common pigeon of the country.
He informs me that they fly in flocks high up in the air and tumble
well. Some of them occasionally appear to become giddy and tum-
ble to the ground, in which respect they resemble some of our
Tumblers.
Sub-race II. Lotan, or Lowtun : Indian Ground Tumblers.
These birds present one of the most remarkable inherited habits or
instincts which have ever been recorded. The specimens sent to me
from Madras by Sir. W. Elliot are white, slightly feathered on the
feet, with the feathers on the head reversed ; and they are rather
smaller than the rock or dovecot pigeon. The beak is proportion-
ally only slightly shorter and rather thinner than in the rock-pigeon.
These birds when gently shaken and placed on the ground immedi-
ately begin tumbling head over heels, and they continue thus to
tumble until taken up and soothed,—the ceremony being generally
to blow in their faces, as in recovering a person from a state of
hypnotism or mesmerism. It is asserted that they will continue to
roll over till they die, if not taken up. There is abundant evidence
with respect to these remarkable peculiarities; but what makes the
case the more worthy of attention is, that the habit has been strictly
inherited since before the year 1600, for the breed is distinctly de-
scribed in the 'Ayeen Akbery.'15 Mr. Evans kept a pair in London,
imported by Captain Vigne; and he assures me that he has seen
them tumble in the air, as well as in the manner above described
on the ground. Sir W. Elliot, however, writes to me from Madras,
that he is informed that they tumble exclusively on the ground,
or at a very small height above it. He also mentions another sub-
variety, called the Kalmi Lotan, which begins to roll over if only
touched on the neck with a rod or wand.
Sub-race III. Common English Tumblers.—These birds have ex-
actly the same habits as the Persian Tumbler, but tumble better.
The English bird is rather smaller than the Persian, and the beak
is plainly shorter. Compared with the rock-pigeon, and proportion-
ally with the size of body, the beak is from 15 to nearly 2 of an
inch shorter, but it is not thinner. There are several varieties of the
common Tumbler, namely, Baldheads, Beards, and Dutch Rollers.
I have kept the latter alive; they have differently shaped heads,
15 English translation, by F. Gladwin,
4th edition, vol. i. The habit of the Lo-
tan is also described in the Persian trea-
tise before alluded to, published about
100 years ago : at this date the Lotans
were generally white and crested as at
present. Mr. Blyth describes these birds
in 'Annals and Mag. of Nat. Hist.,' vol.
xiv., 1847, p. 104: he says that they
"may be seen at any of the Calcutta
bird-dealers."

CHAP. V.
187
DESCRIPTION OF BREEDS.
longer necks, and are feather-footed. They tumble to an extraor-
dinary degree; as Mr. Brent remarks, 16 "Every few seconds over they
"go; one, two, or three summersaults at a time. Here and there a bird
“ gives a very quick and rapid spin, revolving like a wheel, though
they sometimes lose their balance, and make a rather ungraceful
"fall, in which they occasionally hurt themselves by striking some
"object." From Madras I have received several specimens of the
common Tumbler of India, differing slightly from each other in the
length of their beaks. Mr. Brent sent me a dead specimen of a
"House-tumbler," 17 which is a Scotch variety, not differing in gen-
eral appearance and form of beak from the common Tumbler. Mr.
Brent states that these birds generally begin to tumble" almost as soon
as they can well fly; at three months old they tumble well, but still
“fly strong; at five or six months they tumble excessively; and in
“the second year they mostly give up flying, on account of their
"tumbling so much and so close to the ground. Some fly round with
“the flock, throwing a clean summersault every few yards, till they
are obliged to settle from giddiness and exhaustion. These are
"called Air Tumblers, and they commonly throw from twenty to
“thirty summersaults in a minute, each clear and clean. I have one
red cock that I have on two or three occasions timed by my watch
"and counted forty summersaults in the minute. Others tumble
differently. At first they throw a single summersault, then it is
“double, till it becomes a continuous roll, which puts an end to fly-
"ing, for if they fly a few yards over they go, and roll till they
reach the ground. Thus I had one kill herself, and another broke
“his leg. Many of them turn over only a few inches from the ground,
"and will tumble two or three times in flying across their loft. These
"are called House-Tumblers, from tumbling in the house. The act
“of tumbling seems to be one over which they have no control, an
"involuntary movement which they seem to try to prevent. I have
"seen a bird sometimes in his struggles fly a yard or two straight
upwards, the impulse forcing him backwards while he struggles
"to go forwards. If suddenly startled, or in a strange place, they
“seem less able to fly than if quiet in their accustomed loft.” These
House-tumblers differ from the Lotan or Ground Tumbler of India,
in not requiring to be shaken in order to begin tumbling. The
breed has probably been formed merely by selecting the best com-
-16 Journal of Horticulture,' Oct. 22,
1861, p. 76.
17 See the account of the House-tum-
blers kept at Glasgow, in the Cottage
Gardener,'1858, p. 285. Also Mr. Brent's
paper, Journal of Horticulture,'1861, p.
76.

188
CHAP. V.
DOMESTIC PIGEONS.
Fig. 23.-Short-faced English Tumbler.
mon Tumblers, though it is possible that they may have been crossed
at some former period with Lotans.

CHAP. V.
189
DESCRIPTION OF BREEDS.
Sub-race IV. Short-faced Tumblers. These are marvellous birds
and are the glory and pride of many fanciers. In their extremely
short, sharp, and conical beaks, with the skin over the nostrils but
little developed, they almost depart from the type of the Columbidæ.
Their heads are nearly globular and upright in front, so that some
fanciers say 18 " the head should resemble a cherry with a barley-
corn stuck in it." These are the smallest kind of pigeons. Mr. Es
quilant possessed a blue Baldhead, two years old, which when alive
weighed, before feeding-time, only 6 oz. 5 drs. ; two others each
weighed 7 oz. We have seen that a wild rock-pigeon weighed 14
oz. 2 drs, and a Runt 34 oz. 4 drs. Short-faced Tumblers have a
remarkably erect carriage, with prominent breasts, drooping wings,
and very small feet. The length of the beak from the tip to the
feathered base was in one good bird only 4 of an inch ; in a wild
rock-pigeon it was exactly double this length. As these Tumblers
have shorter bodies than the wild rock-pigeons, they ought of course
to have shorter beaks; but proportionally with the size of the body,
the beak is 28 of an inch too short. So, again, the feet of this bird
were actually .45 shorter, and proportionally .21 of an inch shorter,
than the feet of the rock-pigeon. The middle toe has only twelve or
thirteen, instead of fourteen or fifteen scutellæ. The primary wing-
feathers are not rarely only nine instead of ten in number. The im-
proved short-faced Tumblers have almost lost the power of tum-
bling; but there are several authentic accounts of their occasionally
tumbling There are several sub-varieties, such as Baldheads,
Beards, Mottles, and Almonds; the latter are remarkable from not
acquiring the perfectly-coloured plumage until they have moulted
three or four times. There is good reason to believe that most of
these sub-varieties, some of which breed truly, have arisen since the
publication of Moore's treatise in 1735.19
Finally, in regard to the whole group of Tumblers, it is impossi-
ble to conceive a more perfect gradation than I have now lying be-
fore me, from the rock-pigeon, through Persian, Lotan, and Common
Tumblers, up to the marvellous short-faced birds; which latter, no
ornithologist, judging from mere external structure, would place in
the same genus with the rock-pigeon. The differences between the
successive steps in this series are not greater than those which may
be observed between common dovecot-pigeons (C. livia) brought
from different countries.
18 J. M. Eaton's Treatise on Pigeons,'
1852, p. 9.
19 J. M. Eaton's Treatise, edit. 1858,
p. 76,

190
CHAP. V.
DOMESTIC PIGEONS.
RACE VIII.-INDIAN FRILL-BACK.
Beak very short; feathers reversed.
A specimen of this bird, in spirits, was sent to me from Madras
by Sir W. Elliot. It is wholly different from the Frill-back often
exhibited in England. It is a smallish bird, about the size of the
common Tumbler, but has a beak in all its proportions like our
short-faced Tumblers. The beak, measured from the tip to the
feathered base, was only '46 of an inch in length. The feathers
over the whole body are reversed or curl backwards. Had this
bird occurred in Europe, I should have thought it only a monstrous
variety of our improved Tumbler; but as short-faced Tumblers are
not known in India, I think it must rank as a distinct breed. Proba-
bly this is the breed seen by Hasselquist in 1757 at Cairo, and said
to have been imported from India.
RACE IX. JACOBIN.
(Zopf or Perücken-Taube: Non-
nains.)
Feathers of the neck forming a hood; wings and tail
long; beak moderately short.
This pigeon can at once be recognised by its hood, almost enclos-
ing the head and meeting in front of the neck. The hood seems to
be merely an exaggeration of the crest of reversed feathers on the
back of the head, which is common to many sub-varieties, and which
in the Latz-taube 20 is in a nearly intermediate state between a hood
and a crest. The feathers of the hood are elongated. Both the
wings and tail are likewise much elongated ; thus the folded wing
of the Jacobin, though a somewhat smaller bird, is fully 17 inch
longer than in the rock-pigeon. Taking the length of the body
without the tail as the standard of comparison, the folded wing,
proportionally with the wings of the rock-pigeon, is 24 inches too
long, and the two wings, from tip to tip, 51 inches too long. In
disposition this bird is singularly quiet, seldom fying or moving
about, as Bechstein and Riedel have likewise remarked in Ger-
many. The latter author also notices the length of the wings
and tail. The beak is nearly 2 of an inch shorter in proportion to
the size of the body than in the rock-pigeon; but the internal gape
of the mouth is considerably wider.
21
20 Neumeister, Taubenzucht,' Tab. 4,
fig. i.
21 Riedel, Die Taubenzucht,' 1824, S.
26. Bechstein, Naturgeschichte Deutsch.
lands,' Band iv. s. 36, 1795.

CAP. V.
191
DESCRIPTION OF BREEDS.
GROUP IV.
The birds of this group may be characterised by their
resemblance in all important points of structure, espe-
cially in the beak, to the rock-pigeon. The Trumpeter
forms the only well-marked race. Of the numerous other
sub-races and varieties I shall specify only a few of the
most distinct, which I have myself seen and kept alive.
RACE X.-TRUMPETER. (Trommel-Taube; Pigeon tam-
bour; glougou.)
A tuft of feathers at the base of the beak curling for-
ward ; feet much feathered ; voice very peculiar ; size ex-
ceeding that of the rock-pigeon.
This is a well-
marked breed, with a peculiar voice, wholly unlike
that of any other pigeon. The coo is rapidly repeated, and is con-
tinued for several minutes; hence their name of Trumpeters. They
are also characterised by a tuft of elongated feathers, which curls
forward over the base of the beak, and which is possessed by no
other breed. Their feet are so heavily feathered, that they almost
appear like little wings. They are larger birds than the rock-
pigeon, but their beak is of very nearly the same proportional size.
Their feet are rather small. This breed was perfectly characterised
in Moore's time, in 1735. Mr. Brent says that two varieties exist,
which differ in size.
RACE XI.-Scarcely differing in structure from the wild
Columba livia.
Sub-race I. Laughers. Size less than the Rock-pigeon ; voice very
peculiar.-As this bird agrees in nearly all its proportions with the
rock-pigeon, though of smaller size, I should not have thought it
worthy of mention, had it not been for its peculiar voice--a cha-
racter supposed seldom to vary with birds. Although the voice of
the Laugher is very different from that of the Trumpeter, yet one
of my Trumpeters used to utter a single note like that of the
Laugher. I have kept two varieties of Laughers, which differed
only in one variety being turn-crowned; the smooth-headed kind,
for which I am indebted to the kindness of Mr. Brent, besides its
peculiar note, used to coo in a singular and pleasing manner, which,

192
CHAP. V.
DOMESTIC PIGEONS.
independently, struck both Mr. Brent and myself as resembling that
of the turtle dove. Both varieties come from Arabia. This breed
was known by Moore in 1735. A pigeon which seems to say Yak-
roo is mentioned in 1600 in the 'Ayeen Akbery,' and is probably the
same breed. Sir W. Elliot has also sent me from Madras a pigeon
called Yahui, said to have come from Mecca, which does not differ
in appearance from the Laugher; it has “a deep melancholy voice,
like Yahu, often repeated.” Yalu, yahu, means Oh God, Oh God;
and Sayzid Mohammed Musari, in the treatise written about 100
years ago, says that these birds "are not flown, because they repeat
the name of the Most High God." Mr. Keith Abbott, however, in-
forms me that the common pigeon is called Yahoo in Persia.
Sub-race II. Common Frill-back (Die Strupp-Taube). Beak rather
longer than in the Rock-pigeon ; feathers reversed.--This is a con-
siderably larger bird than the rock-pigeon, and with the beak, pro-
portionally with the size of body, a little (viz. by .04 of an inch)
longer. The feathers, especially on the wing-coverts, have their
points curled upwards or backwards.
Sub-race III. Nuns (Pigeons-coquilles).-— These elegant birds are
smaller than the rock-pigeon. The beak is actually 17, and pro-
portionally with the size of the body 1 of an inch shorter than in
the rock-pigeons, although of the same thickness. In young birds
the scutellæ on the tarsi and toes are generally of a leaden-black
colour; and this is a remarkable character (though observed in a
lesser degree in some other breeds), as the colour of the legs in the
adult state is subject to very little variation in any breed. I have
on two or three occasions counted thirteen or fourteen feathers in
the tail ; this likewise occurs in the barely distinct breed called
Helmets. Nuns are symmetrically coloured, with the head, primary
wing-feathers, tail, and tail-coverts of the same colour, namely,
black or red, and with the rest of the body white. This breed has
retained the same character since Aldrovandi wrote in 1600. I have
received from Madras almost similarly colored birds.
Sub-race IV. Spots (Die Blass-Taube: Pigeons heurtés).—These
birds are a very little larger than the rock-pigeon, with the beak a
trace smaller in all its dimensions, and with the feet decidedly
smaller. They are symmetrically coloured, with a spot on the fore-
head, with the tail and tail-coverts of the same colour, the rest of
the body being white. This breed existed in 1676,22 and in 1735
Moore remarks that they breed truly, as is the case at the present
day.
Sub-race V. Stallows.—These birds, as measured from tip to tip
22 Willoughby's 'Ornithology,' edited by Ray.

CHAP. V.
193
DESCRIPTION OF BREEDS.
of wing, or from the end of the beak to the end of the tail, exceed
in size the rock-pigeon ; but their bodies are much less bulky; their
feet and legs are likewise smaller. The beak is of about the same
length, but rather slighter. Altogether their general appearance
is considerably different from that of the rock-pigeon. Their heads
and wings are of the same colour, the rest of the body being white.
Their flight is said to be peculiar. This seems to be a modern
breed, which, however, originated before the year 1795 in Germany.
for it is described by Bechstein.
Besides the several breeds now described, three or four other very
distinct kinds existed lately, or perhaps still exist, in Germany and
France. Firstly, the Karmeliten, or Carme Pigeon, which I have not
seen; it is described as of small size with very short legs, and with
an extremely short beak. Secondly, the Finnikin, which is now
extinct in England. It had, according to Moore's 23 treatise, pub-
lished in 1735, a tuft of feathers on the hinder part of the head,
which ran down its back not unlike a horse's mane. “When it is
salacious it rises over the hen and turns round three or four times,
flapping its wings, then reverses and turns as many times the other
way." The Turner, on the other hand, when it "plays to the
female, turns only one way." Whether these extraordinary state
ments may be trusted I know not; but the inheritance of any habit
may be believed, after what we have seen with respect to the
Ground-tumbler of India. MM. Boitard and Corbié describe a pi.
geon 24 which has the singular habit of sailing for a considerable
time through the air, without flapping its wings, like a bird of
prey. The confusion is inextricable, from the time of Aldrovandi
in 1600 to the present day, in the accounts published of the Draijers,
Smiters, Finnikins, Turners, Claquers, &c., which are all remarkable
from their manner of flight. Mr. Brent informs me that he has
seen one of these breeds in Germany with its wing-feathers injured
from having been so often struck together, but he did not see it
flying. An old stuffed face of a Finnikin in the British Museum
presents no well-marked character. Thirdly, a singular pigeon with
a forked tail is mentioned in some treatises; and as Bechstein 25 briefly
describes and figures this bird, with a tail" having completely the
structure of that of the house-swallow," it must once have existed,
for Bechstein was far too good a naturalist to have confounded any
distinct species with the domestic pigeon. Lastly, an extraordinary
pigeon imported from Belgium has lately been exhibited at the
23 J. M. Eaton's edition (1858) of
Moore, p. 98.
24 Pigeon Patu Plongeur. Les Pi-
geons,' &c., p. 165.
25 Naturgesch, Deutschlands, Band
iv. s. 47.
9

194
CHAP, V.
DOMESTIC PIGEONS,
Philoperisteron Society in London,26 which “conjoins the colour of
an archangel with the head of an owl or barb, its most striking
peculiarity being the extraordinary length of the tail and wing-
feathers, the latter crossing beyond the tail, and giving to the bird
the appearance of a gigantic swift (Cypselus), or long-winged
hawk." Mr. Tegetmeier informs me that this bird weighed only
10 ounces, but in length was 154 inches from tip of beak to end of
tail, and 321 inches from tip to tip of wing ; now the wild rock-
pigeon weighs 141 ounces, and measures from tip of beak to end of
tail 15 inches, and from tip to tip of wing only 264 inches.
I have now described all the domestic pigeons known
to me, and have added a few others on reliable authority.
I have classed them under four Groups, in order to mark
their affinities and degrees of difference; but the third
group is artificial.
The kinds examined by me form
eleven races, which include several sub-races; and even
these latter present differences that would certainly have
been thought of specific value if observed in a state of
nature. The sub-races likewise include many strictly in-
herited varieties; so that altogether there must exist, as
previously stated, above 150 kinds which can be dis-
tinguished, though generally by characters of extremely
slight importance. Many of the genera of the Colum-
bidæ, which are admitted by ornithologists, do not dif-
fer in any great degree from each other; taking this into
consideration, there can be no doubt that several of the
most strongly characterised domestic forms, if found
wild, would have been placed in at least five new gene-
ra. Thus, a new genus would have been formed for the
reception of the improved English Pouter: a second
genus for Carriers and Runts; and this would have been
a wide or comprehensive genus, for it would have ad-
mitted common Spanish Runts without any wattle, short-
beaked Runts like the Tronfo, and the improved English
Carrier: a third genus would have been formed for the
Barb: a fourth for the Fantail: and lastly, a fifth for the
26 Mr, W. B. Tegetmeier, Journal of Horticulture,' Jan. 20th, 1863, p. 58.

CAP. V.
195
DESCRIPTION OF BREEDS.
short-beaked, not-wattled pigeons, such as Turbits and
short-faced Tumblers.
The remaining domestic forms
might have been included in the same genus with the
wild rock-pigeon.
Individual Variability; Variations of a remarkable
nature.
The differences which we have as yet considered are
characteristic of distinct breeds; but there are other dif-
ferences, either confined to individual birds, or often ob-
served in certain breeds but not characteristic of them.
These individual differences are of importance, as they
might in most cases be secured and accumulated by man's
power of selection; and thus an existing breed might be
greatly modified or a new one formed. Fanciers notice
and select only those slight differences which are exter-
nally visible; but the whole organisation is so tied to-
gether by correlation of growth, that a change in one
part is frequently accompanied by other changes. For
our purpose, modifications of all kinds are equally im-
portant, and, if affecting a part which does not common-
ly vary, are of more importance than a modification in
some conspicuous part. At the present day any visible
deviation of character in a well-established breed is re-
jected as a blemish; but it by no means follows that
at an early period, before well-marked breeds had been
formed, such deviations would have been rejected; on
the contrary, they would have been eagerly preserved as
presenting a novelty, and would then have been slowly
augmented, as we shall hereafter more clearly see, by the
process of unconscious selection.
I have made numerous measurements of the various parts of the
body in the several breeds, and have hardly ever found them quite
the same in birds of the same breed, -the differences being greater
than we commonly meet with in wild species. To begin with the
primary feathers of the wing and tail ; but I may first mention, as
some readers may not be aware of the fact, that the number of the

196
CHAP. V.
DOMESTIC PIGEONS.
primary wing and tail feathers in wild birds is generally constant,
and characterises, not only whole genera, but even whole families.
When the tail-feathers are unusually numerous, as for instance in
the swan, they are apt to be variable in number; but this does not
apply to the several species and genera of the Columbidæ, which
never (as far as I can hear) have less than twelve or more than
sixteen tail-feathers; and these numbers characterise, with rare ex-
ception, whole sub-families.27 The wild rock-pigeon has twelve tail-
feathers. With Fantails, as we have seen, the number varies from
fourteen to forty-two. In two young birds in the same nest I
counted twenty-two and twenty-seven feathers. Pouters are very
liable to have additional tail-feathers, and I have seen on several oc-
casions fourteen or fifteen in my own birds. Mr. Bult had a speci-
men, examined by Mr. Yarrell, with seventeen tail-feathers. I had
a Nun with thirteen, and another with fourteen tail-feathers; and in
a Helmet, a breed barely distinguishable from the Nun, I have
counted fifteen, and have heard of other such instances. On the
other hand, Mr. Brent possessed a Dragon, which during its whole life
nover had more than ten tail-feathers; and one of my Dragons, de-
scended from Mr. Brent's, had only eleven. I have seen a Baldhead-
Tumbler with only ten ; and Mr. Brent had an Air-Tumbler with the
same number, but another with fourteen tail-feathers. Two of these
latter Tumblers, bred by Mr. Brent, were remarkable,-one from
having the two central tail-feathers a little divergent, and the other
from having the two outer feathers longer by three-eighths of an
inch than the others; so that in both cases the tail exhibited a
tendency, but in different ways, to become forked. And this shows
us how a swallow-tailed breed, like that described by Bechstein,
might have been formed by careful selection.
With respect to the primary wing-feathers, the number in the Co-
lumbidæ, as far as I can find out, is always nine or ten. In the rock-
pigeon it is ten; but I have seen no less than eight short-faced
Tumblers with only nine primaries, and the occurrence of this
number has been noticed by fanciers, owing to ten flight feathers
of a white colour being one of the points in Short-faced Baldhead-
Tumblers. Mr. Brent, however, had an Air-Tumbler (not short-
faced) which had in both wings eleven primaries. Mr. Corker, the
eminent breeder of prize Carriers, assures me that some of his birds
27 Coup-d'oeil sur l'Ordre des Pigeons,
par C. L. Bonaparte (Comptes Rendus),
1954-55. Mr. Blyth, in ' Annals of Nat.
Hist.,' vol. xix., 1847, p. 41, mentions, as
a very singular fact, that of the two
species of Ectopistes, which are nearly
allied to each other, one should have
fourteen tail-feathers, while the other,
the passenger pigeon of North America,
should possess but the usual number-
twelve."

CHAP. V.
197
INDIVIDUAL VARIABILITY.
had eleven primaries in both wings. I have seen eleven in one
wing in two Pouters. I have been assured by three fanciers that
they have seen twelve in Scanderoons; but as Neumeister asserts
that in the allied Florence Runt the middle flight-feather is often
double, the number twelve may have been caused by two of the ten
primaries having each two shafts to a single feather. The second-
ary wing-feathers are difficult to count, but the number seems to
vary from twelve to fifteen. The length of the wing and tail rela-
tively to the body, and of the wings to the tail, certainly varies; I
have especially noticed this in Jacobins. In Mr. Bult's magnificent
collection of Pouters, the wings and tail varied greatly in length;
and were sometimes so much elongated that the birds could hardly
play upright. In the relative length of the few first primaries I
have observed only a slight degree of variability. Mr. Brent in-
forms me that he has observed the shape of the first feather to vary
very slightly. But the variation in these latter points is extremely
slight compared with what may often be observed in the natural
species of the Columbidæ.
In the beak I have observed very considerable differences in birds
of the same breed, as in carefully bred Jacobins and Trumpeters.
In Carriers there is often a conspicuous difference in the degree of
attenuation and curvature of the beak. So it is indeed in many
breeds: thus I had two strains of black Barbs, which evidently dif-
fered in the curvature of the upper mandible. In width of mouth I
have found a great difference in two swallows. In Fantails of first-
rate merit I have seen some birds with much longer and thinner
necks than in others. Other analogous facts could be given. We
have seen that the oil-gland is aborted in all Fantails (with the ex-
ception of the sub-race from Java), and, I may add, so hereditary is
this tendency to abortion, that some, although not all, of the mon-
grels from the Fantail and Pouter had no oil-gland ; in one Swallow
out of many which I have examined, and in two Nuns, there was no
oil-gland.
The number of the scutellæ on the toes often varies in the same
breed, and sometimes even differs on the two feet of the same indi-
vidual; the Shetland rock-pigeon has fifteen on the middle, and six
on the hinder toe; whereas I have seen a Runt with sixteen on the
middle, and eight on the hind toe ; and a short-faced Tumbler with
only twelve and five on these same toes. The rock-pigeon has no
sensible amount of skin between its toes; but I possessed a Spot and
a Nun with the skin extending for a space of a quarter of an inch
from the fork, between the two inner toes. On the other hand, as
will hereafter be more fully shown, pigeons with feathered feet very
generally have the bases of their outer toes connected by skin. I

198
CHAP. V.
DOMESTIC PIGEONS.
had a red Tumbler, which had a coo unlike that of its fellows, ap-
proaching in tone to that of the Laugher: this bird had the habit,
to a degree which I never saw equalled in any other pigeon, of often
walking with its wings raised and arched in an elegant manner. I
need say nothing on the great variability, in almost every breed, in
size of body, in colour, in the feathering of the feet, and in the
feathers on the back of the head being reversed. But I may men-
tion a remarkable Tumbler 28 exhibited at the Crystal Palace, which
had an irregular crest of feathers on its head, somewhat like the
tuft on the head of the Polish fowl. Mr. Bult reared by accident a
hen Jacobin with the feathers on the thigh so long as to reach the
ground, and a cock having, but in a lesser degree, the same pecu-
liarity: from these two birds he bred others similarly characterised,
which were exhibited at the Philoperisteron Club. I bred a mon-
grel pigeon which had fibrous feathers, and the wing and tail-
feathers so short and imperfect that the bird could not fly even a
foot in height.
There are many singular and inherited peculiarities in
the plumage of pigeons: thus Almond-Tumblers do not
acquire their perfect mottled feathers until they have
moulted three or four times: the Kite-Tumbler is at first
brindled black and red with a barred appearance, but
when “it throws its nest feathers it becomes almost
black, generally with a bluish tail, and a reddish colour
on the inner webs of the primary wing feathers." 29 Neu-
meister describes a breed of a black colour with white
bars on the wing and a white crescent-shaped mark on
the breast; these marks are generally rusty-red before
the first moult, but after the third or fourth moult they
undergo a change; the wing-feathers and the crown of
the head likewise then become white or grey,s
It is an important fact, and I believe there is hardly
an exception to the rule, that the especial characters for
which each breed is valued are eminently variable: thus,
in the Fantail, the number and direction of the tail-feath-
ers, the carriage of the body, and the degree of trembling
28 Described and figured in the Poul-
try Chronicle,' vol. iii., 1855, p. 82.
29 The Pigeon Book,' by Mr. B. P.
Brent, 1859, p. 41.
30 Die Staarhälsige Taube, Das
Ganze, &c.,' s. 21, tab. i. fig. 4.

CHAP. V.
199
SINGULAR VARIATIONS.
are all highly variable points; in Pouters, the degree to
which they pout, and the shape of their inflated crops; in
the Carrier, the length, narrowness, and curvature of the
beak, and the amount of wattle; in Short-faced Tumblers,
the shortness of the beak, the prominence of the forehead,
and general carriage, and in the Almond Tumbler the
colour of the plumage; in common Tumblers, the manner
of tumbling; in the Barb, the breadth and shortness of
the beak and the amount of eye-wattle; in Runts, the
size of body; in Turbits, the frill; and lastly in Trumpet-
ers, the cooing, as well as the size of the tuft of feathers
over the nostrils. These, which are the distinctive and
selected characters of the several breeds, are all eminently
variable.
There is another interesting fact with respect to the
character of the different breeds, namely, that they are
often most strongly displayed in the male bird. In Car-
riers, when the males and females are exhibited in sepa-
rate pens, the wattle is plainly seen to be much more de-
veloped in the males, though I have seen a hen Carrier
belonging to Mr. Haynes heavily wattled. Mr. Teget-
meier informs me that, in twenty Barbs in Mr. P. H.
Jones's possession, the males had generally the largest
eye-wattles; Mr. Esquilant also believes in this rule, but
Mr. H. Weir, a first-rate judge, entertains some doubt on
the subject. Male Pouters distend their crops to a much
greater size than do the females; I have, however, seen a
hen in the possession of Mr. Evans which pouted excel-
lently; but this is an unusual circumstance. Mr. Harri-
son Weir, a successful breeder of prize Fantails, informs
me that his cock birds often have a greater number of
tail-feathers than the hens. Mr. Eaton asserts 92 that, if
a cock and hen Tumbler were of equal merit, the hen
would be worth double the money; and as pigeons al-
31 A Treatise on the Almond Tum-
bler,' by J. M. Eaton, 1852, p. 8, et pas-
sim.
32 A Treatise, &c., p. 10.

200
CHAP. V.
DOMESTIC PIGEONS.
33
ways pair, so that an equal number of both sexes is ne-
cessary for reproduction, this seems to show that high
merit is rarer in the female than in the male. In the de-
velopment of the frill in Turbits, of the hood in Jacobins,
of the tuft in Trumpeters, of tumbling in Tumblers, there
is no difference between the males and females. I may
here add a rather different case, namely, the existence in
France of a wine-coloured variety of the Pouter, in
which the male is generally chequered with black, whilst
the female is never so chequered. Dr. Chapuis also re-
marks 34 that in certain light-coloured pigeons the males
have their feathers striated with black, and these striæ
increase in size at each moult, so that the male ultimately
becomes spotted with black. With Carriers, the wattle,
both on the beak and round the eyes, and with Barbs
that round the eyes, goes on increasing with age. This
augmentation of character with advancing age, and more
especially the difference between the males and females
in the above-mentioned several respects, are highly re-
markable facts, for there is no sensible difference at any
age between the two sexes in the aboriginal rock-pigeon;
and rarely any such difference throughout the whole fam-
ily of the Columbidæ.36
Osteological Characters.
In the skeletons of the various breeds there is much
variability; and though certain differences occur frequent-
ly, and others rarely, in certain breeds, yet none can be
33 Boitard and Corbié, 'Les Pigeons,
&c., 1824, p. 173.
34 'Le Pigeon Voyageur Belge,' 1865,
p. 87.
35 Prof. A. Newton (Proc. Zoolog.
Soc.,' 1865, p. 716) remarks that he
knows no species which presents any re-
markable sexual distinction ; but it is
stated "Naturalist's Library, Birds,' vol.
ix. p. 117) that the excrescence at the
base of the beak in the Carpophaga
oceanica is sexual: this, if correct, is an
interesting point of analogy with the male
Carrier, which has the wattle at the base
of its beak so much more developed than
in the female. Mr. Wallace informs me
that in the sub-family of the Treronidæ
the sexes often differ in vividness of
colour.

CHAP. V.
201
OSTEOLOGICAL DIFFERENCES.
A
said to be absolutely characteristic of
any
breed. Con-
sidering that strongly marked domestic races have been
formed chiefly by man's power of selection, we ought not
to expect to find great and constant differences in the
skeleton; for fanciers can neither see, nor do they care
for, modifications of structure in the internal framework.
Nor ought we to ex-
pect changes in the
skeletons from
changed habits of
life; as every facili-
ty is given to the
most distinct breeds
to follow the same
habits, and the much
modified races
allowed to
wander abroad and
B
are
never
с
D
Fig. 24.—Skulls of Pigeons, viewed laterally, of natural size. A. Wild Rock-
pigeon, Columba livia. B. Short-faced Tumbler. C. English Carrier. D.
Bagadotten Carrier.
procure their own food in various ways. Moreover, I

202
CHAP. V.
DOMESTIC PIGEONS.
find, on comparing the skeletons of Columba livia, oenas,
palumbus, and turtur, which are ranked by all systema-
tists in two or three distinct though allied genera,
that
the differences are extremely slight, certainly less than
between the skeletons of some of the most distinct do-
mestic breeds. How far the skeleton of the wild rock-
pigeon is constant I have no means of judging, as I have
examined only two.
Skull.—The individual bones, especially those at the base, do not
differ in shape. But the whole skull, in its proportions, outline,
and relative direction of the bones, differs greatly in some of the
breeds, as may be seen by comparing the figures of (A) the wild
rock-pigeon, (B) the short-faced tumbler, (C) the English carrier, and
(D) the Bagadotten carrier (of Neumeister), all drawn of the natural
size and viewed laterally. In the carrier, besides the elongation of
the bones of the face, the space between the orbits is proportionally a
little narrower than in the rock-pigeon. In the Bagadotten the upper
mandible is remarkably arched, and the premaxillary bones are pro-
portionally broader. In the short-faced tumbler the skull is more
globular; all the bones of the face are much shortened, and the front
of the skull and descending nasal bones are almost perpendicular; the
maxillo-jugal arch and premaxillary bones form an almost straight
line; the space between the prominent edges of the eye-orbits is
А
B
Fig. 25-Lower jaws, seen from above, of natural size. A. Rock-pigeon. B. Runt.
C. Barb.

CHAP. V.
203
OSTEOLOGICAL DIFFERENCES.
depressed. In the barb the premaxillary bones are much short-
ened, and their anterior portion is thicker than in the rock-pigeon,
as is the lower part of the nasal bone. In two nuns the ascending
branches of the premaxillaries, near their tips, were somewhat at-
tenuated, and in these birds, as well as in some others, for instance
in the spot, the occipital crest over the foramen was considerably
more prominent than in the rock-pigeon.
In the lower jaw, the articular surface is proportionally smaller
in many breeds than in the rock-pigeon; and the vertical diameter
more especially of the outer part of the articular surface is consid-
ably shorter. May not this be accounted for by the lessened use
of the jaws, owing to nutritious food having been given during a
long period to all highly improved pigeons? In runts, carriers,
and barbs (and in a lesser degree in several breeds), the whole side
A
B
Fig. 27.- Lateral view of jaws, of natural size.
A. Rock-pigeon. B. Short-faced Tumbler. C.
Bagadotten Carrier.
of the jaw near the articular end is bent
inwards in a highly remarkable manner;
and the superior margin of the ramus, be-
yond the middle, is reflexed in an equally
Fig. 26.-Skull of Runt, seen
remarkable manner, as may be seen in the
from above, of natural size, accompanying figures, in comparison with
showing the reflexed mar-
the jaw of the rock-pigeon. This reflexion
gin of the distal portion of of the upper margin of the lower jaw is
the lower jaw.
plainly connected with the singularly
wide gape of the mouth, as has been described in runts, carriers, and
barbs. The reflexion is well shown in fig. 26 of the head of a runt
seen from above; here a wide open space may be observed on each

204
CHAP. V.
DOMESTIC PIGEONS.
side, between the edges of the lower jaw and of the premaxillary
bones. In the rock-pigeon, and in several domestic breeds, the
edges of the lower jaw on each side come close up to the premaxil-
lary bones, so that no open space is left. The degree of downward
curvature of the distal half of the lower jaw also differs to an ex-
traordinary degree in some breeds, as may be seen in the drawings
(fig. A) of the rock-pigeon, (B) of the short-faced tumbler, and (C) of
the Bagadotten carrier of Neumeister. In some runts the symphy-
sis of the lower jaw is remarkably solid. No one would readily
have believed that jaws differing so greatly in the several above-
specified points could have belonged to the same species.
Vertebra. All the breeds have twelve cervical vertebræ.36 But
in a Bussorah carrier from India, the twelfth vertebra carried a
small rib, a quarter of an inch in length, with a perfect double arti-
culation,
The dorsal vertebræ are always eight. In the rock-pigeon all eight
bear ribs; the eighth rib being very thin, and the seventh having
no process. In pouters all the ribs are extremely broad, and, in three
out of four skeletons examined by me, the eighth rib was twice or
even thrice as broad as in the rock-pigeon; and the seventh pair had
distinct processes. In many breeds there are only seven ribs, as in
seven out of eight skeletons of various tumblers, and in several ske-
letons of fantails, turbits, and nuns. In all these breeds the seventh
pair was very small, and was destitute of processes, in which respect
it differed from the same rib in the rock-pigeon. In one tumbler,
and in the Bussorah carrier, even the sixth pair had no process.
The hypapophysis of the second dorsal vertebra varies much in de-
velopment; being sometimes (as in several, but not all tumblers)
nearly as prominent as that of the third dorsal vertebra; and the
two hypapophyses together tend to form an ossified arch. The de-
velopment of the arch, formed by the hypapophyses of the third and
fourth dorsal vertebra, also varies considerably, as does the size of
the hypapophysis of the fifth vertebra.
The rock-pigeon has twelve sacral vertebrae ; but these vary in
number, relative size, and distinctness in the different breeds. In
pouters, with their elongated bodies, there are thirteen or even four-
teen, and, as we shall immediately see, an additional number of
caudal vertebræ. In runts and carriers there is generally the pro-
per number, namely twelve; but in one runt, and in the Bussorah
36 I am not sure that I have designa-
ted the different kinds of vertebræ cor-
rectly; but I observe that different ana-
tomists follow in this respect different
rules, and, as I use the same terms in
the comparison of all the skeletons, this,
I hope, will not signify.

CHAP. V.
205
OSTEOLOGICAL DIFFERENCES.
carrier, there were only eleven. In tumblers there are either eleven,
twelve, or thirt en sacral vertebræ.
The caudal vertebra are seven in number in the rock-pigeon. In
fantails, which have their tails so largely developed, there are either
eight or nine, and apparently in one case ten, and they are a little
longer than in the rock-pigeon, and their shape varies considerably.
Pouters, also, have eight or nine caudal vertebra. I have seen eight
in a nun and jacobin, Tumblers, though such small birds, always
have the normal number seven ; as have carriers, with one excep-
tion, in which there were only six.
The following table will serve as a summary, and will show the
most remarkable deviations in the number of the vertebræ and ribs
which I have observed :-
Rock Pigeon.
Pouter, from
Mr. Bult.
Tumbler,
Dutch Roller.
Bussorah
Carrier,
Cervical Vertebræ 12
12
12
12
The 12th bore
a small rib.
Dorsal Vertebræ
8
8
8.
8
Ribs ..
8
8
7
7
The 6th pair with The 6th and The 6th and 7th The 6th and
processes, the 7th 7th pair with pair without 7th pair
pair without a processes. processes.
without pro-
process.
cesses,
Sacral Vertebræ
12
14
11
11
Caudal Vertebræ
7
8 or 9
7
7
Total Vertebrae
39
42 or 43
38
38
The pelvis differs very little in any breed. The anterior margin
of the ilium, however, is sometimes a little more equally rounded on
both sides than in the rock-pigeon. The ischium is also frequently
rather more elongated. The obturator-notch is sometimes, as in
many tumblers, less developed than in the rock-pigeon. The ridges
on the ilium are very prominent in most runts.
In the bones of the extremities I could detect no difference, except
in their proportional lengths; for instance, the metatarsus in a pout-
er was 1.65 inch, and in a short-faced tumbler only .95 in length;
and this is a greater difference than would naturally follow from
their differently-sized bodies; but long legs in the pouter, and small
feet in the tumbler, are selected points. In some pouters the scapula
is rather straighter, and in some tumblers it is straighter, with the
apex less elongated, than in the rock-pigeon: in the woodcut, fig.
28, the scapulæ of the rock-pigeon (A), and of a short-faced tumbler

206
CHAP. V.
DOMESTIC PIGEONS.
B
er.
B
(B), are given. The processes at the sum-
mit of the coracoid, which receive the ex-
tremities of the furcula, form a more perfect
cavity in some tum-
blers than in the
rock-pigeon :
in
pouters these pro-
cesses are larger A
and differently
shaped, and the ex-
terior angle of the
extremity of the
coracoid, which is
articulated to the
A
sternum, is squar-
B
Fig. 28. -Scapulæ, of natu-
ral size. A. Rock-pigeon.
The two arms of
B. Short-faced Tumbler. the furcula in pout-
ers diverge less,
proportionally to their length, than in the
rock-pigeon ; and the symphysis is more so-
lid and pointed. In fantails the degree of di-
vergence of the two arms varies in a remark-
С
able manner. In fig. 29, B and c represent
the furculæ of two fantails; and it will be
seen that the divergence in B is rather less
even than in the furcula of the short-faced,
small-sized tumbler (A); whereas the diver-
gence in c equals that in a rock-pigeon, or
in the pouter (D), though the latter is a much
larger bird. The extremities of the furcu-
la, where articulated to the coracoids, vary
considerably in outline.
In the sternum the differences in form are
slight, except in the size and outline of the
perforations, which, both in the larger and
lesser sized breeds, are sometimes small.
These perforations, also, are sometimes eith-
er nearly circular, or elongated, as is often
the case with carriers. The posterior perfo-
rations occasionally are not complete, being Fig. 29.-Furculæ, of natu-
left open posteriorly. The marginal apophy-
ral size. A. Short-faced
ses forming the anterior perforations vary
Tumbler. B and C. Fan-
tails. D. Pouter.
greatly in development. The degree of
D

CHAP. V.
207
CORRELATION OF GROWTH.
convexity of the posterior part of the sternum differs much, being
sometimes almost perfectly flat. The manubrium is rather more
prominent in some individuals than in others, and the pore imme-
diately under it varies greatly in size.
Correlation of Growth.—By this term I mean that the
whole organisation is so connected, that when one part
varies, other parts vary; but which of two correlated
variations ought to be looked at as the cause and which
as the effect, or whether both result from some common
cause, we can seldom or never tell. The point of interest
for us is that, when fanciers, by the continued selection
of slight variations, have largely modified one part, they
often unintentionally produce other modifications. For
instance, the beak is readily acted on by selection, and,
with its increased or diminished length, the tongue in-
creases or diminishes, but not in due proportion; for, in
a barb and short-faced tumbler, both of which have very
short beaks, the tongue, taking the rock-pigeon as the
standard of comparison, was proportionally not shorten-
ed enough, whilst in two carriers and in a runt the tongue,
proportionally with the beak, was not lengthened enough.
Thus, in a first-rate English carrier, in which the beak from
the tip to the feathered base was exactly thrice as long asin
a first-rate short-faced tumbler, the tongue was only a little
more than twice as long. But the tongue varies in length
independently of the beak: thus, in a carrier with a beak
1.2 inch in length, the tongue was •67 in length; whilst
in a runt which equalled the carrier in length of body
and in stretch of wings from tip to tip, the beak was .92
whilst the tongue was •73 of an inch in length, so that the
tongue was actually longer than in the carrier with its
long beak. The tongue of the runt was also very
broad
at the root. Of two runts, one had its beak longer by
*23 of an inch, whilst its tongue was shorter by '14 than
in the other.
With the increased or diminished length of the beak
the length of the slit forming the external orifice of the

208
CHAP. V.
DOMESTIC PIGEONS.
nostrils varies, but not in due proportion, for, taking the
rock-pigeon as the standard, the orifice in a short-faced
tumbler was not shortened in due proportion with its very
short beak. On the other hand (and this could not have
been anticipated), the orifice in three English carriers, in
the Bagadotten carrier, and in a runt (pigeon cygne), was
longer by above the tenth of an inch than would follow
from the length of the beak proportionally with that of
the rock-pigeon. In one carrier the orifice of the nostrils
was thrice as long as in the rock-pigeon, though in body
and length of beak this bird was not nearly double the
size of the rock-pigeon. This greatly increased length
of the orifice of the nostrils seems to stand partly in cor-
relation with the enlargement of the wattled skin on the
upper mandible and over the nostrils; and this is a cha-
racter which is selected by fanciers. So again, the broad,
naked, and wattled skin round the eyes of carriers and
barbs is a selected character; and in obvious correlation
with this, the eyelids measured longitudinally, are pro-
portionally more than double the length of those of the
rock-pigeon.
The great difference (see woodcut No. 27) in the cur-
vature of the lower jaw in the rock-pigeon, the tumbler,
and Bagadotten carrier, stands in obvious relation to the
curvature of the upper jaw, and more especially to the an-
gle formed by the maxillo-jugal arch with the premaxil-
lary bones. But in carriers, runts, and barbs the singu-
lar reflexion of the upper margin of the middle part of the
lower jaw (see woodcut No. 25) is not strictly correlated
with the width or divergence (as may be clearly seen in
woodcut No. 26) of the premaxillary bones, but with the
breadth of the horny and soft parts of the upper mandible,
which are always overlapped by the edges of the lower
mandible.
In pouters, the elongation of the body is a selected cha-
racter, and the ribs, as we have seen, have generally be-
come very broad, with the seventh pair furnished with

CHAP. V.
209
CORRELATION OF GROWTH.
processes; the sacral and caudal vertebræ have been aug-
mented in number; the sternum has likewise increased in
length (but not in the depth of the crest) by 4 of an inch
more than would follow from the greater bulk of the body
in comparison with that of the rock-pigeon. In fantails,
the length and number of the caudal vertebræ have in-
creased. Hence, during the gradual progress of varia-
tion and selection, the internal bony frame-work and the
external shape of the body have been, to a certain extent,
modified in a correlated manner.
Although the wings and tail often vary in length in-
dependently of each other, it is scarcely possible to doubt
that they generally tend to become elongated or short-
ened in correlation. This is well seen in jacobins, and
still more plainly in runts, some varieties of which have
their wings and tail of great length, whilst others have
both very short. With jacobins, the remarkable length
of the tail and wing-feathers is not a character which is
intentionally selected by fanciers; but fanciers have been
trying for centuries, at least since the year 1600, to in-
crease the length of the reversed feathers on the neck, so
that the hood may more completely enclose the head;
and it may be suspected that the increased length of the
wing and tail-feathers stands in correlation with the in-
creased length of the neck-feathers. Short-faced tumblers
have short wings in nearly due proportion with the re-
duced size of their bodies; but it is remarkable, seeing
that the number of the primary wing-feathers is a con-
stant character in most birds, that these tumblers gene-
rally have only nine instead of ten primaries. I have
myself observed this in eight birds; and the Original
Columbarian Society $reduced the standard for bald-head
tumblers from ten to nine white flight-feathers, thinking
it unfair that a bird which had only nine feathers should
be disqualified for a prize because it had not ten white
37 J. M. Eaton's Treatise, edit. 1858, p. 78.

210
CHAP. V.
DOMESTIC PIGEONS.
flight-feathers. On the other hand, in carriers and runts,
which have large bodies and long wings, eleven primary
feathers have occasionally been observed.
Mr. Tegetmeier has informed me of a curious and in-
explicable case of correlation, namely, that young pigeons
of all breeds, which when mature become white, yellow,
silver, (i.e. extremely pale blue), or dun-coloured, are born
almost naked; whereas other coloured pigeons are born
well clothed with down. Mr. Esquilant, however, has
observed that young dun carriers are not so bare as
young dun barbs and tumblers. Mr. Tegetmeier has
seen two young birds in the same nest, produced from
differently coloured parents, which differed greatly in
the degree to which they were at first clothed with
down.
I have observed another case of correlation which at
first sight appears quite inexplicable, but on which, as
we shall see in a future chapter, some light can be thrown
by the law of homologous parts varying in the same
manner. The case is, that, when the feet are much
feathered, the roots of the feathers are connected by a
web of skin, and apparently in correlation with this the
two outer toes become connected for a considerable space
by skin. I have observed this in very many specimens
of pouters, trumpeters, swallows, roller-tumblers (like-
wise observed in this breed by Mr. Brent), and in a lesser
degree in other feather-footed pigeons.
The feet of the smaller and larger breeds are of course
much smaller or larger than those of the rock-pigeon;
but the scutellæ or scales covering the toes and tarsi
have not only decreased or increased in size, but likewise
in number. To give a single instance, I have counted
eight scutellä on the hind toe of a runt, and only five on
that of a short-faced tumbler. With birds in a state of
nature the number of the scutellæ on the feet is usually
a constant character. The length of the feet and the
length of the beak apparently stand in correlation; but

CHAP. V.
211
ON THE EFFECTS OF DISUSE.
as disuse apparently has affected the size of the feet, this
case may come under the following discussion.
On the Effects of Disuse. In the following discussion
on the relative proportions of the feet, sternum, furcula,
scapulæ, and wings, I may premise, in order to give some
confidence to the reader, that my ineasurements were all
made in the same manner, and that all the measurements
of the external parts were made without the least inten-
tion of applying them to the following purpose.
I measured most of the birds which came into my possession, from
the feathered base of the beak (the length of beak itself being so
variable) to the end of the tail, and to the oil-gland, but unfortu-
nately (except in a few cases) not to the root of the tail ; I measured
each bird from the extreme tip to tip of wing; and the length of
the terminal folded part of the wing, from the extremity of the
primaries to the joint of the radius. I measured the feet without
the claws, from the end of the middle toe to the end of the hind
toe; and the tarsus together with the middle toe. I have taken in
every case the mean measurement of two wild rock-pigeons from
the Shetland Islands, as the standard of comparison. The following
table shows the actual length of the feet in each bird ; and the
difference between the length which the feet ought to have had
according to the size of body of each, in comparison with the size
of body and length of feet of the rock-pigeon, calculated (with a
few specified exceptions) by the standard of the length of the body
from the base of the beak to the oil-gland. I have preferred this
standard, owing to the variability of the length of tail. But I have
made similar calculations, taking as the standard the length from
tip to tip of wing, and likewise in most cases from the base of the
beak to the end of the tail; and the result has always been closely
similar. To give an example: the first bird in the table, being a
short-faced tumbler, is much smaller than the rock-pigeon, and
would naturally have shorter feet; but it is found on calculation to
have feet too short by 11 of an inch, in comparison with the feet of
the rock-pigeon, relatively to the size of the body in these two birds,
as measured from the base of beak to the oil-gland. So again,
when this same tumbler and the rock-pigeon were compared by the
length of their wings, or by the extreme length of their bodies, the
feet of the tumbler were likewise found to be too short in very
nearly the same proportion. I am well aware that the measure-

212
CHAP. V.
DOMESTIC PIGEONS.
ments pretend to greater accuracy than is possible, but it was less
trouble to write down the actual measurements given by the com-
passes in each case than an approximation.
TABLE I.
Pigeons with their beaks generally shorter than that of the Rock-
pigeon, proportionally with the size of their bodies.
Name of Breed.
Actual
length
of
Feet.
Difference between
actual and calculated
length of feet, in
proportion to length of
feet and size of body
in the Rock-pigeon.
Wild rock-pigeon (mean measure-
ment)..
2.02
Too short
by
Too long
by
. .
1:57
1.60
1.75
Short-faced Tumbler, bald-head..
almond ..
Tumbler, red magpie
red common (by standard
to end of tail)
common bald-head
roller ..
Turbit
1.85
Jacobin
Trumpeter, white
mottled..
Fantail (by standard to end of tail)..
0.11
0.16
0.19
0.07
0:18
0.06
0.17
0.01
0.15
0.02
0:06
0.18
0.15
0.15
0.0
0.19
0.03
0.02
0.16
0.03
0.0
1.85
1.80
1.5
1.80
1.84
1.90
2:02
1.95
1.85
1.95
1.95
1.80
2.10
1.82
1.65
2.00
2.00
1.90
1.90
1.85
2.00
2.42
2 30
2.17
crested var.
Indian Frill-back
English Frill back
Nun..
Laugher
Barb
0.03
Spot
0:02
.
0.07
0.18
Swallow, red
blue
Pouter
German.
Bussorah Carrier
0.03
0.11
0.09
0.09
Number of specimens ..
28
22
5

CHAP. V.
213
ON THE EFFECTS OF DISUSE.
TABLE II.
Pigeons with their beaks longer than that of the Rock-pigeon,
proportionally with the size of their bodies.
Name of Breed.
Actual
length
Difference between
actual and calculated
length of feet, in
proportion to length of
feet and size of body
in the Rock-pigeon.
of
Feet.
Wild rock-pigeon (mean measurement) 2:02
Too short Too long
by
by
Carrier ..
2.60
2.60
2.40
2.25
2.80
2.80
2.85
2.75
Dragon
Bagadotten Carrier
Scanderoon, white
Pigeon cygne
Runt
0.31
0.25
0-21
0.06
0.56
0:37
0.29
0.27
. .
Number of specimens
8.
8
In these two tables we see in the first column the actual length
of the feet in thirty-six birds belonging to various breeds, and in the.
two other columes we see by how much the feet are too short or too
long, according to the size of bird, in comparison with the rock-pi-
geon. In the first table twenty-two specimens have their feet too
short, on an average by a little above the tenth of an inch (viz. 107);
and five specimens have their feet on an average a very little too
long, namely, by :07 of an inch. But some of these latter and ex-
ceptional cases can be explained; for instance, with pouters the legs
and feet are selected for length, and thus any natural tendency to a
diminution in the length of the feet will have been counteracted.
In the swallow and barb, when the calculation was made on any
standard of comparison excepting the one above used (viz. length
of body from base of beak to oil-gland), the feet were found to be too
small.
in the second table we have eight birds, with their beaks much
longer than in the rock pigeon, both actually and proportionally with
the size of body, and their feet are in an equally marked manner
longer, namely, in proportion, on an average by .29 of an inch. I
should here state that in Table I. there are a few partial exceptions
to the beak being proportionally shorter than in the rock-pigeon:

214
CHAP. V.
DOMESTIC PIGEONS.
38
thus the beak of the English frill-back is just perceptibly longer,
and that of the Bussorah carrier of the same length or slightly longer,
than in the rock-pigeon. The beaks of spots, swallows, and laugh-
ers are only a very little shorter, or of the same proportional length,
but slenderer. Nevertheless, these two tables, taken conjointly, in-
dicate pretty plainly some kind of correlation between the length
of the beak and the size of the feet. Breeders of cattle and horses
believe that there is an analogous connection between the length of
the limbs and head; they assert that a race-horse with the head of
a dray-horse, or a greyhound with the head of a bulldog, would be
a monstrous production. As fancy pigeons are generally kept in
small aviaries, and are abundantly supplied with food, they must
walk about much less than the wild rock-pigeon; and it.may be ad-
mitted as highly probable that the reduction in the size of the feet
in the twenty-two birds in the first table has been caused by disuse,
and that this reduction has acted by correlation on the beaks of the
great majority of the birds in Table I. When, on the other hand,
the beak has been much elongated, by the continued selection of
successive slight increments of length, the feet by correlation have
likewise become much elongated in comparison with those of the
wild rock-pigeon, notwithstanding their lessened use.
As I had taken measures from the end of the middle toe to the
heel of the tarsus in the rock-pigeon and in the above thirty-six birds,
I have made calculations analogous with those above given, and the
result is the same,-namely, that in the short-beaked breeds, with
equally few exceptions as in the former case, the middle toe con-
jointly with the tarsus has decreased in length; whereas in the long-
beaked breeds it has increased in length, though not quite so uni-
formly as in the former case, for the leg in some varieties of the runt
varies much in length.
As fancy pigeons are generally confined in aviaries of moderate
size, and as even when not confined they do not search for their own
food, they must during many generations have used their wings in-
comparably less than the wild rock-pigeon. Hence it seemed to me
probable that all the parts of the skeleton subservient to flight
would be found to be reduced in size. With respect to the sternum,
I have carefully measured its extreme length in twelve birds of dif-
ferent breeds, and in two wild rock-pigeons from the Shetland Is-
lands. For the proportional comparison I have tried with all twelve
birds three standards of measurement, namely the length from the
38 In an analogous, but converse man-
ner, certain natural groups of the Colum-
bidæ, from being more terrestrial in their
habits than other allied groups, have
larger ſeet. See Prince Bonaparte's 'Coup.
d'oeil sur l'Ordre des Pigeons.'

CHAP. V.
215
ON THE EFFECTS OF DISUSE. .
base of the beak to the oil-gland, to the end of the tail, and from the
extreme tip to tip of wings. The result has been in each case nearly
the same, the sternum being invariably found to be shorter than the
wild rock-pigeon. I will give only a single table, as calculated by
the standard from the base of the beak to the oil-gland; for the re-
sult in this case is nearly the mean between the result obtained by
the two other standards.
Length of Sternum.
Name of Breed.
Actual
Too
Length. Short by
Inches.
Name of Breed.
Actual
Too
Length. Short by
Inches.
Wild Rock-pigeon
Pied Scanderoon
Bagadotten Carrier
Dragon
Carrier
Short-faced Tumbler
2.55
2.80
2.80
2.45
2.75
2:05
0.60
0:17
0.41
0.35
0.28
Barb..
Nun
German Pouter
Jacobin
English Frill-back
Swallow
2.35
2.27
2.36
2.33
2.40
2.45
0.34
0.15
0.51
0.22
0.43
0.17
This table shows that in these twelve breeds the sternum is on an
average one-third of an inch (exactly 332) shorter than in the rock-
pigeon, proportionally with the size of their bodies ; so that the
sternum has been reduced by between one-seventh and one-eighth
of its entire length; and this is a considerable reduction.
I have also measured in twenty-one birds, including the above
dozen, the prominence of the crest of the sternum relatively to its
length, independently of the size of the body. In two of the
twenty-one birds the crest was prominent in the same relative de-
gree as in the rock pigeon ; in seven it was more prominent; but in
five out of these seven, namely, in a fantail, two scanderoons, and two
English carriers, this greater prominence may to a certain extent be
explained, as a prominent breast is admired and selected by fanciers;
in the remaining twelve birds the prominence was less. Hence it
follows that the crest exhibits a slight, though uncertain, tendency
to become reduced in prominence in a greater degree than does the
length of the sternum relatively to the size of body in comparison
with the rock-pigeon.
I have measured the length of the scapula in nine different large
and small-sized breeds, and in all the scapula is proportionally
shorter taking the same standard as before) than in the wild rock-
pigeon. The reduction in length on an average is very nearly one-
fifth of an inch, or about one-ninth of the length of the scapula in
the rock-pigeon.
The arms of the furcula in all the specimens which I compared,
diverged less, proportionally with the size of body, than in the

216
CHAP. V.
DOMESTIC PIGEONS.
rock-pigeon ; and the whole furcula was proportionally shorter.
Thus in a runt, which measured from tip to tip of wings 384 inches,
the furcula was only a very little longer (with the arms hardly more
divergent) than in a rock-pigeon which measured from tip to tip 26
inches. In a barb, which in all its measurements was a little larger
than the same rock-pigeon, the furcula was a quarter of an inch
shorter. In a pouter, the furcula had not been lengthened propor-
tionally with the increased length of the body. In a short-faced
tumbler, which measured from tip to tip of wings 24 inches, there-
fore only 24 inches less than the rock-pigeon, the furcula was barely
two thirds of the length of that of the rock-pigeon.
We thus clearly see that the sternum, scapulæ, and
furcula are all reduced in proportional length; but when
we turn to the wings we find what at first appears a
wholly different and unexpected result. I may here re-
mark that I have not picked out specimens, but have
used every measurement made by me. Taking the length
from the base of beak to the end of the tail as the
standard of comparison, I find that, out of thirty-five
birds of various breeds, twenty-five have wings of
greater, and ten have them of less proportional length,
than in the rock-pigeon. But from the frequently corre-
lated length of the tail and wing-feathers, it is better to
take as the standard of comparison the length from the
base of the beak to the oil-gland; and by this standard,
out of twenty-six of the same birds which had been thus
measured, twenty-one had wings too long, and only five
had them too short. In the twenty-one birds the wings ex-
ceeded in length those of the rock-pigeon, on an average,
by 13 inch ; whilst in the five birds they were less in
length by only •8 of an inch. As I was much surprised
that the wings of closely confined birds should thus so
frequently have been increased in length, it occurred to
me that it might be solely due to the greater length of
the wing-feathers; for this certainly is the case with the
jacobin, which has wings of unusual length. As in al-
most every case I had measured the folded wings, I sub-
tracted the length of this terminal part from that of the

CHAP. V.
217
ON THE EFFECTS OF DISUSE.
expanded wings, and thus I obtained, with a moderate
degree of accuracy, the length of the wings from the
ends of the two radii, answering from wrist to wrist in
our arms. The wings, thus measured in the same twenty-
five birds, now gave a widely different result; for they
were proportionally with those of the rock-pigeon too
short in seventeen birds, and in only eight too long. Of
these eight birds, five were long-beaked, and this fact
perhaps indicates that there is some correlation between
the length of the beak and the length of the bones of the
wings, in the same manner as with the feet and tarsi.
The shortening of the humerus and radius in the seven-
teen birds may probably be attributed to disuse, as in the
case of the scapulæ and furcula to which the wing-bones
are attached ;-the lengthening of the wing-feathers, and
consequently the expansion of the wings from tip to
tip, being, on the other hand, as completely independent
of use and disuse as is the growth of the hair or wool on
our long-haired dogs or long-woolled sheep.
To sum up: we may confidently admit that the length
of the sternum, and frequently the prominence of its
crest, the length of the scapulæ and furcula, have all been
reduced in size in comparison with the same parts in the
rock-pigeon. And I presume that this may be safely at-
tributed to disuse or lessened exercise.
The wings, as
measured from the ends of the radii, have likewise been
generally reduced in length; but, owing to the increased
growth of the wing-feathers, the wings, from tip to
tip, are commonly longer than in the rock-pigeon. The
feet, as well as the tarsi conjointly with the middle toe,
have likewise in most cases become reduced ; and this it
is probable has been caused by their lessened use; but
39 It perhaps deserves notice that be-
sides these five birds two of the eight
were barbs, which, as I have shown,
must be classed in the same group with
the long-beaked carriers and runts.
Barbs may properly be called short-
beaked carriers. It would, therefore,
appear as if, during the reduction of
their beaks, their wings had retained a
little of that excess of length which is
characteristic of their nearest relations
and progenitors.
10

218
CHAP. V.
DOMESTIC PIGEONS.
the existence of some sort of correlation between the
feet and beak is shown more plainly than the effects of
disuse. We have also some faint indication of a similar
correlation between the main bones of the wing and the
beak.
Summary on the Points of Difference between the
several Domestic Races, and between the individual
Birds.—The beak, together with the bones of the face,
differ remarkably in length, breadth, shape, and curva-
ture. The skull differs in shape, and greatly in the
angle formed by the union of the premaxillary, nasal,
and maxillo-jugal bones. The curvature of the lower
jaw and the reflexion of its upper margin, as well as the
gape of the mouth, differ in a highly remarkable manner.
The tongue varies much in length, both independently
and in correlation with the length of the beak. The
development of the naked, wattled skin over the nostrils
and round the eyes varies in an extreme degree. The
eyelids and the external orifices of the nostrils vary in
length, and are to a certain extent correlated with the
degree of development of the wattle. The size and form
of the esophagus and crop, and their capacity for in-
flation, differ immensely. The length of the neck varies.
With the varying shape of the body, the breadth and
number of the ribs, the presence of processes, the number
of the sacral vertebra, and the length of the sternum,
all vary. The number and size of the coccygeal verte-
bræ vary, apparently in correlation with the increased
size of the tail. The size and shape of the perforations
in the sternum, and the size and divergence of the arms
of the furcula, differ. The oil-gland varies in develop-
ment, and is sometimes quite aborted. The direction
and length of certain feathers have been much modified,
as in the hood of the Jacobin and the frill of the Turbit.
The wing and tail feathers generally vary in length to-
gether, but sometimes independently of each other and
of the size of the body. The number and position of the

CHAP, V.
219
SUMMARY OF DIFFERENCES.
tail-feathers vary to an unparalleled degree. The primary
and secondary wing-feathers occasionally vary in num.
ber, apparently in correlation with the length of the
wing. The length of the leg and the size of the feet,
and, in connection with the latter, the number of the
scutellæ, all vary. A web of skin sometimes connects
the bases of the two inner toes, and almost invariably the
two outer toes when the feet are feathered.
The size of the body differs greatly: a runt has been
known to weigh more than five times as much as a short-
faced tumbler. The eggs differ in size and shape. Ac-
cording to Parmentier, some races use much straw in
building their nests, and others use little; but I cannot
hear of any recent corroboration of this statement. The
length of time required for hatching the eggs is uniform
in all the breeds. The period at which the characteristic
plumage of some breeds is acquired, and at which cer-
tain changes of colour supervene, differs. The degree
to which the young birds are clothed with down when
first hatched is different, and is correlated in a singular
manner with the future colour of the plumage. The
manner of flight, and certain inherited movements, such
as clapping the wings, tumbling either in the air or on
the ground, and the manner of courting the female, pre-
sent the most singular differences. In disposition the
several races differ. Some races are very silent; others
coo in a highly peculiar manner.
Although many different races have kept true in cha-
racter during several centuries, as we shall hereafter
more fully see, yet there is far more individual variability
in the truest breeds than in birds in a state of nature.
There is hardly any exception to the rule that those cha-
racters vary most which are now most valued and at-
tended to by fanciers, and which consequently are now
being improved by continued selection. This is indi-
40 Temminck, 'Hist. Nat. Gén. des Pigeons et des Gallinacés,' tom. 1., 1813, p. 170.

220
CHAP. V.
DOMESTIC PIGEONS,
rectly admitted by fanciers when they complain that it
is much more difficult to breed high fancy pigeons up to
the proper standard of excellence than the so-called
toy pigeons, which differ from each other merely in
colour; for particular colours when once acquired are
not liable to continued improvement or augmentation.
Some characters become attached, from quite unknown
causes, more strongly to the male than to the female
sex; so that we have, in certain races, a tendency to-
wards the appearance of secondary sexual characters."
of which the aboriginal rock-pigeon displays not a trace.
41 This term was used by John Hunter
for such differences in structure between
the males and females, as are not di-
rectly connected with the act of repro-
duction, as the tail of the peacock, the
horns of deer, &c.

CHAP, VI,
221
THEIR PARENTAGE.
CHAPTER VI.
9
PIGEONS-continued.
ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC
RACES HABITS OF LIFE WILD RACES OF THE ROCK-PIGEON
DOVECOT-PIGEONS - PROOFS OF THE DESCENT OF THE SEV-
ERAL 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 --SELEC-
TION - 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 PERMA-
NENT, WHILST OTHERS CHANGE — SUMMARY.
The differences described in the last chapter between
the eleven chief domestic races and between individual
birds of the same race, would be of little significance,
if they had not all descended from a single wild stock.
The question of their origin is therefore of fundamental
importance, and must be discussed at considerable length.
No one will think this superfluous who considers the
great amount of difference between the races, who knows
how ancient many of them are, and how truly they
breed at the present day. Fanciers almost unanimously
believe that the different races are descended from sev-
eral wild stocks, whereas most naturalists believe that
all are descended from the Columba livia or rock pigeon.
Temminck has well observed, and Mr. Gould has
1 Temminck, Hist. Nat. Gén, des Pigeons,' &c., tom. i. p. 191.

222
CHAP. VI.
DOMESTIC PIGEONS.
made the same remark to me, that the aboriginal parent
must have been a species which roosted and built its nest
on rocks; and I may add that it must have been a social
bird. For all the domestic races are highly social, and
none are known to build or habitually to roost on trees.
The awkward manner in which some pigeons, kept by
me in a summer-house near an old walnut-tree, occasion-
ally alighted on the barer branches, was evident. Nev-
ertheless, Mr. R. Scot Skirving informs me that he often
saw crowds of pigeons in Upper Egypt settling on the
low trees, but not on the palms, in preference to the mud
hovels of the natives. In India Mr. Blyth® has been
assured that the wild C. livia, var. intermedia, some-
times roosts in trees. I may here give a curious instance
of compulsion leading to changed habits: the banks of
the Nile above lat. 28° 30' are perpendicular for a long
distance, so that when the river is full the pigeons cannot
alight on the shore to drink, and Mr. Skirying repeat-
edly saw whole flocks settle on the water, and drink
whilst they floated down the stream. These flocks seen
from a distance resembled flocks of gulls on the surface
of the sea.
If any domestic race had descended from a species
which was not social, or which built its nest or roosted in
trees," the sharp eyes of fanciers would assuredly have de-
tected some vestige of so different an aboriginal habit.
For we have reason to believe that aboriginal habits are
long retained under domestication. Thus with the com-
2 I have heard through Sir C. Lyell
from Miss Buckley, that some half-bred
carriers kept during many years near
London regularly settled by day on some
adjoining trees, and, after being dis-
turbed in their loft by their young being
taken, roosted on them at night.
3. Annals and Mag. of Nat. Hist.,'
2nd ser., vol. xx., 1857, p. 509; and in a
late volume of the Journal of the Asiatic
Society.
4 In works written on the pigeon by
fanciers I have sometimes observed the
mistaken belief expressed that the spe-
cies which naturalists call ground-pi-
geons (in contradistinction to arboreal
pigeons) do not perch and build on
trees. In these same works wild spe-
cies resembling the chief domestic races
are often said to exist in various parts
of the world, but such species are quite
unknown to naturalists.

CHAP, VI.
223
THEIR PARENTAGE.
mon ass we see signs of its original desert life in its
strong dislike to cross the smallest stream of water, and
in its pleasure in rolling in the dust. The same strong
dislike to cross a stream is common to the camel, which
has been domesticated from a very ancient period.
Young pigs, though so tame, sometimes squat when
frightened, and thus try to conceal themselves even on an
open and bare place. Young turkeys, and occasionally
even young fowls, when the hen gives the danger-cry, run
away and try to hide themselves, like young partridges
or pheasants, in order that their mother may take flight,
of which she has lost the power. The musk-duck (Den-
drocygna viduata) in its native country often perches and
roosts on trees,' and our domesticated musk-ducks, though
such sluggish birds, “are fond of perching on the tops of
barns, walls, &c., and, if allowed to spend the night in
the hen-house, the female will generally go to roost by
the side of the hens, but the drake is too heavy to mount
thither with ease. We know that the dog, however
well and regularly fed, often buries, like the fox, any su-
perflous food; and we see him turning round and round
on a carpet, as if to trample down grass to form a bed;
we see him on bare pavements scratching backwards as
if to throw earth over his excrement, although, as I be-
lieve, this is never effected even where there is earth. In
the delight with which lambs and kids crowd together
and frisk on the smallest hillock, we see a vestige of their
former alpine habits.
We have therefore good reason to believe that all the
domestic races of the pigeon are descended either from
some one or from several species which both roosted and
built their rests on rocks, and were social in disposition.
As only five or six wild species with these habits and
making any near approach in structure to the domes-
96
6 Sir R. Schomburgk, in Journal R.
Geograph. Soc.,' vol. xiii. 1944, p. 32.
6 Rev. E. S. Dixon, 'Ornamental Poul-
try,' 1848, pp. 63, 66.

224
CHAP. VI.
DOMESTIC PIGEONS.
ticated pigeon are known to exist, I will enumerate
them.
Firstly, the Columba leuconota resembles certain domestic varie-
ties in its plumage, with the one marked and never-failing difference
of a white band which crosses the tail at some distance from the ex-
tremity. This species, moreover, inhabits the Himalaya, close to
the limit of perpetual snow; and therefore, as Mr. Blyth has re-
marked, is not likely to have been the parent of our domestic breeds,
which thrive in the hottest countries. Secondly, the C. rupestris,
of Central Asia, which is intermediate between the C. leuconota and
livia ; but has nearly the same coloured tail with the former species.
Thirdly, the Columba littoralis builds and roosts, according to Tem-
minck, on rocks in the Malayan archipelago; it is white, excepting
parts of the wing and the tip of the tail, which are black ; its legs
are livid-coloured, and this is a character not observed in any adult
domestic pigeon ; but I need not have mentioned this species or the
closely-allied C. luctuosa, as they in fact belong to the genus Carpo-
phaga. Fourthly, Columba Guinea, which ranges from Guinea 8 to
the Cape of Good Hope, and roosts either on trees or rocks, according
to the nature of the country. This species belongs to the genus
Strictonas of Reichenbach, but is closely allied to true Columba ; it
is to some extent coloured like certain domestic races, and has been
said to be domesticated in Abyssinia ; but Mr. Mansfield Parkyns,
who collected the birds of that country and knows the species, informs
me that this is a mistake. Moreover the C. Guinea is characterized
by the feathers of the neck having peculiar notched tips,-a charac-
ter not observed in any domestic race. Fifthly, the Columba anas
of Europe, which roosts on trees, and builds its nest in holes, either
in trees or the ground; this species, as far as external characters go,
might be the parent of several domestic races; but, though it crosses
readily with the true rock-pigeon, the offspring, as we shall pre-
sently see, are sterile hybrids, and of such sterility there is not a
trace when the domestic races are intercrossed. It should also be
observed that if we were to admit, against all probability, that any
of the foregoing five or six species were the parents of some of our
domestic pigeons, not the least light would be thrown on the chief
differences between the eleven most strongly-marked races.
We now come to the best known rock-pigeon, the Columba livia,
7 Proc. Zoolog. Soc., 1859, p. 400.
8 Temminck, Hist. Nat. Gén, des
Pigeons,' tom. i.; also "Les Pigeons,'
par Mad. Knip and Temminck. Bona-
parte however, in his 'Coup-d'ail,' be-
lieves that two closely allied species are
confounded together under this name.
The C. leucocephal of the West Indies
is stated by Temminck to be a rock-
pigeon; but I am informed by Mr. Gosse
that this is an error.

CHAP. VI.
225
THEIR PARENTAGE.
which is often designated in Europe pre-eminently as the Rock-pi-
geon, and which naturalists believe to be the parent of all the do-
mesticated breeds. This bird agrees in every essential character
with the breeds which have been only slightly modified. It differs
from all other species in being of a slaty-blue colour, with two black
bars on the wings, and with the croup (or loins) white. Occasion-
ally birds are seen in Faroe and the Hebrides with the black bars
replaced by two or three black spots; this form has been named by
Brehm ° C. amalice, but this species has not been admitted as distinct
by other ornithologists. Graba 10 even found a difference between
the wing-bars of the same bird in Faroe. Another and rather mor
distinct form is either truly wild or has become feral on the cliffs
of England, and was doubtfully named by Mr. Blyth" as C. affinis,
but is now no longer considered by him as a distinct species. C. af-
finis is rather smaller than the rock-pigeon of the Scottish islands,
and has a very different appearance owing to the wing-coverts being
chequered with black, with similar marks often extending over the
back. The chequering consists of a large black spot on the two
sides, but chiefly on the outer side, of each feather. The wing. bars
in the true rock-pigeon and in the chequered variety are, in fact, due
to similar though larger spots symmetrically crossing the secondary
wing-feather and the larger coverts. Hence the chequering arises
merely from an extension of these marks to other parts of the
plumage. Chequered birds are not confined to the coasts of Eng-
land; for they were found by Graba at Faroe; and W. Thompson
says that at Islay fully half the wild rock-pigeons were chequered.
Colonel King, of Hythe, stocked his dovecot with young wild
birds which he himself procured from nests at the Orkney Islands;
and several specimens, kindly sent to me by him, were all plainly
chequered. As we thus see that chequered birds occur mingled with
the true rock-pigeon at three distinct sites, namely, Faroe, the Ork-
ney Islands, and Islay, no importance can be attached to this natu-
ral variation in the plumage.
Prince C. L. Bonaparte, 13 a great divider of species, enumerates,
with a mark of interrogation, as distinct from C. livia, the C. turri-
cola of Italy, the C. rupestris of Daouria, and the C. Schimperi of
Abyssinia ; but these birds differ from C. livia in characters of the
12
Handbuch der Naturgesch. Vogel
Deutschlands.'
10 Tagebuch Reise nach Färo,' 1830,
8. 62.
ing.
12 Natural History of Ireland,' Birds,
vol.ii. (1850), p. 11. For Graba, see pre-
vious reference.
13 Coup-d'oeil sur l'Ordre des Pi-
ons,' Comptes Rendus, 1854-55.
11 Annals and Mag. of Nat. Hist.,'
vol. xix., 1847, p. 102. This excellent
paper on pigeons is well worth consult

226
CHAP, VÍ.
DOMESTIC PIGEONS.
most trifling value. In the British Museum there is a chequered
pigeon, probably the C. Schimperi of Bonaparte, from Abyssinia.
To these may be added the C.gymnocyclus of G. R. Gray from W.
Africa, which is slightly more distinct, and has rather more naked
skin round the eyes than the rock pigeon ; but from information
given me by Dr. Daniell, it is doubtful whether this is a wild
bird, for dovecot pigeons (which I have examined) are kept on the
coast of Guinea.
The wild rock-pigeon of India (C. intermedia of Strickland) has
been more generally accepted as a distinct species. It chiefly dif-
fers in the croup being blue instead of snow-white; but as Mr.
Blyth informs me, the tint varies, being sometimes albescent. When
this form is domesticated chequered birds appear, just as occurs in
Europe with the truly wild C. livia. Moreover we shall immediate-
ly have proof that the blue and white croup is a highly variable
character; and Bechstein 14 asserts that with dovecot-pigeons in
Germany this is the most variable of all the characters of the plu-
mage. Hence it may be concluded that C. intermedia cannot be
ranked as specifically distinct from C. livia.
In Madeira there is a rock-pigeon which a few ornithologists
have suspected to be distinct from C.livia. I have examined numer-
ous specimens collected by Mr. E. V. Harcourt and Mr. Mason.
They are rather smaller than the rock-pigeon from the Shetland
Islands, and their beaks are plainly thinner, but the thickness of
the beak varied in the several specimens. In plumage there is
remarkable diversity ; some specimens are identical in every feather
(I speak after actual comparison) with the rock-pigeon of the Shet-
land Islands; others are chequered, like C. affinis from the cliffs of
England, but generally to a greater degree, being almost black over
the whole back; others are identical with the so-called C. intermedia
of India in the degree of blueness of the croup; whilst others have
this part very pale or very dark blue, and are likewise chequered.
So much variability raises a strong suspicion that these birds are
domestic pigeons which have become feral.
From these facts it can hardly be doubted that C. Tiria, affinis, in-
termedia, and the forms marked with an interrogation by Bonaparte,
ought all to be included under a single species. But it is quite im-
material whether or not they are thus ranked, and whether some
one of these forms or all are the progenitors of the various domestic
kinds, as far as any light is thus thrown on the differences between
the more strongly-marked races. That common dovecot-pigeons,
14 Naturgesch, Deutschlands,' Band iv., 1795, s. 14.

CHAP. VI.
227
THEIR PARENTAGE.
which are kept in various parts of the world, are descended from
one or from several of the above-mentioned wild varieties of C. livia,
no one who compares them will doubt. But before making a few
remarks on dovecot-pigeons, it should be stated that the wild rock-
pigeon has been found easy to tame in several countries. We have
seen that Colonel King at Hythe stocked his dovecot more than
twenty years ago with young wild birds taken at the Orkney Is-
lands, and since this time they have greatly multiplied. The accu-
rate Macgillivray 15 asserts that he completely tamed a wild rock-
pigeon in the Hebrides; and several accounts are on record of these
pigeons having bred in dovecots in the Shetland Islands. In India,
as Captain Hutton informs me, the wild rock-pigeon is easily tamed,
and breeds readily with the domestic kind; and Mr. Blyth 16 asserts
that wild birds come frequently to the dovecots and mingle freely
with their inhabitants. In the ancient 'Ayeen Akbery' it is writ-
ten that, if a few wild pigeons be taken, "they are speedily joined
by a thousand others of their kind.”
Dovecot-pigeons are those which are kept in dovecots in a semi-
domesticated state; for no special care is taken of them, and they
procure their own food, except during the severest weather. In
England, and, judging from MM. Boitard and Corbie's work, in
France, the common dovecot-pigeon exactly resembles the chequered
variety of C. livia, but I have seen dovecots brought from York-
shire, without any trace of chequering, like the wild rock-pigeon of
the Shetland Islands. The chequered dovecots from the Orkney
Islands, after having been domesticated by Colonel King for more
than twenty years, differed slightly from each other in the darkness
(f their plumage, and in the thickness of their beaks; the thinnest
beak being rather thicker than the thickest one in the Madeira
birds. In Germany, according to Bechstein, the common dovecot-
pigeon is not chequered. In India they often become chequered,
and sometimes pied with white; the croup also, as I am informed
by Mr. Blyth, 'becomes nearly white. I have received from Sir J.
Brooke some dovecot-pigeons, which originally came from the S.
Natunas Islands in the Malay archipelago, and which had been
15 History of British Birds,' vol. i. pp.
275-284. Mr. Andrew Duncan tamed a
rock-pigeon in the Shetland Islands. Mr.
James Barclay, and Mr. Smith of Uyea
Sound, both say that the wild rock-pi.
geon can be easily tamed; and the for-
mer gentleman asserts that the tame
birds breed four times a year. Dr. Law-
rence Edmondstone informs me that a
wild rock-pigeon came and settled in his
dovecot in Balta Sound in the Shetland
Islands, and bred with his pigeons; he
has also given me other instances of the
wild rock-pigeon having been taken
young and breeding in captivity.
16. Annals and Mag. of Nat. History,
vol. xix., 1847, p. 103, and vol. for 1857,
p. 512.

228
CHAP, VI.
DOMESTIC PIGEONS.
crossed with the Singapore dovecots; they were small, and the dark-
est variety was extremely like the dark chequered variety with a
blue croup from Madeira; but the beak was not so thin, though
decidedly thinner than in the rock-pigeon from the Shetland Islands.
A dovecot-pigeon sent to me by Mr. Swinhoe from Foochow, in
China, was likewise rather small, but differed in no other respect.
I have also received, through the kindness of Dr. Daniell, four liv.
ing dovecot-pigeons from Sierra Leone ; 17 these were fully as large
as the Shetland rock-pigeon, with even bulkier bodies. In plumage
some of them were identical with the Shetland rock-pigeon, but
with the metallic tints apparently rather more brilliant; others had
a blue croup and resembled the chequered variety of C. intermedia
of India, and some were so much chequered as to be nearly black.
In these four birds the beak differed slightly in length, but in all it
was decidedly shorter, more massive, and stronger than in the wild
rock-pigeon from the Shetland Islands, or in the English dovecot.
When the beaks of these African pigeons were compared with the
thinnest beaks of the wild Madeira specimens, the contrast was
great; the former being fully one-third thicker in a vertical direc-
tion than the latter; so that any one at first would have felt inclined
to rank these birds as specifically distinct; yet so perfectly gradua-
ted a series could be formed between the above-mentioned varieties,
that it was obviously impossible to separate them.
To sum up: the wild Columba livia, including under
this name C. affinis, intermedia, and the other still more
closely-affined geographical races, has a vast range from
the southern coast of Norway and the Faroe Islands to
the shores of the Mediterranean, to Madeira and the Ca-
nary Islands, to Abyssinia, India, and Japan. It varies
greatly in plumage, being in many places chequered with
black, and having either a white or blue croup or loins;
it varies also slightly in the size of the beak and body.
Dovecot-pigeons, which no one disputes are descended
from one or more of the above wild forms, present a sim-
ilar but greater range of variation in plumage, in the size
of body, and in the length and thickness of the beak.
17 Domestic pigeons of the common
kind are mentioned as being pretty nu-
merous in John Barbut's Description
of the Coast of Guinea,' (9.215), publish-
ed in 1746; they are said, in accordance
with the name which they bear, to have
been imported.

CHAP. VI.
229
THEIR PARENTAGE.
There seems to be some relation between the croup being
blue or white, and the temperature of the country inhab-
ited by both wild and dovecot-pigeons; for nearly all the
dovecot-pigeons in the northern parts of Europe have a
white croup, like that of the wild European rock-pigeon ;
and nearly all the dovecot-pigeons of India have a blue
croup like that of the wild C. intermedia of India. As
in various countries the wild rock-pigeon has been found
easy to tame, it seems extremely probable that the dove-
cot-pigeons throughout the world are the descendants of
at least two and perhaps more wild stocks, but these, as
we have just seen, cannot be ranked as specifically distinct.
With respect to the variation of C. livia, we may
without fear of contradiction go one step further. Those
pigeon-fanciers who believe that all the chief races, such
as Carriers, Pouters, Fantails, &c., are descended from
distinct aboriginal stocks, yet admit that the so-called
toy-pigeons, which differ from the rock-pigeon in little
except in colour, are descended from this bird. By toy-
pigeons are meant such birds as Spots, Nuns, Helmets,
Swallows, Priests, Monks, Porcelains, Swabians, Arch-
angels, Breasts, Shields, and others in Europe, and many
others in India. It would indeed be as puerile to sup-
pose that all these birds are descended frorn so many
distinct wild stocks as to suppose this to be the case
with the many varieties of the gooseberry, heartsease,
or dahlia. Yet these pigeons all breed true, and many
of them present sub-varieties which likewise truly trans-
mit their character. They differ greatly from each
other and from the rock-pigeon in plumage, slightly
in size and proportions of body, in size of feet, and
in the length and thickness of their beaks. They differ
from each other in these respects more than do dovecot-
pigeons. Although we may safely admit that the latter,
which vary slightly, and that the toy-pigeons, which vary
in a greater degree in accordance with their more highly-
domesticated condition, are descended from C. livia, in-

230
CHAP, VÌ.
DOMESTIC PIGEONS.
cluding under this name the above-enumerated wild geo-
graphical races; yet the question becomes far more
difficult when we consider the eleven principal races,
most of which have been so profoundly modified. It can,
however, be shown, by indirect evidence of a perfectly
conclusive nature, that these principal races are not de-
scended from so many wild stocks; and if this be once
admitted, few will dispute that they are the descendants
of C. livia, which agrees with them so closely in habits
and in most characters, which varies in a state of nature,
and which has certainly undergone a considerable amount
of variation, as in the toy-pigeons. We shall moreover
presently see how eminently favourable circumstances
have been for a great amount of modification in the more
carefully tended breeds.
The reasons for concluding that the several principal
races have not descended from so many aboriginal and
unknown stocks
may be grouped under the following six
heads :
-Firstly, if the eleven chief races have not arisen
from the variation of some one species, together with its
geographical races, they must be descended from several
extremely distinct aboriginal species; for no amount of
crossing between only six or seven wild forms could pro-
duce races so distinct as pouters, carriers, runts, fantails,
turbits, short-faced tumblers, jacobines, and trumpeters.
How could crossing produce, for instance, a pouter or a
fantail, unless the two supposed aboriginal parents pos-
sessed the remarkable characters of these breeds? I
am aware that some naturalists, following Pallas, believe
that crossing gives a strong tendency to variation, inde-
pendently of the characters inherited from either parent.
They believe that it would be easier to raise a pouter or
fantail pigeon from crossing two distinct species, neither
of which possessed the characters of these races, than
from any single species. I can find few facts in support
of this doctrine, and believe in it only to a limited degree;
but in a future chapter I shall have to recur to this sub-

CHAP. VI.
231
THEIR PARENTAGE.
ject. For our present purpose the point is not material.
The question which concerns us is, whether or not many
new and important characters have arisen since man first
domesticated the pigeon. On the ordinary view, varia-
bility is due to changed conditions of life; on the Palla-
sian doctrine, variability, or the appearance of new cha-
racters, is due to some mysterious effect from the crossing
of two species, neither of which possess the characters in
question. In some few instances it is credible, though
for several reasons not probable, that well-marked races
have been formed by crossing; for instance, a barb might
perhaps have been formed by a cross between a long-beak-
ed carrier, having large eye-wattles, and some short-beak-
ed pigeon. That many races have been in some degree
modified by crossing, and that certain varieties which are
distinguished only by peculiar tints have arisen from
crosses between differently-coloured varieties, may be ad-
mitted as almost certain. On the doctrine, therefore,
that the chief races owe their differences to their descent
from distinct species, we must admit that at least eight
or nine, or more probably a dozen species, all having the
same habit of breeding and roosting on rocks and living
in society, either now exist somewhere, or formerly exist-
ed but have become extinct as wild birds. Considering
how carefully wild pigeons have been collected through-
out the world, and what conspicuous birds they are, es-
pecially when frequenting rocks, it is extremely impro-
bable that eight or nine species, which were long ago
domesticated and therefore must have inhabited some
anciently known country, should still exist in the wild
state and be unknown to ornithologists.
The hypothesis that such species formerly existed, but
have become extinct, is in some slight degree more pro-
bable. But the extinction of so many species within the
historical period is a bold hypothesis, seeing how little
influence man has had in exterminating the common
rock-pigeon, which agrees in all its habits of life with the

232
CHAP. VÌ.
DOMESTIC PIGEONS.
domestic races. The C. livia now exists and flourishes
on the small northern islands of Faroe, on many islands
off the coast of Scotland, on Sardinia and the shores of
the Mediterranean, and in the centre of India. Fanciers
have sometimes imagined that the several supposed pa-
rent-species were originally confined to small islands,
and thus might readily have been exterminated; but the
facts just given do not favour the probability of their
extinction, even on small islands. Nor is it probable,
from what is known of the distribution of birds, that
the islands near Europe should have been inhabited
by peculiar species of pigeons; and if we assume that
distant oceanic islands were the homes of the supposed
parent-species, we must remember that ancient voyages
were tediously slow, and that ships were then ill-pro-
vided with fresh food, so that it would not have been
easy to bring home living birds. I have said ancient
voyages, for nearly all the races of the pigeon were
known before the year 1600, so that the supposed wild
species must have been captured and domesticated before
that date.
Secondly.-The doctrine that the chief domestic races
have descended from several aboriginal species, implies
that several species were formerly so thoroughly domes-
ticated as to breed readily when confined. Although it
is easy to tame most wild birds, experience shows us
that it is difficult to get them to breed freely under con-
finement; although it must be owned that this is less
difficult with pigeons than with most other birds. Dur-
ing the last two or three hundred years, many birds have
been kept in aviaries, but hardly one has been added
to our list of thoroughly reclaimed species; yet on the
above doctrine we must admit that in ancient times
nearly a dozen kinds of pigeons, now unknown in the
wild state, were thoroughly domesticated.
Thirdly.-Most of our domesticated animals have run
wild in various parts of the world; but birds, owing

CHAP. VI.
233
THEIR PARENTAGE.
apparently to their partial loss of the power of flight,
less often than quadrupeds. Nevertheless I have met
with accounts showing that the common fowl has become
feral in South America and perhaps in West Africa, and
on several islands: the turkey was at one time almost
feral on the banks of the Parana ; and the Guinea-fowl
has become perfectly wild at Ascension and in Jamaica.
In this latter island the peacock, also, “has become a
maroon bird." The common duck wanders from its
home and becomes almost wild in Norfolk. Hybrids
between the common and musk-duck which have become
wild have been shot in North America, Belgium, and
near the Caspian Sea. The goose is said to have run
wild in La Plata. The common dovecot-pigeon has be-
come wild at Juan Fernandez, Norfolk Island, Ascension,
probably at Madeira, on the shores of Scotland, and, as
is asserted, on the banks of the Hudson in North Ame-
rica.18 But how different is the case, when we turn to
the eleven chief domestic races of the pigeon, which are
supposed by some authors to be descended from so many
distinct species ! no one has ever pretended that any one
of these races has been found wild in any quarter of the
world; yet they have been transported to all countries,
and some of them must have been carried back to their
native homes. On the view that all the races are the
18 With respect to feral pigeons-for American Ornithology,' and Selys-
Juan Fernandez, see Bertero in Annal. Longchamp's 'Hybrides dans la Famille
des Sc. Nat.,' tom xxi. p. 351. For Nor- des Anatides. For the goose, Isidore
folk Island, sce Rev. E. S. Dixon in the Geoffroy St. Hilaire, Hist. Nat. Gén.,'
Dovecote,' 1851, p. 14, on the authority tom. iii. p. 498. For guinea-fowls, see
of Mr. Gould, For Ascension I rely on Gosse's 'Naturalist's Sojourn in Jamai-
MS. information given me by Mr. Layard. ca,' p. 124; and his 'Birds of Jamaica'
For the banks of the Hudson, see Blyth for fuller particulars. I say the wild
in 'Annals of Nat. Hist.,' vol. xx., 1857, Guinea-fowl in Ascension. For the pea-
p. 511. For Scotland, see Macgillivray, cock, see 'A Week at Port Royal,' by a
British Birds, vol. i. p. 275; also competent authority, Mr. R. Hill, p. 42.
Thompson's Nat. History of Ireland, For the turkey I rely on oral informa-
Birds,' vol. ii. p. 11. For ducks, see Rev. tion; I ascertained that they were not
E. S. Dixon, Ornamental Poultry,' 1817, Curassows. With respect to fowls I
For the feral hybrids of the will give the references in the next
common and musk-ducks, see Audubon's chapter.
p. 122.

234
CHAP, VỊ.
DOMESTIC PIGEONS. .
product of variation, we can understand why they have
not become feral, for the great amount of modification
which they have undergone shows how long and how
thoroughly they have been domesticated; and this would
unfit them for a wild life.
Fourthly.—If it be assumed that the characteristic dif-
ferences between the various domestic races are due to
descent from several aboriginal species, we must con-
clude that man chose for domestication in ancient times,
either intentionally or by chance, a most abnormal set
of pigeons; for that species resembling such birds as
pouters, fantails, carriers, barbs, short-faced tumblers,
turbits, &c., would be in the highest degree abnormal,
as compared with all the existing members of the great
pigeon-family, cannot be doubted. Thus we should have
to believe that man not only formerly succeeded in
thoroughly domesticating several highly abnormal spe-
cies, but that these same species have since all become
extinct, or are at least now unknown. This double acci-
dent is so extremely improbable that the assumed exist-
ence of so many abnormal species would require to be
supported by the strongest evidence. On the other hand,
if all the races are descended from C. livia, we can un-
derstand, as will hereafter be more fully explained, how
any slight deviation in structure which first appeared
would continually be augmented by the preservation of
the most strongly marked individuals; and as the power
of selection would be applied according to man's fancy,
and not for the bird's own good, the accumulated amount
of deviation would certainly be of an abnormal nature in
comparison with the structure of pigeons living in a
state of nature.
I have already alluded to the remarkable fact, that the
characteristic differences between the chief domestic
races are eminently variable: we see this plainly in the
great difference in the number of the tail-feathers in the
fantail, in the development of the crop in pouters, in the

CHAP. VI,
235
THEIR PARENTAGE.
length of the beak in tumblers, in the state of the wattle
in carriers, &c. If these characters are the result of suc-
cessive variations added together by selection, we can
understand why they should be so variable: for these
are the very parts which have varied since the domesti-
cation of the pigeon, and therefore would be likely still
to vary; these variations moreover have been recently,
and are still being accumulated by man's selection;
therefore they have not as yet become firmly fixed.
Fifthly. All the domestic races pair readily together,
and, what is equally important, their mongrel offspring
are perfectly fertile. To ascertain this fact I made many
experiments, which are given in the note below; and
recently Mr. Tegetmeier has made similar experiments
with the same result. 19 The accurate Neumeister
serts that when dovecots are crossed with pigeons of any
20
as-
19 I have drawn out a long table of
the various crosses made by fanciers
between the several domestic breeds,
but I do not think it worth pub-
lishing. I have myself made for this
special purpose many crosses, and all
were perfectly fertile. I have united
in one bird five of the most distinct
races, and with patience I might un-
doubtedly have thus united all. The
case of five distinct breeds being blended
together with unimpaired fertility is
important, because Gärtner has shown
that it is a very general, though not,
as he thought, universal rule, that com-
plex crosses between several species are
excessively sterile. I have met with only
two or three cases of reported sterility
in the offspring of certain races when
crossed. Von Pistor (Das Ganze
der Feld-taubenzucht,' 1831, S. 15)
asserts that the mongrels from barbs
and fantails are sterile: I have proved
this to be erroneous, not only by cross-
ing these hybrids with several other
hybrids of the same parentage, but by
the more severe test of pairing brother
and sister hybrids inter se, and they
were perfectly fertile. Temminck has
stated (Hist. Nat. Gén. des Pigeons,'
tom. i. p. 197) that the turbit or owl
will not cross readily with other
breeds: but my turbits crossed, when
left free, with almond tumblers and
with trumpeters; the same thing has
occurred (Rev. E. S. Dixon, The
Dovecot,' p. 107) between turbits
and dovecots and nuns. I have
crossed turbits with barbs, as has M.
Boitard (p. 31), who says the hybrids
were very fertile. Hybrids from a turbit
and fantail have been known to breed
inter se (Riedel, Taubenzucht, s. 25,
and Bechstein, Naturgesch. Deutsch.'
B. iv. s. 44). Turbits (Riedel, s. 26)
have been crossed with pouters and with
jacobins, and with a hybrid jacobin-
trumpeter (Riedel, s. 27). The latter
author has, however, made some vague
statements (s. 22) on the sterility of
turbits when crossed with certain other
crossed breeds. But I have little doubt
that the Rev. E. S. Dixon's explanation
of such statements is correct, viz. that
individual birds both with turbits and
other breeds are occasionalls sterile.
20 Das Ganze der Taubenzucht,'
s. 18.

236
CHAP. VI.
DOMESTIC PIGEONS.
other breed, the mongrels are extremely fertile and
hardy. MM. Boitard and Corbié - affirm, after their great
experience, that with crossed pigeons the more distinct
the breeds, the more productive are their mongrel off-
spring. I admit that the doctrine first broached by
Pallas is highly probable, if not actually proved, namely,
that closely allied species, which in a state of nature or
when first captured would have been in some degree
sterile when crossed, lose this sterility after a long course
of domestication; yet when we consider the great differ-
ence between such races as pouters, carriers, runts, fan-
tails, turbits, tumblers, &c., the fact of their perfect, or
even increased, fertility when intercrossed in the most
complicated manner becomes a strong argument in favour
of their having all descended from a single species. This
argument is rendered much stronger when we hear (I
append in a note 22 all the cases which I have collected)
21 "Les Pigeons,' &c., p. 35.
22 Domestic pigeons pair readily with
the allied C. oenas (Bechstein, Natur-
gesch. Deutschlands,' B. iv. s. 3); and
Mr. Brent has made the same cross seve-
ral times in England, but the young were
very apt to die at about ten days old;
one hybrid which he reared (from C.
oenas and a male Antwerp carrier) pair-
ed with a dragon, but never laid eggs.
Bechstein further states (s. 26) that the
domestic pigeon will cross with C. pa-
lumbus, Turtur risoria, and T. vulga-
ris, but nothing is said of the fertility of
the hybrids, and this would have been
mentioned had the fact been ascertained.
In the Zoological Gardens (MS. report to
me from Mr. James Hunt) a male hybrid
from Turtur vulgaris and a domestic
pigeon "paired with several different
species of pigeons and doves, but none
of the eggs were good." Hybrids from
C. oenas and gymnophthalmos were
sterile. In Loudon's Mag. of Nat. Hist.'
vol. vii. 1834, p. 154, it is said that a
male hybrid (from Turtur vulgaris
male, and the cream-coloured T. risoria
female) paired during two years with a fe-
male T. risoria, and the latter laid many
eggs, but all were sterile." MM. Boitard
and Corbié ("Les Pigeons,' p. 235) state
that the hybrids from these two turtle-
doves are invariably sterile both inter
se and with either pure parent. The ex-
periment was tried by M. Corbié "avec
une espèce d'obstination;" and likewise
by M. Manduyt, and by M. Vieillot. Tem-
minck also found the hybrids from these
two species quite barren. Therefore,
when Bechstein (Naturgesch. Vogel.
Deutschlands,' B. 4, s. 101) asserts that
the hybrids from these two turtle doves
propagate inter se equally well with pure
species, and when a writer in the 'Field'
newspaper (in a letter dated Nov. 10th,
1858) makes a similar assertion, it would
appear that there must be some mistake;
though what the mistake is I know not,
as Bechstein at least must have known
the white variety of T. risoria: it would
be an unparalleled fact if the same two
species sometimes produced extremely
fertile, and sometimes extremely barren,
offspring. In the MS. report from the
Zoological Gardens it is said that hybrids
from Turtur vulgaris and suratensis,

CHAP. VI.
237
THEIR PARENTAGE.
that hardly a single well-ascertained instance is known
of hybrids between two true species of pigeons being fer-
tile, inter se, or even when crossed with one of their
pure
parents.
Sixthly.--Excluding certain important characteristic
differences, the chief races agree most closely both with
each other and with C. livia in all other respects. As
previously observed, all are eminently sociable; all dis-
like to perch or roost, and refuse to build in trees; all
lay two eggs, and this is not a universal rule with the
Columbida; all, as far as I can hear, require the same
time for hatching their eggs; all can endure the same
great range of climate; all prefer the same food, and are
passionately fond of salt; all exhibit (with the asserted
exception of the finnikin and turner, which do not differ
much in any other character) the same peculiar gestures
when courting the females; and all (with the exception of
trumpeters and laughers, which likewise do not differ much
in any other character) coo in the same peculiar manner,
unlike the voice of any other wild pigeon. All the co-
loured breeds display the same peculiar metallic tints on
the breast, a character far from general with pigeons.
Each race presents nearly the same range of variation in
colour; and in most of the races we have the same singu-
lar correlation between the development of down in the
young and the future colour of plumage. All have the
proportional length of their toes, and of their primary
wing-feathers, nearly the same, - characters which are
apt to differ in the several members of the Columbidæ.
and from T. vulgaris and Ectopistes mi-
gratorius, were sterile. Two of the latter
male hybrids paired with their pure pa-
rents, viz. Turtur vulgaris and the
Ectopistes, and likewise with T. risoria
and with Columba oenas, and many eggs
were produced, but all were barren.
At Paris, hybrids have been raised (Isid.
Geoffroy Saint Hilaire, Hist. Nat. Géné-
rale,' tom. iii. p. 180) from Turtur auri-
tus with T. cambayensis and with T.
suratensis, but nothing is said of their
fertility. At the Zoological Gardens of
London the Goura coronata and victo-
rice produced a hybrid, which paired
with the pure G.coronata, and laid seve-
ral eggs, but these proved barren.
In
1860 Columba gymnophthalmos and
maculosa produced hybrids in these
same gardens.

238
CHAP. VI.
DOMESTIC PIGEONS.
In those races which present some remarkable deviation
of structure, such as the tail of fantails, crop of pouters,
beak of carriers and tumblers, &c., the other parts re-
main nearly unaltered. Now every naturalist will admit
that it would be scarcely possible to pick out a dozen
natural species in any Family, which should agree closely
in habits and in general structure, and yet should differ
greatly in a few characters alone. This fact is explicable
through the doctrine of natural selection; for each suc-
cessive modification of structure in each natural species
is preserved, solely because it is of service; and such
modifications when largely accumulated imply a great
change in the habits of life, and this will almost certainly
lead to other changes of structure throughout the whole
organisation. On the other hand, if the several races of
the pigeon have been produced by man through selection
and variation, we can readily understand how it is that
they should still all resemble each other in habits and in
those many characters which man has not cared to modi-
fy, whilst they differ to so prodigious a degree in those
parts which have struck his eye or pleased his fancy.
Besides the points above enumerated, in which all the
domestic races resemble C. livia and each other, there is
one which deserves special notice. The wild rock-pigeon
is of a slaty-blue colour; the wings are crossed by two
black bars; the croup varies in colour, being generally
white in the pigeon of Europe, and blue in that of India ;
the tail has a black bar close to the end, and the outer
webs of the outer tail-feathers are edged with white, ex-
cept near the tips. These combined characters are not
found in any wild pigeon besides C. livia. I have looked
carefully through the great collection of pigeons in the
British Museum, and I find that a dark bar at the end
of the tail is common; that the white edging to the outer
tail-feathers is not rare; but that the white croup is ex-
tremely rare, and the two black bars on the wings occur
in no other pigeon, excepting the alpine C. leuconota and

CHAP. VI.
239
THEIR REVERSION IN COLOUR.
C. rupestris of Asia. Now if we turn to the domestic
races, it is highly remarkable, as an eminent fancier, Mr.
Wicking, observed to me, that, whenever a blue bird ap-
pears in any race, the wings almost invariably show the
double black bars.23 The primary wing-feathers may be
white or black, and the whole body may be of any colour,
but if the wing-coverts alone are blue, the two black bars
surely appear. I have myself seen, or acquired trust-
worthy evidence, as given below, 24 of blue birds with
black bars on the wing, with the croup either white or
very pale or dark blue, with the tail having a terminal
black bar, and with the outer feathers externally edged
with white or very pale coloured, in the following races,
sures
23 There is one exception to the rule,
namely in a sub-variety of the swallow
of German origin, which is figured by
Neumeister, and was shown to me by
Mr. Wicking. This bird is blue, but
has not the black wing-bars; for our
object, however, in tracing the descent
of the chief races, this exception signi-
fies the less as the swallow approaches
closely in structure to c. livia. In
many sub-varieties, the black bars are
replaced by bars of various colours. The
figures given by Neumeister are sufficient
to show that, if the wings alone are blue,
the black wing-bars appear.
24 I have observed blue birds with all
the above mentioned marks in the fol-
lowing races, which seemed to be per-
fectly pure, and were shown at various
exhibitions. Pouters, with the double
black wing-bars, with white croup, dark
bar to end of tail, and white edging to
outer tail-feathers. Turbits, with all
these same characters. Fantails, with the
same; but the group in some was bluish
or pure blue: Mr. Wicking bred blue
fantails from two black birds. Carriers
(including the Bagadotten of Neumei-
ster), with all the marks: two birds
which I examined had white, and two
had blue croups; the white edging to
the outer tail-feathers was not present
in all. Mr. Corker, a great breeder, as-
me that, if black carriers are
matched for many successive genera-
tions, the offspring become first ash-
coloured, and then blue with black
wing-bars. Runts of the elongated
breed had the same marks, but the
croup was pale blue; the outer tail-
feathers had white edges. Neumeister
figures the great Florence Runt of a blue
colour with black bars. Jacobins are
very rarely blue, but I have received
authentic accounts of at least two in-
stances of the blue variety with black
bars having appeared in England: blue
jacobins were bred by Mr. Brent from
two black birds. I have seen common
tumblers, both Indian and English, and
short-faced tumblers, of a blue colour,
with black wing-bars, with the black
bar at the end of the tail, and with the
outer tail-feathers edged with white;
the croup in all was blue, or extremely
pale blue, never absolutely white. Blue
barbs and trumpeters seem to be ex-
cessively rare; but Neumeister, who
may be implicitly trusted, figures blue
varieties of both, with black wing-bars.
Mr. Brent informs me that he has seen
a blue barb; and Mr. H. Weir, as I am
informed by Mr. Tegetmeier, once bred
a silver (which means very pale blue)
barb from two yellow birds.

240
CHAP. VI.
DOMESTIC PIGEONS.
which, as I carefully observed in each case, appeared to
be perfectly pure: namely, in Pouters, Fantails, Tumblers,
Jacobins, Turbits, Barbs, Carriers, Runts of three dis-
tinct varieties, Trumpeters, Swallows, and in many
other
toy-pigeons, which, as being closely allied to C. livia, are
not worth enumerating. Thus we see that, in purely-
bred races of every kind known in Europe, blue birds oc-
casionally appear having all the marks which charac-
terise C. livia, and which concur in no other wild species.
Mr. Blyth, also, has made the same observation with re-
spect to the various domestic races known in India.
Certain variations in the plumage are equally common
in the wild C. livia, in dovecot-pigeons, and in all the
most highly modified races. Thus, in all, the croup varies
from white to blue, being most frequently white in Eu-
rope, and very generally blue in India.25 We have seen
that the wild C. livia in Europe, and dovecots in all parts
of the world, often have the upper wing-coverts chequered
with black; and all the most distinct races, when blue,
are occasionally chequered in precisely the same manner.
Thus I have seen Pouters, Fantails, Carriers, Turbits,
Tumblers (Indian and English), Swallows, Bald-pates,
and other toy-pigeons blue and chequered; and Mr. Es-
quilant has seen a chequered Runt. I bred from two
pure
blue Tumblers a chequered bird.
The facts hitherto given refer to the occasional appear-
ance in pure races of blue birds with black wing-bars,
and likewise of blue and chequered birds; but it will
now be seen that when two birds belonging to distinct
races are crossed, neither of which have, nor probably
25 Mr. Blyth informs me that all the
domestic races in India have the croup
blue; but this is not invariable, for I
possess a very pale blue Simmali pigeon
with the croup perfectly wbite, sent to
me by Sir W. Elliot from Madras. A
slaty-blue and chequered Nakshi pigeon
has some white feathers on the croup
alone. In some other Indian pigeons
there were a few white feathers confined
to the croup, and I have noticed the same
fact in a carrier from Persia. The Java
fantail (imported into Amoy, and thence
sent me) has a perfectly white croup.

CHAP. VI,
241
THEIR REVERSION IN COLOUR.
.
have had during many generations, a trace of blue in
their plumage, or a trace of wing-bars and the other cha-
racteristic marks, they very frequently produce mongrel
offspring of a blue colour, sometimes chequered, with
black wing-bars, &c.; or if not of a blue colour, yet with
the several characteristic marks more or less plainly de-
veloped. I was led to investigate this subject from MM.
Boitard and Corbié 24 having asserted that from crosses
between certain breeds it is rare to get anything but bi-
sets or dovecot-pigeons, which, as we know, are blue birds
with the usual characteristic marks. We shall hereafter
see that this subject possesses, independently of our pre-
sent object, considerable interest, so that I will give the
results of my own trials in full. I selected for experiment
races which, when pure, very seldom produce birds of a
blue colour, or have bars on their wings and tail.
The nun is white, with the head, tail, and primary
wing-feathers black; it is a breed which was established
as long ago as the year 1600. I crossed a male nun with
a female red common tumbler, which latter variety gen-
erally breeds true. Thus neither parent had a trace of
blue in the plumage, or of bars on the wing and tail. I
should premise that common tumblers are rarely blue in
England. From the above cross I reared several young:
one was red over the whole back, but with the tail as
blue as that of the rock-pigeon; the terminal bar, how.
ever, was absent, but the outer feathers were edged with
white: a second and third nearly resembled the first, but
the tail in both presented a trace of the bar at the end: a
fourth was brownish, and the wings showed a trace of
the double bar: a fifth was pale blue over the whole
breast, back, croup, and tail, but the neck and primary
wing-feathers were reddish; the wings presented two dis-
tinct bars of a red colour; the tail was not barred, but
the outer feathers were edged with white. I crossed this
26 Les Pigeons,' &c., p. 37.
11

242
CHAP. VI.
DOMESTIC PIGEONS.
last curiously colored bird with a black mongrel of com-
plicated descent, namely, from a black barb, a spot, and
almond tumbler, so that the two young birds produced
from this cross included the blood of five varieties, none
of which had a trace of blue, or of wing and tail bars :
one of the two young birds was brownish-black, with
black wing-bars; the other was reddish-dun, with red-
dish wing-bars, paler than the rest of the body, with the
croup pale blue, the tail bluish, with a trace of the termi-
nal bar.
Mr. Eaton 27 matched two short-faced tumblers, name-
ly, a splash cock and kite hen (neither of which are blue
or barred), and from the first nest he got a perfect blue
bird, and from the second a silver or pale blue bird, both
of which, in accordance with all analogy, no doubt pre-
sented the usual characteristic marks.
I crossed two male black barbs with two female red
spots. These latter have the whole body and wings
white, with a spot on the forehead, the tail and tail-cov-
erts red; the race existed as least as long ago as 16976,
and now breeds perfectly true, as was known to be the
case in the year 1735.28 Barbs alle uniformly-colored
birds, with rarely even a trace of bars on the wing or
tail; they are known to breed very true. The mongrels
thus raised were black or nearly black, or dark or pale
brown, sometimes slightly piebald with white: of these
birds no less than six presented double wing-bars; in
two the bars were conspicuous and quite black; in seven
some white feathers appeared on the croup; and in two
or three there was a trace of the terminal bar to the tail,
but in none were the outer tail-feathers edged with white.
I crossed black barbs (of two excellent strains) with
purely-bred, snow-white fantails. The mongrels were
generally quite black, with a few of the primary wing
27 Treatise on Pigeons,'1858p. 145.
28 J. Moore's Columbarium,' 1735, in J. M. Eaton's edition, 1852, p. 71,

CHAP. VI.
243
THEIR REVERSION IN COLOUR.
and tail-feathers white: others were dark reddish-brown,
and others snow-white: none had a trace of wing-bars
or of the white croup. I then paired together two of
these mongrels, namely, a brown and black bird, and
their offspring displayed wing-bars, faint, but of a darker
brown than the rest of body. In a second brood from
the same parents a brown bird was produced, with seve-
ral white feathers confined to the croup. .
I crossed a male dun dragon belonging to a family
which had been dun-coloured without wing-bars during
several generations, with a uniform red barb (bred from
two black barbs); and the offspring presented decided
but faint traces of wing-bars. I crossed a uniform red
male runt with a white trumpeter; and the offspring
had a slaty-blue tail, with a bar at the end, and with the
outer feathers edged with white. I also crossed a female
black and white chequered trumpeter (of a different
strain from the last) with a male almond-tumbler, neither
of which exhibited a trace of blue, or of the white croup,
or of the bar at end of tail: nor is it probable that the
progenitors of these two birds had for many generations
exhibited
any
of these characters, for I have never even
heard of a blue trumpeter in this country, and my al-
mond-tumbler was purely bred; yet the tail of this
mongrel was bluish, with a broad black bar at the end,
and the croup was perfectly white. It
It may be observed
in several of these cases, that the tail first shows a ten-
dency to become by reversion blue; and this fact of the
persistency of colour in the tail and tail-coverts will
surprise no one who has attended to the crossing of
pigeons.
29 I could give numerous examples;
two will suffice. A mongrel, whose four
grandparents were a white turbit, white
trumpeter, white fantail, and blue pouter,
was white all over, except a very few
feathers about the head and on the
wings, but the whole tail and tail-coverts
were dark bluish-grey. Another mon-
grel, whose four grandparents were a
red runt, white trumpeter, white fantail,
and the same blue pouter, was pure
white all over, except the tail and upper
tail-coverts, which were pale fawn, and
except the faintest trace of double wing.
bars of the same pale fawn tint,

244
CHAP. VI.
DOMESTIC PIGEONS.
The last case which I will give is the most curious. I
paired a mongrel female barb-fantail with a mongrel
male barb-spot; neither of which mongrels had the
least blue about them. Let it be remembered that blue
barbs are excessively rare; that spots, as has been
already stated, were perfectly characterized in the year
1676, and breed perfectly true; this likewise is the case
with white fantails, so much so that I have never heard
of white fantails throwing any other colour. Neverthe-
less the offspring from the above two mongrels was of
exactly the same blue tint as that of the wild rock-pigeon
from the Shetland Islands over the whole back and wings;
the double black wing-bars were equally conspicuous;
the tail was exactly alike in all its characters, and the
croup was pure white; the head, however, was tinted
with a shade of red, evidently derived from the spot, and
was of a paler blue than in the rock-pigeon, as was the
stomach. So that two black barbs, a red spot, and a
white fantail, as the four purely-bred grandparents, pro-
duced a bird of the same general blue colour, together
with every characteristic mark, as in the wild Columba
livia.
.
With respect to crossed breeds frequently producing
blue birds chequered with black, and resembling in all
respects both the dovecot-pigeon and the chequered
wild variety of the rock-pigeon, the statement before
referred to by MM. Boitard and Corbié would almost
suffice; but I will give three instances of the appearance
of such birds from crosses in which one alone of the pa-
rents or great-grandparents was blue, but not chequered.
I crossed a male blue turbit with a snow-white trumpet-
er, and the following year with a dark, leaden-brown,
short-faced tumbler; the offspring from the first cross
were as perfectly chequered as any dovecot-pigeon; and
from the second, so much so as to be nearly as black as
the most darkly chequered rock-pigeon from Madeira.
Another bird, whose great-grandparents were a white

CHAP, VI.
245
THEIR REVERSION IN COLOUR.
If we
trumpeter, a white fantail, a white red-spot, a red runt,
and a blue pouter, was slaty-blue and chequered exactly
like a dovecot-pigeon. I may here add a remark made to
me by Mr. Wicking, who has had more experience than
any other person in England in breeding pigeons of
various colours: namely, that when a blue, or a blue and
chequered bird, having black wing-bars, once appears in
any race and is allowed to breed, these characters are so
strongly transmitted that it is extremely difficult to
eradicate them.
What, then, are we to conclude from this tendency in
all the chief domestic races, both when purely bred and
more especially when intercrossed, to produce offspring
of a blue colour, with the same characteristic marks, vary-
ing in the same manner, as in Columba livia?
admit that these races have all descended from C. livia,
no breeder will doubt that the occasional appearance of
blue birds thus characterised is accounted for on the well-
known principle of “throwing back” or reversion. Why
crossing should give so strong a tendency to reversion,
we do not with certainty know; but abundant evidence
of this fact will be given in the following chapters. It is
probable that I might have bred even for a century pure
black barbs, spots, nuns, white fantails, trumpeters, &c.,
without obtaining a single blue or barred bird; yet by
crossing these breeds I reared in the first and second
gen-
eration, during the course of only three or four years, a
considerable number of young birds, more or less plainly
coloured blue, and with most of the characteristic marks.
When black and white, or black and red birds, are crossed,
it would appear that a slight tendency exists in both pa-
rents to produce blue offspring, and that this, when com-
bined, overpowers the separate tendency in either parent
to produce black, or white, or red offspring.
If we reject the belief that all the races of the pigeon
are the modified descendants of C. livia, and suppose
that
they are descended from several aboriginal stocks, then

246
CAAP. VI.
DOMESTIC PIGEONS.
we must choose between the three following assumptions:
firstly, that at least eight or nine species formerly existed
which were aboriginally coloured in various ways, but
have since varied in so exactly the same manner as to
assume the colouring of C. livia ; but this assumption
throws not the least light on the appearance of such col-
ours and marks when the races are crossed. Or secondly,
we may assume that the aboriginal species were all col-
oured blue, and had the wing-bars and other characteristic
marks of C. livia,-a supposition which is highly im-
probable, as besides this one species no existing member
of the Columbidæ presents these combined characters;
and it would not be possible to find any other instance of
several species identical in plumage, yet as different in
important points of structure as are pouters, fantails, car-
riers, tumblers, &c. Or lastly, we may assume that all
the races, whether descended from C.livia or from several
aboriginal species, although they have been bred with so
much care and are so highly valued by fanciers, have all
been crossed within a dozen or score of generations with
C. livia, and have thus acquired their tendency to produce
blue birds with the several characteristic marks. I have
said that it must be assumed that each race has been cross-
ed with C. livia within a dozen, or, at the utmost, within
a score of generations; for there is no reason to believe
that crossed offspring ever revert to one of their ances-
tors when removed by a greater number of generations.
In a breed which has been crossed only once, the tendency
to reversion will naturally become less and less in the suc-
ceeding generations, as in each there will be less and less
of the blood of the foreign breed; but when there has been
no cross with a distinct breed, and there is a tendency in
both parents to revert to some long-lost character, this
tendency, for all that we can see to the contrary, may be
transmitted undiminished for an indefinite number of
generations. These two distinct cases of reversion are

CHAP. VI. THEIR REVERSION IN COLOUR.
247
often confounded together by those who have written on
inheritance.
Considering, on the one hand, the improbability of the
three assumptions which have just been discussed, and,
on the other hand, how simply the facts are explained on
the principle of reversion, we may conclude that the occa-
sional appearance in all the races, both when purely bred
and more especially when crossed, of blue birds, some-
times chequered, with double wing-bars, with white or
blue croups, with a bar at the end of the tail, and with the
outer tail-feathers edged with white, affords an argument
of the greatest weight in favour of the view that all are
descended from Columba livia, including under this name
the three or four wild varieties or sub-species before enu-
merated.
To sum up the six foregoing arguments, which are
opposed to the belief that the chief domestic races are
the descendants of at least eight or nine or perhaps a
dozen species; for the crossing of any less number would
not yield the characteristic differences between the seve-
ral races. Firstly, the improbability that so many spe-
cies should still exist somewhere, but be unknown to
ornithologists, or that they should have become within
the historical period extinct, although man has had
so little influence in exterminating the wild C. livia.
Secondly, the improbability of man in former times
having thoroughly domesticated and rendered fertile
under confinement so many species. Thirdly, these sup-
posed species having nowhere become feral. Fourthly,
the extraordinary fact that man should, intentionally or
by chance, have chosen for domestication several species,
extremely abnormal.in character; and furthermore, the
points of structure which render these supposed species
so abnormal being now highly variable. Fifthly, the
fact of all the races, though differing in many important
points of structure, producing perfectly fertile mongrels;
whilst all the hybrids which have been produced between

248
CHIP. VI.
DOMESTIC PIGEONS.
even closely allied species in the pigeon-family are sterile.
Sixthly, the remarkable statements just given on the
tendency in all the races, both when purely bred and
when crossed, to revert in numerous minute details of
colouring to the character of the wild rock-pigeon, and
to vary in a similar manner. To these arguments may
be added the extreme improbability that a number of
species formerly existed, which differed greatly from each
other in some few points, but which resembled each other
as closely as do the domestic races in other points of
structure, in voice, and in all their habits of life. When
these several facts and arguments are fairly taken into
consideration, it would require an overwhelming amount
of evidence to make us admit that the chief domestic
races are descended from several aboriginal stocks; and
of such evidence there is absolutely none.
The belief that the chief domestic races are descended
from several wild stocks no doubt has arisen from the
apparent improbability of such great modifications of
structure having been effected since man first domesti-
cated the rock-pigeon. Nor am I surprised at any degree
of hesitation in admitting their common origin: formerly,
when I went into my aviaries and watched such birds as
pouters, carriers, barbs, fantails, and short-faced tumblers,
&c., I could not persuade myself that they had all
descended from the same wild stock, and that man had
consequently in one sense created these remarkable modi-
fications. Therefore I have argued the question of their
origin at great, and, as some will think, superfluous
length.
Finally, in favour of the belief that all the races are
descended from a single stock, we have in Columbia livia
a still existing and widely distributed species, which can
be and has been domesticated in various countries. This
species agrees in most points of structure and in all its
habits of life, as well as occasionally in every detail of
plumage, with the several domestic races. It breeds

CHAP. VI.
249
FORMATION OF RACES.
freely with them, and produces fertile offspring. It
varies in a state of nature, sº and still more so when semi-
domesticated, as shown by comparing the Sierra Leone
pigeons with those of India, or with those which appa-
rently have run wild in Madeira. It has undergone
a still greater amount of variation in the case of the
numerous toy-pigeons, which no one supposes to be
descended from distinct species; yet some of these toy-
pigeons have transmitted their character truly for centu-
ries. Why, then, should we hesitate to believe in that
greater amount of variation which is necessary for the
production of the eleven chief races? It should be borne
in mind that in two of the most strongly-marked races,
namely, carriers and short-faced tumblers, the extreme
forms can be connected with the parent-species by gradu-
ated differences not greater than those which may be
observed between the dovecot-pigeons inhabiting differ-
ent countries, or between the various kinds of toy-
pigeons,-gradations which must certainly be attributed
to variation.
That circumstances have been eminently favourable for
the modification of the pigeon through variation and
selection will now be shown. The earliest record, as has
been pointed out to me by Professor Lepsius, of pigeons
in a domesticated condition, occurs in the fifth Egyptian
dynasty, about 3000 B.C. ; 31 but Mr. Birch, of the British
Museum, informs me that the pigeon appears in a bill of
fare in the previous dynasty. Domestic pigeons are
mentioned in Genesis, Leviticus, and Isaiah. In the
32
30 It deserves notice, as bearing on the
general subject of variation, that not only
C. livia presents several wild forms, re-
garded by some naturalists as species
and by others as sub-species or as mere
varieties, but that the species of several
allied genera are in the same predica-
ment. This is the case, as Mr. Blyth
has remarked to me, with Treron, Palum-
bus, and Turtur.
81 Denkmaler,' Abth. ii. Bl. 70.
32 The Dovecote,' by the Rev. E. S.
Dixon, 1851, pp. 11-13. Adolphe Pictet
(in his 'Les Origines Indo-Européennes,'
1859, p. 399) states that there are in the
ancient Sanscrit language between 25
and 30 names for the pigeon, and other
15 or 16 Persian names; none of these
are common to the European languages.
This fact indicates the antiquity of the
domestication in the East of the pigeon.
11*

250
CHAP. VI.
DOMESTIC PIGEONS.
time of the Romans, as we hear from Pliny,'' immense
prices were given for pigeons; “nay, they are come to
this pass, that they can reckon up their pedigree and
race.” In India, about the year 1600, pigeons were much,
valued by Akber Khan: 20,000 birds were carried about
with the court, and the merchants brought valuable
collections. “The monarchs of Iran and Turan sent him
some very rare breeds. His Majesty," says the courtly
historian, "by crossing the breeds, which method was
never practised before, has improved them astonish-
ingly."
1934 Akber Khan possessed seventeen distinct kinds,
eight of which were valuable for beauty alone. At about
this same period of 1600 the Dutch, according to Aldro-
vandi, were as eager about pigeons as the Romans had
formerly been. The breeds which were kept during the
fifteenth century in Europe and in India apparently dif-
fered from each other. Tavernier, in his Travels in 1677,
speaks, as does Chardin in 1735; of the vast number of
pigeon-houses in Persia; and the former remarks that, as
Christians were not permitted to keep pigeons, some of
the vulgar actually turned Mahometans for this sole
purpose. The Emperor of Morocco had his favourite
keeper of pigeons, as is mentioned in Moore's treatise,
published 1737. In England, from the time of Willughby
in 1678 to the present day, as well as in Germany and in
France, numerous treatises have been published on the
pigeon. In India, about a hundred years ago, a Persian
treatise was written; and the writer thought it no light
affair, for he begins with a solemn invocation, "in the
name of God, the gracious and merciful.” Many large
towns, in Europe and the United States, now have their
societies of devoted pigeon-fanciers: at present there are
three such societies in London. In India, as I hear from
Mr. Blyth, the inhabitants of Delhi and of some other
83 English translation, 1601, book x.
ch. xxxvii.
34 'Ayeen Akbery,' translated by F.
Gladvin, 4to. edit., vol. i. p. 270.

CHAP. VI.
251
FORMATION OF RACES.
great cities are eager fanciers. Mr. Layard informs me
that most of the known breeds are kept in Ceylon. In
China, according to Mr. Swinhoe of Amoy, and Dr.
Lockhart of Shangai, carriers, fantails, tumblers, and
other varieties are reared with care, especially by the
bonzes or priests. The Chinese fasten a kind of whistle
to the tail-feathers of their pigeons, and as the flock
wheels through the air they produce a sweet sound. In
Egypt the late Abbas Pacha was a great fancier of
fantails. Many pigeons are kept at Cairo and Constanti-
nople, and these have lately been imported by native
merchants, as I hear from Sir W. Elliot, into Southern
India, and sold at high prices.
The foregoing statements show in how many countries,
and during how long a period, many men have been pas-
sionately devoted to the breeding of pigeons. Hear how
an enthusiastic fancier at the present day writes: “If it
were possible for noblemen and gentlemen to know the
amazing amount of solace and pleasure derived from
Almond Tumblers, when they begin to understand their
properties, I should think that scarce any nobleman or
gentleman would be without their aviaries of Almond
Tumblers." 35 The pleasure thus taken is of
paramount
importance, as it leads amateurs carefully to note and
preserve each slight deviation of structure which strikes
their fancy. Pigeons are often closely confined during
their whole lives; they do not partake of their naturally
varied diet; they have often been transported from one
climate to another; and all these changes in their condi-
tions of life would be likely to cause variability. Pigeons
have been domesticated for nearly 5000 years, and have
been kept in many places, so that the numbers reared
under domestication must have been enormous; and this
is another circumstance of high importance, for it obvi-
ously favours the chance of rare modifications of structure
35 J. M. Eaton, Treatise on the Almond Tumbler,' 1851; Preface, p. vi.

252
CHAP, VI.
DOMESTIC PIGEONS.
occasionally appearing. Slight variations of all kinds
would almost certainly be observed, and, if valued, would,
owing to the following circumstances, be preserved and
propagated with unusual facility. Pigeons, differently
from any other domesticated animal, can easily be mated
for life, and, though kept with other pigeons, they rarely
prove unfaithful to each other. Even when the male
does break his marriage vow, he does not permanently
desert his mate. I have bred in the same aviaries many
pigeons of different kinds, and never reared a single bird
of an impure strain. Hence a fancier can with the great-
est ease select and match his birds. He will also soon
see the good results of his care; for pigeons breed with
extraordinary rapidity. He may freely reject inferior
birds, as they serve at an early age as excellent food. To
sum up, pigeons are easily kept, paired, and selected; vast
numbers have been reared; great zeal in breeding them
has been shown by many men in various countries; and
this would lead to their close discrimination, and to a
strong desire to exhibit some novelty, or to surpass other
fanciers in the excellence of already established breeds.
36
History of the principal Races of the Pigeon.
Before discussing the means and steps by which the chief races
have been formed, it will be advisable to give some historical details,
for more is known of the history of the pigeon, little though this be,
than of any other domesticated animal. Some of the cases are in-
teresting as proving how long domestic varieties may be propagated
with exactly the same or nearly the same characters; and other cases
are still more interesting as showing how slowly but steadily races
have been greatly modified during successive generations. In the
last chapter I stated that Trumpeters and Laughers, both so remark-
able for their voices, seem to have been perfectly characterized in
1735 ; and Laughers were apparently known in India before the
year 1600. Spots in 1676, and Nuns in the time of Aldrovandi, be-
fore 1600, were coloured exactly as they now are. Common Tum-
36 As, in the following discussion I state that this chapter was completed in
often speak of the present time, I should the year 1855.

CEAP. VI. HISTORY OF THE PRINCIPAL RACES. - 253
blers and Ground Tumblers exhibited in India, before the year 1600,
the same extraordinary peculiarities of flight as at the present day,
for they are well described in the 'Ayeen Akbery. These breeds
may all have existed for a much longer period ; we know only that
they were perfectly characterized at the dates above given. The
average length of life of the domestic pigeon is probably about five or
six years; if so, some of these races have retained their character
perfectly for at least forty or fifty generations.
Pouters.—These birds, as far as a very short description serves for
comparison, appear to have been well characterized in Aldrovandi's
time, 37 before the year 1600. Length of body and length of leg are
at the present time the two chief points of excellence. In 1735
Moore said (see Mr. J. M. Eaton's edition)-and Moore was a first-
rate fancier—that he once saw a bird with a body 20 inches in length,
" though 17 or 18 inches is reckoned a very good length;" and he
has seen the legs very nearly 7 inches in length, yet a leg 6 or 68
long “must be allowed to be a very good one." Mr. Bult, the most
successful breeder of Pouters in the world, informs me that at pre-
sent (1858) the standard length of the body is not less than 18 inches;
but he has measured one bird 19 inches in length, and has heard of
20 and 22 inches, but doubts the truth of these latter statements.
The standard length of the leg is now 7 inches, but Mr. Bult has
recently measured two of his own birds with legs 71 long. So that
in the 123 years which have elapsed since 1735 there has been hardly
any increase in the standard length of the body ; 17 or 18 inches
was formerly reckoned a very good length, and now 18 inches is the
minimum standard ; but the length of leg seems to have increased,
as Moore never saw one quite 7 inches long; now the standard is 7,
and two of Mr. Bult's birds measured 74 inches in length. The ex-
tremely slight improvement in Pouters, except in the length of the
leg, during the last 123 years, may be partly accounted for by the
neglect which they suffered, as I am informed by Mr. Bult, until
within the last 20 or 30 years. About 1765 38 there was a change of
fashion, stouter and more feathered legs being preferred to thin and
nearly naked legs.
Fantails.-The first notice of the existence of this breed is in In-
dia, before the year 1600, as given in the 'Ayeen Akbery ;'
date, judging from Aldrovandi, the breed was unknown in Europe.
In 1677 Willughby speaks of a Fantail with 26 tail-feathers; in
37 Ornithologie,' 1600, vol. ii. p. 360.
38 A Treatise on Domestic Pigeons,'
dedicated to Mr. Mayor, 1765. Preface,
39 Mr. Blyth has given a translation
of part of the 'Ayeen Akbery' in An-
nals and Mag. of Nat. Hist.,' vol. xix.,
1847, p. 104.
P. xiv.

254
CHAP. VI.
DOMESTIC PIGEONS.
1735 Moore saw one with 36 feathers; and in 1824 MM. Boitard and
Corbié assert that in France birds can easily be found with 42 tail-
feathers. In England, the number of the tail-feathers is not at
present so much regarded as their upward direction and expansion.
The general carriage of the bird is likewise now much regarded.
• The old descriptions do not suffice to show whether in these latter
respects there has been much improvement; but if fantails had
formerly existed with their heads and tails touching each other, as
at the present time, the fact would almost certainly have been
noticed. The Fantails which are now found in India probably show
the state of the race, as far as carriage is concerned, at the date of
their introduction into Europe; and some, said to have been brought
from Calcutta, which I kept alive, were in a marked manner in-
ferior to our exhibition birds. The Java Fantail shows the same
difference in carriage; and although Mr. Swinhoe has counted 18
and 24 tail-feathers in his birds, a first-rate specimen sent to me
had only 14 tail-feathers.
Jacobins. This breed existed before 1600, but the hood, judging
from the figure given by Aldrovandi, did not enclose the head nearly
so perfectly as at present: nor was the head then white; nor were
the wings and tail so long, but this last character might have been
overlooked by the rude artist. In Moore's time, in 1735, the Jacobin
was considered the smallest kind of pigeon, and the bill is said to
be very short. Hence either the Jacobin, or the other kinds with
which it was then compared, must have been since considerably
modified; for Moore's description (and it must be remembered that
he was a first-rate judge) is clearly not applicable, as far as size of
body and length of beak are concerned, to our present Jacobins.
In 1795, judging from Bechstein, the breed had assumed its present
character.
Turbits.—It has generally been supposed by the older writers on
pigeons, that the Turbit is the Cortbeck of Aldrovandi; but if this
be the case, it is an extraordinary fact that the characteristic frill
should not have been noticed. The beak, moreover, of the Cortbeck
is described as closely resembling that of the Jacobin, which shows
a change in the one or the other race. The Turbit, with its
characteristic frill and bearing its present name, is described by
Willughby in 1677; and the bill is said to be like that of the bull-
finch,-a good comparison, but now more strictly applicable to the
beak of the Barb. The sub-breed called the Owl was well known in
Moore's time, in 1735.
Tumblers.—Common Tumblers, as well as Ground Tumblers,
perfect as far as tumbling is concerned, existed in India before the
year 1600; and at this period diversified modes of flight, such as

CHAP. VI. HISTORY OF THE PRINCIPAL RACES.
255
flying at night, the ascent to a great height, and manner of descent,
seem to have been much attended to, as at the present time, in India.
Belon 40 in 1555 saw in Paphlagonia what he describes as "a very
new thing, viz. pigeons which flew so high in the air that they were
lost to view, but returned to their pigeon-house without separating."
This manner of flight is characteristic of our present Tumblers, but
it is clear that Belon would have mentioned the act of tumbling if
the pigeons described by him had tumbled. Tumblers were not
known in Europe in 1600, as they are not mentioned by Aldrovandi,
who discusses the flight of pigeons. They are briefly alluded to by
Willughby, in 1687, as small pigeons "which show like footballs in
the air.” The short-faced race did not exist at this period, as Wil-
lughby could not have overlooked birds so remarkable for their
small size and short beaks. We can even trace some of the steps
by which this race has been produced. Moore in 1735 enumerates
correctly the chief points of excellence, but does not give any de-
scription of the several sub-breeds; and from this fact Mr. Eaton in-
fers 41 that the short-faced Tumbler had not then come to full per-
fection. Moore even speaks of the Jacobin as being the smallest
pigeon. Thirty years afterwards, in 1765, in the Treatise dedicated
to Mayor, short-faced Almond Tumblers are fully described, but the
author, an excellent fancier, expressly states in his Preface (p. xiv.)
that," from great care and expense in breeding them, they have
arrived to so great perfection and are so different from what they
were 20 or 30 years past, that an old fancier would have condemned
them for no other reason than because they are not like what used
to be thought good when he was in the fancy before.” Hence it
would appear that there was a rather sudden change in the charac-
ter of the short-faced Tumbler at about this period; and there is
reason to suspect that a dwarfed and half-monstrous bird, the parent-
form of the several short-faced sub-breeds, then appeared. I suspect
this because short-faced Tumblers are born with their beaks (ascer-
tained by careful measurement) as short, proportionally with the
size of their bodies, as in the adult bird ; and in this respect they
differ greatly from all other breeds, which slowly acquire during
growth their various characteristic qualities.
Since the year 1765 there has been some change in one of the
chief characters of the short-faced Tumbler, namely, in the length
of the beak. Fanciers measure the "head and beak” from the tip.
of the beak to the front corner of the eyeball. About the year 1765
40 L'Hist. de la Nature des Oiseaux,' p. 314.
41 Treatise on Pigeons,' 1852, p. 64.

256
CHAP. VI.
DOMESTIC PIGEONS.
a "head and beak” was considered good,42 which, measured in the
usual manner, was of an inch in length; now it ought not to ex-
ceed of an inch ; "it is however possible," as Mr. Eaton candidly
confesses, " for a bird to be considered as pleasant or neat even at
of an inch, but exceeding that length it must be looked upon as un-
worthy of attention." Mr. Eaton states that he has never seen in
the course of his life more than two or three birds with the "head
and beak” not exceeding half an inch in length; "still I believe in
the course of a few years that the head and beak will be shortened,
and that balf-inch birds will not be considered so great a curiosity
as at the present time.” That Mr. Eaton's opinion deserves atten-
tion cannot be doubted, considering his success in winning prizes
at our exhibitions. Finally in regard to the Tumbler it may be con-
cluded from the facts above given that it was originally introduced
into Europe, probably first into England, from the East; and that it
then resembled our common English Tumbler, or more probably the
Persian or Indian Tumbler, with a beak only just perceptibly shorter
than that of the common dovecot-pigeon. With respect to the
short-faced Tumbler, which is not known to exist in the East, there
can hardly be a doubt that the whole wonderful change in the size
of the head, beak, body, and feet, and in general carriage, has been
produced during the last two centuries by continued selection, aided
probably by the birth of a semi-monstrous bird somewhere about the
year 1750
Runts. Of their history little can be said. In the time of Pliny
the pigeons of Campania were the largest known; and from this
fact alone some authors assert that they were Runts. In Aldrovandi's
time, in 1600, two sub-breeds existed ; but one of them, the short-
beaked, is now extinct in Europe.
Barbs.-Notwithstanding statements to the contrary, it seems to
me impossible to recognise the barb in Aldrovandi's descriptions and
figures; four breeds, however existed in the year 1600 which were
evidently allied both to Barbs and Carriers. To show how difficult
it is to recognise some of the breeds described by Aldrovandi, I will
give the different opinions in regard to the above four kinds, named
by him C. Indica, Cretensis, Gutturosa, and Persica. Willughby
thought that the Columba Indica was a Turbit, but the eminent
fancier Mr. Brent believes that it was an inferior Barb: C. Cretensis,
with a short beak and a swelling on the upper mandible, cannot be
recognised: C. (falsely called) gutturosa, which from its rostrum,
breve, crassum, et tuberosum seems to me to come nearest to the
42 J. M. Eaton's Treatise on the Tumbler,' 1851. Compare p. V. of Pre-
Breeding and Managing of the Almond face, p. 9, and p. 32.

Chap. VI. HISTORY OF THE PRINCIPAL RACES. 257
Barb, Mr. Brent believes to be a Carrier; and lastly, the C. Persica
et Turcica, Mr. Brent thinks, and I quite concur with him, was a
short-beaked Carrier with very little wattle. In 1687 the Barb was
known in England, and Willughby describes the beak as like that
of the Turbit; but it is not credible that his Barb should have had
a beak like that of our present birds, for so accurate an observer
could not have overlooked its great breadth.
English Carrier.- We may look in vain in Aldrovandi's work
for any bird resembling our prize Carriers ; the C. Persica et Turcica
of this author comes the nearest, but is said to have had a short
thick beak; therefore it must have approached in character a Barb,
and have differed greatly from our Carriers. In Willughby's time,
in 1677, we can clearly recognise the Carrier, but he adds," the bill
is not short, but of a moderate length," a description which no one
would apply to our present Carriers, so conspicuous for the extra-
ordinary length of their beaks. The old names given in Europe
to the Carrier, and the several names now in use in India, indicate
that Carriers-originally came from Persia; and Willughby's de-
scription would perfectly apply to the Bussorah Carrier as it now
exists in Madras. In later times we can partially trace the progress
of change in our English Carriers : Moore in 1735 says "an inch
and a half is reckoned a long beak, though there are very good
Carriers that are found not to exceed an inch and a quarter.”
These birds must have resembled, or perhaps been a little superior
to, the Carriers previously described, which are now found in
Persia. In England at the present day “there are," as Mr. Eaton 43
states, “beaks that would measure (from edge of eye to tip of beak)
one inch and three-quarters, and some few even two inches in
length."
From these historical details we see that nearly all the
chief domestic races existed before the year 1600. Some
remarkable only for colour appear to have been identical
with our present breeds, some were nearly the same,
some considerably different, and some have since become
extinct. Several breeds, such as Finnikins and Turners,
the swallow-tailed pigeon of Bechstein and the Carme-
lite, seem both to have originated and to have disap-
peared within this same period. Any one now visiting
a well stocked English aviary would certainly pick out
43 Treatise on Pigeons,' 1852, p. 41.

258
CHAP. VÌ
DOMESTIC PIGEONS.
as the most distinct kinds, the massive Runt, the Carrier
with its wonderfully elongated beak and great wattles,
the Barb with its short broad beak and eye-wattles, the
short-faced Tumbler with its small conical beak, the
Pouter with its great crop, long legs and body, the Fan-
tail with its upraised, widely-expanded, well-feathered
tail, the Turbit with its frill and short blunt beak, and
the Jacobin with its hood. Now, if this same person
could have viewed the pigeons kept before 1600 by Akber
Khan in India and by Aldrovandi in Europe, he would
have seen the Jacobin with a less perfect hood; the Tur-
bit apparently without its frill; the Pouter with shorter
legs, and in every way less remarkable—that is, if Aldro-
vandi's Pouter resembled the old German kind; the Fan-
tail would have been far less singular in appearance, and
would have had much fewer feathers in its tail ; he would
have seen excellent flying Tumblers, but he would in
vain have looked for the marvellous short-faced breeds ;
he would have seen birds allied to barbs, but it is ex-
tremely doubtful whether he would have met with our
actual Barbs; and lastly, he would have found Carriers
with beaks and wattle incomparably less developed than
in our English Carriers. He might have classed most of
the breeds in the same groups as at present; but the
differences between the groups were then far less strong-
ly pronounced than at present. In short, the several
breeds had at this early period not diverged in so great
a degree from their aboriginal common parent, the wild
rock-pigeon.
Manner of Formation of the chief Races.
We will now consider more closely the probable steps
by which the chief races have been formed. As long as
pigeons are kept semi-domesticated in dovecots in their
native country, without any care in selecting and match-
ing them, they are liable to little more variation than the

CHAP. VI.
259
FORMATION OF CHIEF RACES.
wild C. livia, namely, in the wings becoming chequered
with black, in the croup being blue or white, and in the
size of the body. When, however, dovecot-pigeons are
transported into diversified countries, such as Sierra
Leone, the Malay archipelago, and Madeira (where the
wild C. livia is not known to exist), they are exposed to
new conditions of life; and apparently in consequence
they vary in a somewhat greater degree. When closely
confined, either for the pleasure of watching them, or to
prevent their straying, they must be exposed, even under
their native climate, to considerably different conditions ;
for they cannot obtain their natural diversity of food ;
and, what is probably more important, they are abun-
dantly fed, whilst debarred from taking much exercise.
Under these circumstances we might expect to find, from
the analogy of all other domesticated animals, a greater
amount of individual variability than with the wild
pigeon; and this is the case. The want o exercise ap-
parently tends to reduce the size of the feet and organs
of flight; and then, from the law of correlation of
growth, the beak apparently becomes affected. From
what we now see occasionally taking place in our aviaries,
we may conclude that sudden variations or sports, such
as the appearance of a crest of feathers on the head, of
feathered feet, of a new shade of colour, of an additional
feather in the tail or wing, would occur at rare intervals
during the many centuries which have elapsed since the
pigeon was first domesticated. At the present day such
sports” are generally rejected as blemishes; and there
is so much mystery in the breeding of pigeons that, if
a valuable sport did occur, its history would often be
concealed. Before the last hundred and fifty years, there
is hardly a chance of the history of
any
such sport hav-
ing been recorded. But it by no means follows from
this that such sports in former times, when the pigeon
had undergone much less variation, would have been re-
jected. We are profoundly ignorant of the cause of

260
CHAP. VI.
DOMESTIC PIGEONS.
each sudden and apparently spontaneous variation, as
well as of the infinitely numerous shades of difference
between the birds of the same family. But in a future
chapter we shall see that all such variations appear to be
the indirect result of changes of some kind in the con-
ditions of life.
Hence, after a long course of domestication, we might
expect to see in the pigeon much individual variability,
and occasional sudden variations, as well as slight modi-
fications from the lessened use of certain parts, together
with the effects of correlation of growth. But without
selection all this would produce only a trifling or no re-
sult; for without such aid differences of all kinds would,
from the two following causes, soon disappear. In a
healthy and vigorous lot of pigeons many more young
birds are killed for food or die than are reared to matu-
rity; so that an individual having any peculiar character,
if not selected, would run a good chance of being destroy-
ed, and if not destroyed, the peculiarity in question would
almost certainly be obliterated by free intercrossing. It
might, however, occasionally happen that the same varia-
• tion repeatedly occurred, owing to the action of peculiar
and uniform conditions of life, and in this case it would pre-
vail independently of selection. But when selection is
brought into play all is changed; for this is the foundation-
stone in the formation of new races; and with the pigeon,
circumstances, as we have already seen, are eminently fa-
vourable for selection. When a bird presenting some
conspicuous variation has been preserved, and its offspring
have been selected, carefully matched, and again propa-
gated, and so onwards during successive generations, the
principle is so obvious that nothing more need be said
about it. This may be called methodical selection, for
the breeder has a distinct object in view, namely, to pre-
serve some character which has actually appeared; or to
create some improvement already pictured in his mind.
Another form of selection has hardly been noticed by

CHAP. VI.
261
FORMATION OF CHIEF RACES.
those authors who have discussed this subject, but is even
more important. This form may be called unconscious
selection, for the breeder selects his birds unconsciously,
unintentionally, and without method, yet he surely though
slowly produces a great result. I refer to the effects which
follow from each fancier at first procuring and afterwards
rearing as good birds as he can, according to his skill,
and according to the standard of excellence at each suc-
cessive period. He does not wish permanently to modify
the breed; he does not look to the distant future, or
speculate on the final result of the slow accumulation dur-
ing many generations of successive slight changes: he is
content if he possesses a good stock, and more than con-
tent if he can beat his rivals. The fancier in the time of
Aldrovandi, when in the year 1600 he admired his own
jacobins, pouters, or carriers, never reflected what their
descendants in the year 1860 would become; he would
have been astonished could he have seen our jacobins, our
improved English carriers, and our pouters; he would
probably have denied that they were the descendants of
his own once admired stock, and he would perhaps not
have valued them, for no other reason, as was written in
1765, “than because they were not like what used to be
thought good when he was in the fancy.” No one will
attribute the lengthened beak of the carrier, the shorten-
ed beak of the short-faced tumbler, the lengthened leg
of the pouter, the more perfectly-enclosed hood of the
jacobin, &c.,-changes effected since the time of Aldro-
vandi, or even since a much later period, -to the direct
and immediate action of the conditions of life. For these
several races have been modified in various and even in
directly opposite ways, though kept under the same cli-
mate and treated in all respects in as nearly uniform a
manner as possible. Each slight change in the length or
shortness of the beak, in the length of leg, &c., has no
doubt been indirectly and remotely caused by some change
in the conditions to which the bird has been subjected,

262
CHAP. VI.
DOMESTIC PIGEONS.
but we must attribute the final result, as is manifest in
those cases of which we have any historical record, to the
continued selection and accumulation of many slight suc-
cessive variations.
The action of unconscious selection, as far as pigeons
are concerned, depends on a universal principle in human
nature, namely, on our rivalry, and desire to outdo our
neighbours. We see this in every fleeting fashion, even
in our dress, and it leads the fancier to endeavour to ex-
aggerate every peculiarity in his breeds. A great autho-
rity on pigeons 44 says, “Fanciers do not and will not ad-
mire a medium standard, that is, half and half, which is
neither here nor there, but admire extremes.” After re-
marking that the fancier of short-faced beard tumblers
wishes for a very short beak, and that the fancier of long-
faced beard tumblers wishes for a very long beak, he
says, with respect to one of intermediate length, “Don't
deceive yourself. Do you suppose for a moment the short
or the long-faced fancier would accept such a bird as a
gift ? Certainly not; the short-faced fancier could see no
beauty in it; the long-faced fancier would swear there
was no use in it, &c.” In these comical passages, written
seriously, we see the principle which has ever guided fan-
ciers, and has led to such great modifications in all the
domestic races which are valued solely for their beauty
or curiosity.
Fashions in pigeon-breeding endure for long periods;
we cannot change the structure of a bird as quickly as
we can the fashion of our dress. In the time of Aldro-
vandi, no doubt the more the pouter inflated his crop, the
more he was valued. Nevertheless, fashions do to a cer-
tain extent change; first one point of structure and then
another is attended to; or different breeds are admired at
different times and in different countries. As the author
just quoted remarks, “the fancy ebbs and flows; a tho-
44 Eaton's "Treatise on Pigeons,' 1858, p. 86.

CHAP. VI.
263
FORMATION OF CHIEF RACES.
rough fancier now-a-days never stoops to breed toy-birds;"
yet these very “toys” are now most carefully bred in
Germany. Breeds which at the present time are highly
valued in India are considered worthless in England.
No doubt, when breeds are neglected, they degenerate;
still we may believe that, as long as they are kept under
the same conditions of life, characters once gained will be
partially retained for a long time, and may form the
starting-point for a future course of selection.
Let it not be objected to this view of the action of un-
conscious selection that fanciers would not observe or
care for extremely slight differences. Those alone who
have associated with fanciers can be thoroughly aware of
their accurate powers of discrimination acquired by long
practice, and of the care and labour which they bestow
on their birds. I have known a fancier deliberately study
his birds day after day to settle which to match together
and which to reject. Observe how difficult the subject
appears to one of the most eminent and experienced fan-
ciers. Mr. Eaton, the winner of many prizes, says, “I
would here particularly guard you against keeping too
great a variety of pigeons, otherwise you will know a lit-
tle about all the kinds, but nothing about one as it ought
to be known.” “It is possible there may be a few fan-
ciers that have a good general knowledge of the several
fancy pigeons, but there are many who labour under the
delusion of supposing they know what they do not."
Speaking exclusively of one sub-variety of one race,
namely, the short-faced almond tumbler, and after saying
that some fanciers sacrifice every property to obtain a
good head and beak, and that other fanciers sacrifice
everything for plumage, he remarks: “Some young fan-
ciers who are over covetous go in for all the five proper-
ties at once, and they have their reward by getting
nothing." In India, as I hear from Mr. Blyth, pigeons
are likewise selected and matched with the greatest care.
But we must not judge of the slight differences which

264
CHAP. VI.
DOMESTIC PIGEONS.
would have been valued in ancient days, by those which
are now valued after the formation of many races, each
with its own standard of perfection, kept uniform by our
numerous Exhibitions. The ambition of the most ener-
getic fancier may be fully satisfied by the difficulty of ex-
celling other fanciers in the breeds already established,
without trying to form a new one.
A difficulty with respect to the power of selection will
perhaps already have occurred to the reader, namely,
what could have led fanciers first to attempt to make
such singular breeds as pouters, fantails, carriers, &c. ?
But it is this very difficulty which the principle of un-
conscious selection removes. Undoubtedly no fancier
ever did intentionally make such an attempt. All that
we need suppose is that a variation occurred sufficiently
marked to catch the discriminating eye of some ancient
fancier, and then unconscious selection carried on for
many generations, that is, the wish of succeeding fanciers
to excel their rivals, would do the rest. In the case of
the fantail we may suppose that the first progenitor of
the breed had a tail only slightly erected, as may now
be seen in certain runts, with some increase in the
number of the tail-feathers, as now occasionally occurs
with nuns.
In the case of the pouter we may suppose
that some bird inflated its crop a little more than other
pigeons, as is now the case in a slight degree with the
esophagus of the turbit. We do not in the least know
the origin of the common tumbler, but we may suppose
that a bird was born with some affection of the brain,
leading it to make somersaults in the air, and the diffi-
culty in this case is lessened, as we know that, before
the year 1600, in India, pigeons remarkable for their
diversified manner of flight were much valued, and by
45 See Neumeister's figure of the Florence runt, tab. 13, in "Das Ganze der Tau-
benzucht.'

CHAP, VI.
FORMATION OF CHIEF RACES.
265
the order of the Emperor Akber Khan were sedulously
trained and carefully matched.
In the foregoing cases we have supposed that a sudden
variation, conspicuous enough to catch a fancier's eye,
first appeared; but even this degree of abruptness in the
process of variation is not necessary for the formation of
a new breed. When the same kind of pigeon has been
kept pure, and has been bred during a long period by
two or more fanciers, slight differences in the strain can
often be recognised. Thus I have seen first-rate jacobins
in one man's possession which certainly differed slightly
in several characters from those kept by another. I pos-
sessed some excellent barbs descended from a pair which
had won a prize, and another lot descended from a stock
formerly kept by that famous fancier Sir John Sebright,
and these plainly differed in the form of the beak; but
the differences were so slight, that they could hardly be
described by words. Again, the common English and
Dutch tumbler differ in a somewhat greater degree, both
in length of beak and shape of head. What first caused
these slight differences cannot be explained any more
than why one man has a long nose and another a short
In the strains long kept distinct by different fan-
ciers, such differences are so common that they cannot
be accounted for by the accident of the birds first chosen
for breeding having been originally as different as they
The explanation no doubt lies in selection of a
slightly different nature having been applied in each
case; for no two fanciers have exactly the same taste,
and consequently no two, in choosing and carefully
matching their birds, prefer or select exactly the same.
As each man naturally admires his own birds, he goes on
continually exaggerating by selection whatever slight
peculiarities they may possess. This will more especially
happen with fanciers living in different countries, who
do not compare their stocks and aim at a common stand-
ard of perfection. Thus, when a mere strain has once
12
one.
now are.

266
CHAP. VI.
DOMESTIC PIGEONS.
been formed, unconscious selection steadily tends to
augment the amount of difference, and thus converts the
strain into a sub-breed, and this ultimately into a well-
marked breed or race.
The principle of correlation of growth should never be
lost sight of. Most pigeons have small feet, apparently
caused by their lessened use, and from correlation, as it
would appear, their beaks have likewise become reduced
in length. The beak is a conspicuous organ, and, as
soon as it had thus become perceptibly shortened, fanciers
would almost certainly strive to reduce it still more by
the continued selection of birds with the shortest beaks ;
whilst at the same time other fanciers, as we know has
actually been the case, would, in other sub-breeds, strive
to increase its length. With the increased length of the
beak, the tongue would become greatly lengthened, as
would the eyelids with the increased development of
the eye-wattles; with the reduced or increased size of
the feet the number of the scatellæ would vary; with the
length of the wing the number of the primary wing-
feathers would differ; and with the increased length of
the body in the pouter the number of the sacral vertebræ
would be augmented. These important and correlated
differences of structure do not invariably characterise
any breed; but if they had been attended to and selected
with as much care as the more conspicuous external
differences, there can hardly be a doubt that they would
have been rendered constant. Fanciers could assuredly
have made a race of tumblers with nine instead of ten
primary wing-feathers, seeing how often the number nine
appears without any wish on their part, and indeed in
the case of the-white-winged varieties in opposition to
their wish. In a similar manner, if the vertebra had
been visible and had been attended to by fanciers, as-
suredly an additional number might easily have been
fixed in the pouter. If these latter characters had once
been rendered constant we should never have suspected

CHAP. VI.
267
FORMATION OF CHIEF RACES.
that they had at first been highly variable, or that they
had arisen from correlation, in the one case with the
shortness of the wings, and in the other case with the
length of the body.
In order to understand how the chief domestic races
have become distinctly separated from each other, it is
important to bear in mind, that fanciers constantly try
to breed from the best birds, and consequently that those
which are inferior in the requisite qualities are in each
generation neglected; so that after a time the less im-
proved parent-stocks and many subsequently formed
intermediate grades become extinct. This has occurred
in the case of the pouter, turbit, and trumpeter, for these
highly improved breeds are now left without any links
closely connecting them either with each other or with the
aboriginal rock-pigeon. In other countries, indeed, where
the same care has not been applied, or where the same
fashion has not prevailed, the earlier forms may long
remain unaltered or altered only in a slight degree, and
we are thus sometimes enabled to recover the connecting
links. This is the case in Persia and India with the tum-
bler and carrier, which there differ but slightly from the
rock-pigeon in the proportions of their beaks. So again
in Java, the fantail sometimes has only fourteen caudal
feathers, and the tail is much less elevated and expanded
than in our improved birds; so that the Java bird forms
a link between a first-rate fantail and the rock-pigeon.
Occasionally a breed may be retained for some particu-
lar quality in a nearly unaltered condition in the same
country, together with highly modified offshoots or sub-
breeds, which are valued for some distinct property. We
see this exemplified in England, where the common tum-
bler, which is valued only for its flight, does not differ
much from its parent-form, the Eastern tumbler; whereas
the short-faced tumbler has been prodigiously modified,
from being valued, not for its flight, but for other quali-
ties. But the common-flying tumbler of Europe has

268
CHAP. VI.
DOMESTIC PIGEONS.
already begun to branch out into slightly different sub-
breeds, such as the common English tumbler, the Dutch
roller, the Glasgow house-tumbler, and the long-faced
beard tumbler, &c.; and in the course of centuries,
unless fashions greatly change, these sub-breeds will
diverge through the slow and insensible process of uncon-
scious selection, and become modified, in a greater and
greater degree. After a time the perfectly graduated
links, which now connect all these sub-breeds together,
will be lost, for there would be no object and much diffi-
culty in retaining such a host of intermediate sub-varie-
ties.
The principle of divergence, together with the extinc-
tion of the many previously existing intermediate forms,
is so important for understanding the origin of domestic
races, as well as of species in a state of nature, that I
will enlarge a little more on this subject. Our third main
group includes carriers, barbs, and runts, which are
plainly related to each other, yet wonderfully distinct in
several important characters. According to the view given
in the last chapter, these three races have probably de-
scended from an unknown race having an intermediate cha-
racter and this from the rock-pigeon. Their characteristic
differences are believed to be due to different breeders
having at an early period admired different points of
structure; and then, on the acknowledged principle of
admiring extremes, having gone on breeding, without
any thought of the future, as good birds as they could, -
carrier-fanciers preferring long beaks with much wat-
tle,-barb-fanciers preferring short thick beaks with
much eye-wattle,—and runt-fanciers not caring about the
beak or wattle, but only for the size and weight of the
body. This process will have led to the neglect and final
extinction of the earlier, inferior, and intermediate birds;
and thus it has come to pass, that in Europe these three
races are now so extraordinarily distinct from each other.
But in the East, whence they were originally brought,

CHAP. VI.
269
FORMATION OF CHIEF RACES. .
the fashion has been different, and we there see breeds
which connect the highly modified English carrier with
the rock-pigeon, and others which to a certain extent
connect carriers and runts. Looking back to the time
of Aldrovandi, we find that there existed in Europe,
before the year 1600, four breeds which were closely
allied to carriers and barbs, but which competent au-
thorities cannot now identify with our present barbs and
carriers; nor can Aldrovandi's runts be identified with
our present runts. These four breeds certainly did not
differ from each other nearly so much as do our existing
English carriers, barbs, and runts. All this is exactly
what might have been anticipated. If we could collect
all the pigeons which have ever lived, from before the
time of the Romans to the present day, we should be
able to group them in several lines, diverging from the
parent rock-pigeon. Each line would consist of almost
insensible steps, occasionally broken by some slightly
greater variation or sport, and each would culminate in
one of our present highly modified forms. Of the many
former connecting links, some would be found to have
become absolutely extinct without having left any issue,
whilst others though extinct would be seen to be the
progenitors of the existing races.
I have heard it remarked as a strange circumstance
that we occasionally hear of the local or complete ex-
tinction of domestic races, whilst we hear nothing of their
origin. How, it has been asked, can these losses be com-
pensated, and more than compensated, for we know that
with almost all domesticated animals the races have largely
increased in number since the time of the Romans? But
on the view here given, we can understand this apparent
contradiction. The extinction of a race within historical
times is an event likely to be noticed; but its gradual
and scarcely sensible modification through unconscious
selection, and its subsequent divergence, either in the
same or more commonly in distant countries, into two or

270
CHAP. VI.
DOMESTIC PIGEONS.
more strains, and their gradual conversion into sub-breeds,
and these into well-marked breeds, are events which
would rarely be noticed. The death of a tree, that has
attained gigantio dimensions, is recorded; the slow
growth of smaller trees and their increase in number
excite no attention.
In accordance with the belief of the great power of
selection, and of the little direct power of changed con-
ditions of life, except in causing general variability or
plasticity of organisation, it is not surprising that dovecot-
pigeons have remained unaltered from time immemorial;
and that some toy-pigeons, which differ in little else
besides colour from the dovecot-pigeon, have retained the
same character for several centuries. For when one of
these toy-pigeons had once become beautifully and sym-
metrically coloured,—when, for instance, a Spot had been
produced with the crown of its head, its tail, and tail-
coverts of a uniform colour, the rest of the body being
snow-white, no alteration or improvement would be
desired. On the other hand, it is not surprising that
during this same interval of time our highly-bred pigeons
have undergone an astonishing amount of change; for in
regard to them there is no defined limit to the wish of the
fancier, and there is no known limit to the variability of
their characters. What is there to stop the fancier
desiring to give to his carrier a longer and longer beak,
or to his tumbler a shorter and shorter beak? nor has the
extreme limit of variability in the beak, if there be any
such limit, as yet been reached. Notwithstanding the
great improvement effected within recent times in the
short-faced almond tumbler, Mr. Eaton remarks, “the
field is still as open for fresh competitors as it was one
hundred years ago;
years ago;” but this is perhaps an exaggerated
assertion, for the young of all highly improved fancy birds
are extremely liable to disease and death.
I have heard it objected that the formation of the
several domestic races of the pigeon throws no light on

CHAP. VI.
271
FORMATION OF CHIEF RACES.
the origin of the wild species of the Columbidæ, because
their differences are not of the same nature. The
domestic races for instance do not differ, or differ hardly
at all, in the relative lengths and shapes of the primary
wing-feathers, in the relative length of the hind toe, or in
habits of life, as in roosting and building in trees. But
the above objection shows how completely the principle
of selection has been misunderstood. It is not likely that
characters selected by the caprice of man should resemble
differences preserved under natural conditions, either from
being of direct service to each species, or from standing
in correlation with other modified and serviceable struc-
tures. Until man selects birds differing in the relative
length of the wing-feathers or toes, &c., no sensible
change in these parts should be expected. Nor could
man do anything unless these parts happened to vary
under domestication: I do not positively assert that this
is the case, although I have seen traces of such variability
in the wing-feathers, and certainly in the tail-feathers. It
would be a strange fact if the relative length of the hind
toe should never vary, seeing how variable the foot is
both in size and in the number of the scutellæ. With
respect to the domestic races not roosting or building in
trees, it is obvious that fanciers would never attend to or
select such changes in habits; but we have seen that the
pigeons in Egypt, which do not for some reason like
settling on the low mud hovels of the natives, are led,
apparently by compulsion, to perch in crowds on the
trees. We may even affirm that, if our domestic races
had become greatly modified in any of the above specified
respects, and it could be shown that fanciers had never
attended to such points, or that they did not stand in
correlation with other selected characters, the fact, on the
principles advocated in this chapter, would have offered
a serious difficulty.
Let us briefly sum up the last two chapters on the
pigeon. We may conclude with confidence that all the

272
CHAP, VI.
DOMESTIC PIGEONS.
domestic races, notwithstanding their great amount of
difference, are descended from the Columba livia, includ-
ing under this name certain wild races. But the differ-
ences between these latter forms throw no light whatever
on the characters which distinguish the domestic races.
In each breed or sub-breed the individual birds are more
variable than birds in a state of nature; and occasional-
ly they vary in a sudden and strongly-marked manner.
This plasticity of organisation apparently results from
changed conditions of life. Disuse has reduced certain
parts of the body. Correlation of growth so ties the or-
ganisation together, that when one part varies other parts
vary at the same time. When several breeds have once
been formed, their intercrossing aids the progress of modi-
fication, and has even produced new sub-breeds. But as,
in the construction of a building, mere stones or bricks
are of little avail without the builder's art, so, in the
pro-
duction of new races, selection has been the presiding
power. Fanciers can act by selection on excessively
slight individual differences, as well as on those greater
differences which are called sports. Selection is followed
methodically when the fancier tries to improve and modi-
fy a breed according to a prefixed standard of excellence;
or he acts unmethodically and unconsciously, by merely
trying to rear as good birds as he can, without any wish
or intention to alter the breed. The progress of selection
almost inevitably leads to the neglect and ultimate extinc-
tion of the earlier and less improved forms, as well as of
many intermediate links in each long line of descent,
Thus it has come to pass that most of our present races
are so marvellously distinct from each other, and from the
aboriginal rock-pigeon.

CHAP. VII.
273
FOWLS.
.
CHAPTER VII.
FOWLS.
BRIEF DESCRIPTIONS OF THE CHIEF BREEDS ARGUMENTS IN FA-
VOUR OF THEIR DESCENT FROM SEVERAL SPECIES ARGUMENTS
IN FAVOUR OF ALL THE BREEDS HAVING DESCENDED FROM GAL-
LUS BANKIVA-KEVERSION TO THE PARENT-STOCK IN COLOUR
ANALOGOUS VARIATIONS - ANCIENT HISTORY OF THE FOWL-EX-
TERNAL DIFFERENCES BETWEEN THE SEVERAL BREEDS — EGGS
CHICKENS SECONDARY SEXUAL CHARACTERS - WING- AND
TAIL-FEATHERS, VOICE, DISPOSITION, ETC. OSTEOLOGICAL DIF-
FERENCES IN THE SKULL, VERTEBRÆ, ETC. — EFFECTS OF USE
AND DISUSE ON CERTAIN PARTS — CORRELATION OF GROWTH.
As some naturalists may not be familiar with the chief
breeds of the fowl, it will be advisable to give a con-
densed description of them. From what I have read and
seen of specimens brought from several quarters of the
world, I believe that most of the chief kinds have been
imported into England, but many sub-breeds are proba-
bly still here unknown. The following discussion on the
origin of the various breeds and on their characteristic
differences does not pretend to completeness, but may be
of some interest to the naturalist. The classification of
the breeds cannot, as far as I can see, be made natural.
1 I have drawn up this brief synopsis
from various sources, but chiefly from
information given me by Mr. Tegetmeier.
This gentleman has kindly looked
through the whole of this chapter, and
from his well-known knowledge, the
statements here given may be fully trust-
ed. Mr. Tegetmeier has likewise assist-
ed me in every possible way in obtaining
for me information and specimens. I
must not let this opportunity pass with-
out expressing my cordial thanks to Mr.
B. P. Brent, a well-known writer on poul-
try, for indefatigable assistance and the
gift of many specimens.
12*

274
CHAP. VII.
FOWLS.
They differ from each other in different degrees, and do
not afford characters in subordination to each other, by
which they can be ranked in group under group. They
seem all to have diverged by independent and different
Fig. 50.-Spanish Fowl.
roads from a single type. Each chief breed includes dif-
ferently coloured sub-varieties, most of which can be truly
propagated, but it would be superfluous to describe them.
I have classed the various crested fowls as sub-breeds
under the Polish fowl; but I have great doubts whether

CHAP. VII.
275
DESCRIPTION OF BREEDS.
this is a natural arrangement, showing true affinity or
blood relationship. It is scarcely possible to avoid laying
stress on the commonness of a breed ; and if certain for-
eign sub-breeds had been largely kept in this country they
would perhaps have been raised to the rank of main-
breeds. Several breeds are abnormal in character; that
is, they differ in certain points from all wild Gallinaceous
birds. At first I made a division of the breeds into nor-
mal and abnormal, but the result was wholly unsatisfac-
tory.
1. GAME BREED.—This may be considered as the typical breed,
as it deviates only slightly from the wild Gallus bankiva, or, as per-
haps more correctly named, ferrugineus. Beak strong; comb single
and upright. Spurs long and sharp. Feathers closely adpressed
to the body. Tail with the normal number of 14 feathers. Eggs
often pale-buff. Disposition indomitably courageous, exhibited even
in the hens and chickens. An unusual number of differently col-
oured varieties exist, such as black and brown-breasted reds, duck-
wings, blacks, whites, piles, &c., with their legs of various colours.
2. MALAY BREED.—Body of great size, with head, neck, and legs
elongated; carriage erect; tail small, sloping downwards, generally
formed of 16 feathers; comb and wattle small; ear-lobe and face
red ; skin yellowish ; feathers closely adpressed to the body; neck-
hackles short, narrow, and hard: Eggs often pale buff. Chickens
feather late. Disposition savage. Of Eastern origin.
3. COCHIN, OR SHANGAI BREED.-Size great; wing-feathers short,
arched, much hidden in the soft downy plumage ; barely capable of
flight ; tail short, generally formed of 16 feathers, developed at a
late period in the young males; legs thick, feathered ; spurs short,
thick; nail of middle toe flat and broad; an additional toe not
rarely developed ; skin yellowish. Comb and wattle well devel-
oped. Skull with deep medial furrow; occipital foramen, sub-
triangular, vertically elongated. Voice peculiar. Eggs rough, buff-
coloured. Disposition extremely quiet. Of Chinese origin.
4. DORKING BREED.—Size great; body square, compact ; feet
with an additional toe; comb well developed, but varies much in
form ; wattles well developed ; colour of plumage various. Skull
remarkably broad between the orbits. Of English origin.
The white Dorking may be considered as a distinct sub-breed,
being a less massive bird.
5. SPANISH BREED.—Tall, with stately carriage; tarsi long: comb

276
CHAP. VII.
FOWLS.
single, deeply serrated, of immense size; wattles largely developed ;
the large ear-lobes and sides of face white. Plumage black glossed
with green. Do not incubate. Tender in constitution, the comb
being often injured by frost. Eggs white, smooth, of large size.
Chickens feather late, but the young cocks show their masculine
characters, and crow at an early age. Of Mediterranean origin.
The Andalusians may be ranked as a sub-breed: they are of a
slaty blue colour, and their chickens are well feathered. A smaller,
short-legged Dutch sub-breed has been described by some authors
as distinct.
6. HAMBURGH BREED (fig. 31).-Size moderate ; comb flat, pro-
duced backwards, covered with numerous small points; wattle of
moderate dimensions; ear-lobe white; legs blueish, thin. Do not
incubate. Skull, with the tips of the ascending branches of the
premaxillary and with the nasal bones standing a little separate
from each other; anterior margin of the frontal bones less de-
pressed than usual.
There are two sub-breeds; the spangled Hamburgh, of English
origin, with the tips of the feathers marked with a dark spot; and
the pencilled Hamburgh, of Dutch origin, with dark transverse lines
across each feather, and with the body rather smaller. Both these
sub-breeds include gold and silver varieties, as well as some other
sub-varieties. Black Hamburghs have been produced by a cross
with the Spanish breed.
7. CRESTED OR POLISH BREED (fig. 32).-Head with a large,
rounded crest of feathers, supported on a hemispherical protube-
rance of the frontal bones, which includes the anterior part of the
brain. The ascending branches of the premaxillary bones and the
inner nasal processes are much shortened. The orifice of the nos-
trils raised and crescentic. Beak short. Comb absent, or small and
of crescentic shape; wattles either present or replaced by a beard.
like tuft of feathers. Legs leaden-blue. Sexual differences appear
late in life. Do not incubate. There are several beautiful varieties
which differ in colour and slightly in other respects.
The following sub-breeds agree in having a crest, more or less
developed, with the comb, when present, of crescentic shape. The
skull presents nearly the same remarkable peculiarities of structure
as in the true Polish fowl.
Sub-breed (a) Sultans.-A Turkish breed, resembling white Polish
fowls, with a large crest and beard, with short and well-feathered
legs. The tail is furnished with additional sickle feathers. Do not
incubate 2
2 The best account of Sultans is by
Miss Watts in The Poultry Yard,' 1856,
p. 79. I owe to Mr. Brent's kindness
the examination of some specimens of
this breed.

CHAP. VII.
277
DESCRIPTION OF BREEDS.
Sub-breed (6) Ptarmigans.-An inferior breed closely allied to the
last, white, rather small, legs much feathered, with the crest pointed ;
comb small, cupped ; wattles small.
Fig. 31.-Hamburgh Fowl.
Sub-breed (c) Ghoondooks.—Another Turkish breed having an ex-
traordinary appearance; black and tailless; crest and beard large;
legs feathered. The inner processes of the two nasal bones come
into contact with each other, owing to the complete absorption of
the ascending branches of the premaxillaries. I have seen an allied,
white, tailless breed from Turkey.
Sub-breed (d) Crève-cour.-A French breed of large size, barely
capable of flight, with short black legs, head crested, comb produced
into two points or horns, sometimes a little branched like the horns
of a stag; both beard and wattles present. Eggs large. Disposi-
tion quiet.
3 A good description with figures is given of this sub-breed in the Journal of Hor-
ticulture,' June 10th, 1862, p. 206.

278
CHAP. VII.
FOWLS.
Sub-breed (e) Horned fowl.—With a small crest; comb produced
into two great points, supported on two bony protuberances.
Sub-breed (f) Houdan.- A French breed ; of moderate size, short-
legged with five toes, wings well developed ; plumage invariably
mottled with black, white, and straw-yellow; head furnished with
a crest, and a triple comb placed transversely; both wattles and
beard present.
Sub-breed (9) Guelderlands.- No comb, head said to be surmoun-
ted by a longitudinal crest of soft velvety feathers; nostrils said to
be crescentic; wattles well developed ; legs feathered ; colour black.
From North America. The Breda fowl seems to be closely allied to
the Guelderland.
8. BANTAM BREED. — Originally from Japan, characterized by
small size alone; carriage bold and erect. There are several sub-
breeds, such as the Cochin, Game, and Sebright Bantams, some of
which have been recently formed by various crosses. The Black
Bantam has a differently shaped skull, with the occipital foramen like
that of the Cochin fowl.
9. RUMP-LESS FOWLs.-- These are so variable in character that
they hardly deserve to be called a breed. Any one who will exam-
ine the caudal vertebrae will see how monstrous the breed is.
10. CREEPERS OR JUMPERS.-These are characterized by an al-
most monstrous shortness of legs, so that they move by jumping
rather than by walking; they are said not to scratch up the ground.
I have examined a Burmese variety, which had a skull of rather un-
usual shape.
11. FRIZZLED OR CAFFRE FOWLS.-Not uncommon in India, with
the feathers curling backwards, and with the primary feathers of the
wing and tail imperfect; periosteum of bones black.
12. SILK FOWLS.-Feathers silky, with the primary wing and tail-
feathers imperfect; skin and periosteum of bones black; comb and
wattles dark leaden-blue; ear lappets tinged with blue; legs thin,
often furnished with an additional toe. Size rather small.
13. SOOTY FOWLs. - An Indian breed, of a white colour stained
with soot, with black skin and periosteum. The hens alone are thus
characterized.
From this synopsis we see that the several breeds differ
4 A description, with figures, is given
of this breed in Journal of Horticul-
ture,' June 3rd, 1862, p. 186. Some
writers describe the comb as two-horned.
6 Mr. Crawfurd, 'Descript. Dict of the
Indian Islands,' p. 113. Bantams are
mentioned in an ancient native Japanese
Encyclopædia, as I am informed by Mr.
Birch of the British Museum.
8 Ornamental and Domestic Poul-
try,' 1848.

CHAP. VII.
279
DESCRIPTION OF BREEDS.
considerably, and they would have been nearly as inter
esting for us as pigeons, if there had been equally good
Fig. 32.-Polish Fowl.
evidence that all had descended from one parent-species.
Most fanciers believe that they are descended from sev-
eral primitive stocks. The Rev. E. S. Dixon" argues
strongly on this side of the question; and one fancier
even denounces the opposite conclusion by asking, “Do
we not perceive pervading this spirit, the spirit of the
Ornamental and Domestic Poultry,' 1848.

280
CHAP. VII,
FOWLS.
Deist?” Most naturalists, with the exception of a few,
such as Temminck, believe that all the breeds have pro-
ceeded from a single species; but authority on such a
point goes for little. Fanciers look to all parts of the
world as the possible sources of their unknown stocks ;
thus ignoring the laws of geographical distribution. They
know well that the several kinds breed truly even in
colour. They assert, but, as we shall see, on very weak
grounds, that most of the breeds are extremely ancient.
They are strongly impressed with the great difference
between the chief kinds, and they ask with force, can dif-
ferences in climate, food, or treatment have produced
birds so different as the black stately Spanish, the dimi-
nutive elegant Bantam, the heavy Cochin with its many
peculiarities, and the Polish fowl with its great top-knot
and protuberant skull? But fanciers, whilst admitting
and even overrating the effects of crossing the various
breeds, do not sufficiently regard the probability of the
occasional birth, during the course of centuries, of birds
with abnormal and hereditary peculiarities; they overlook
the effects of correlation of growth of the long-continued
use and disuse of parts, and of some direct result from
changed food and climate, though on this latter head I
have found no sufficient evidence; and lastly, they all,
as far as I know, entirely overlook the all-important sub-
ject of unconscious or unmethodical selection, though
they are well aware that their birds differ individually,
and that by selecting the best birds for a few generations
they can improve their stocks.
An amateur writes 8 as follows. “The fact that poul-
try' have until lately received but little attention at the
hands of the fancier, and been entirely confined to the
domains of the producer for the market, would alone
suggest the improbability of that constant and unremit-
ting attention having been observed in breeding, which
8 Ferguson's Illustrated Series of Rare and Prize Poultry,' 1854, p. vi., Preface.

CIAP. VII.
281
THEIR PARENTAGE.
is requisite to the consummating, in the offspring of any
two birds, transmittable forms no texhibited by the pa-
rents.” This at first sight appears true. But in a future
chapter on Selection, abundant facts will be given show-
ing not only that careful breeding, but that actual selec-
tion was practised during ancient periods, and by barely
civilised races of man. In the case of the fowl I can ad-
duce no direct facts showing that selection was ancient-
ly practised; but the Romans at the commencement of
the Christian era kept six or seven breeds, and Columella
“particularly recommends as the best, those sorts that
have five toes and white ears.” In the fifteenth century
several breeds were known and described in Europe; and
in China, at nearly the same period, seven kinds were
named. A more striking case is that at present, in one
of the Philippine Islands, the semi-barbarous inhabitants
have distinct native names for no less than nine sub-
breeds of the Game Fowl.º Azara, who wrote towards
the close of the last century, states that in the interior
parts of South America, where I should not have ex-
pected that the least care would have been taken of
poultry, a black-skinned and black-boned breed is kept,
m being considered fertile and its flesh good for sick
persons. Now every one who has kept poultry knows
how impossible it is to keep several breeds distinct unless
the utmost care be taken in separating the sexes. Will
it then be pretended that those persons who in ancient
times and in semi-civilised countries took pains to keep
the breeds distinct, and who therefore valued them,
would not occasionally have destroyed inferior birds and
occasionally have preserved their best birds? This is all
that is required. It is not pretended that any one in an-
9 Rev. E. S. Dixon, in his Ornamen-
tal Poultry,' p. 203, gives an account of
Columella's work.
10 Mr. Crawfurd On the Relation of
the Domesticated Animals to Civiliza-
tion,' separately printed, p. 6; first read
before the Brit. Assoc. at Oxford, 1869.
11 Quadrupèdes du Paraguay,' tom.
ii. p. 324.

282
CHAP. VII.
FOWLS.
cient times intended to form a new breed, or to modify
an old breed according to some ideal standard of excel-
lence. He who cared for poultry would merely wish to
obtain, and afterwards to rear, the best birds which he
could ; but this occasional preservation of the best birds
would in the course of time modify the breed, as surely,
though by no means as rapidly, as does methodical selec-
tion at the present day. If one person out of a hundred
or out of a thousand attended to the breeding of his
birds, this would be sufficient; for the birds thus tended
would soon become superior to others, and would form
a new strain; and this strain would, as explained in the
last chapter, slowly have its characteristic differences
augmented, and at last be converted into a new sub-
breed or breed. But breeds .would often be for a time
neglected and would deteriorate; they would, however,
partially retain their character, and afterwards might
again come into fashion and be raised to a standard of
perfection higher than their former standard ; as has ac-
tually occurred quite recently with Polish fowls. If, how-
ever, a breed were utterly neglected, it would become
extinct, as has recently happened with one of the Polish
sub-breeds. Whenever in the course of past centuries a
bird appeared with some slight abnormal structure, such
as with a lark-like crest on its head, it would probably
often have been preserved from that love of novelty
which leads some persons in England to keep rumpless
fowls, and others in India to keep frizzled fowls. And
after a time any such abnormal appearance would be
carefully preserved, from being esteemed a sign of the
purity and excellence of the breed; for on this principle
the Romans eighteen centuries ago valued the fifth toe
and the white ear-lobe in their fowls.
Thus from the occasional appearance of abnormal cha-
racters, though at first only slight in degree; from the
effects of the use and the disuse of parts; possibly from
the direct effects of changed climate and food; from cor-

Crap. VII.
283
THEIR PARENTAGE.
relation of growth; from occasional reversions to old
and long-lost characters; from the crossing of breeds,
when more than one had once been formed; but, above
all, from unconscious selection carried on during many
generations, there is no insuperable difficulty, to the best
of my judgment, in believing that all the breeds have
descended from some one parent-source. Can any sin-
gle species be named from which we may reasonably
suppose that all have descended? The Gallus bankiva
apparently fulfils every requirement. I have already
given as fair an account as I could of the arguments in
favour of the multiple origin of the several breeds; and
now I will give those in favour of their common descent
from G. bankiva.
But it will be convenient first briefly to describe all the known
species of Gallus. The G. Sonneratii does not range into the north-
ern parts of India ; according to Colonel Sykes,12 it presents at dif-
ferent heights on the Ghauts, two strongly marked varieties, perhaps
deserving to be called species. It was at one time thought to be the
primitive stock of all our domestic breeds, and this shows that it
closely approaches the common fowl in general structure; but its
hackles partially consist of highly peculiar, horny laminæ, trans-
versely banded with three colours; and I have met with no au-
thentic account of any such character having been observed in any
domestic breed.13 This species also differs greatly from the com-
mon fowl, in the comb being finely serrated, and in the loins being
destitute of true hackles. Its voice is utterly different. It crosses
readily in India with domestic hens; and Mr. Blyth 14 raised nearly
100 hybrid chickens; but they were tender and mostly died whilst
young. Those which were reared were absolutely sterile when
crossed inter se or with either parent. At the Zoological Gardens,
however, some hybrids of the same parentage were not quite so
sterile: Mr. Dixon, as he informed me, made, with Mr. Yarrell's
aid, particular inquiries on this subject, and was assured that out
12 'Proc. Zoloog. Soc. 1832, p. 151.
13 I have examined the feathers of
some hybrids raised in the Zoological
Garders between the male G. Son-
neratii and a red game-hen, and these
feathers exhibited the true character of
those of G. Sonneratii, except that the
horny laminæ were much smaller.
14 See also an excellent letter on the
Poultry of India, by Mr. Blyth, in
Gardener's Chronicle,' 1851, p. 619.

284
CHAP. VII.
FOWLS.
of 50 eggs only five or six chickens were reared. Some, however,
of these half-bred birds were crossed with one of their parents,
namely, a Bantam, and produced a few extremely feeble chickens.
Mr. Dixon also procured some of these same birds and crossed them
in several ways, but all were more or less infertile. Nearly similar
experiments have recently been tried on a great scale in the Zoolo-
gical Gardens with almost the same result.16 Out of 500 eggs,
,
raised from various first crosses and hybrids, between G. Sonneratii,
bankiva, and varius, only 12 chickens were reared, and of these only
three were the product of hybrids inter se. From these facts, and
from the above-mentioned strongly-marked differences in structure
between the domestic fowl and G. Sonneratii, we may reject this
latter species as the parent of any domestic breed.
Ceylon possesses a fowl peculiar to the island, viz. G. Stanleyic;
this species approaches so closely (except in the colouring of the
comb) to the domestic fowl, that Messrs. E. Layard and Kellaert 16
would have considered it, as they inform me, as one of the parent-
stocks, had it not been for its singularly different voice. This bird,
like the last, crosses readily with tame hens, and even visits solitary
farms and ravishes them. Two hybrids, a male and female, thus
produced, were found by Mr. Mitford to be quite sterile: both in-
herited the peculiar voice of G. Stanleyii. This species, then, may
in all probability be rejected as one of the primitive stocks of the
domestic fowl.
Java and the islands eastward as far as Flores are inhabited by
G. varius (or furcatus), which differs in so many characters-green
plumage, unserrated comb, and single median wattle—that no
one supposes it to have been the parent of any one of our breeds;
yet, as I am informed by Mr. Crawfurd, 17 hybrids are commonly
raised between the male G. varius and the common hen, and are
kept for their great beauty, but are invariably sterile; this, how-
ever, was not the case with some bred in the Zoological Gardens.
These hybrids were at one time thought to be specifically distinct,
and were named G. aneus. Mr. Blyth and others believe that the
G. Temminckile (of which the history is not known) is a similar
hybrid. Sir J. Brooke sent me some skins of domestic fowls from
Borneo, and across the tail of one of these, as Mr, Tegetmeier ob-
served, there were transverse blue bands like those which he had
15 Mr. S. J. Salter, in Natural His- 17 See also Mr. Crawfurd's Descrip-
tory Review,' April, 1863, p. 276.
tive Dict. of the Indian Islands,' 1856,
16 See also Mr. Layard's paper in p. 113.
Annals and Mag. of Nat. History, 18 Described by Mr. G. R. Gray,
2nd series, vol. xiv. p. 62.
Proc. Zoolog. Soc.', 1949, p. 62.

CPAP. VII.
285
THEIR PARENTAGE.
seen on the tail-feathers of hybrids from G. varius, reared in the
Zoological Gardens. This fact apparently indicates that some of
the fowls of Borneo have been slightly affected by crosses with G.
varius, but the case may possibly be one of analogous variation. I
may just allude to the G. giganteus, so often referred to in works on
poultry as a wild species; but Marsden, the first describer, speaks
of it as a tame breed; and the specimen in the British Museum
evidently has the aspect of a domestic variety.
The last species to be mentioned, namely, Gallus bankiva, has a
much wider geographical range than the three previous species; it
inhabits Northern India as far west as Sinde, and ascends the Hi-
malaya to a height of 4000 ft.; it inhabits Burmah, the Malay pe-
ninsula, the Indo-Chinese countries, the Philippine Islands, and the
Malayan archipelago as far eastward as Timor. This species varies
considerably in the wild state. Mr. Blyth informs me that the speci-
mens, both male and female, brought from near the Himalaya, are
rather paler coloured than those from other parts of India; whilst
those from the Malay peninsula and Java are brighter coloured than
the Indian birds. I have seen specimens from these countries, and
the difference of tint in the hackles was conspicuous. The Malayan
hens were a shade redder on the breast and neck than the Indian
hens. The Malayan males generally had a red ear-lappet, instead of
a white one as in India ; but Mr. Blyth has seen one Indian specimen
without the white ear-lappet. The legs are leaden blue in the Indi-
an, whereas they show some tendency to be yellowish in the Malayan
and Javan specimens. In the former Mr. Blyth finds the tarsus re-
markably variable in length. According to Temminck 20 the Timor
specimens differ as a local race from that of Java. These several
wild varieties have not as yet been ranked as distinct species; if
they should, as is not unlikely, be hereafter thus ranked, the cir-
cumstance would be quite immaterial as far as the parentage and dif-
ferences of our domestic breeds are concerned. The wild G. bankiva
agrees most closely with the black-breasted red Game-breed, in co-
louring and in all other respects, except in being smaller, and in the
tail being carried more horizontally. But the manner in which the
tail is carried is highly variable in many of our breeds, for, as Mr.
Brent informs me, the tail slopes much in the Malays, is erect in
the Games and some other breeds, and is more than erect in Dork-
ings, Bantams, &c. There is one other difference, namely, that in
19 The passage from Marsden is
given by Mr. Dixon in his 'Poultry
Book,' p. 176. No ornithologist now
ranks this bird as a distinct species.
20 Coup-d'oeil général sur l'Inde
Archipélagique,' tom. iii. (1849), p. 177;
see also Mr. Blyth in Indian Sporting
Review,' vol. ii. p. 5, 1856.

286
CHAP. VII.
FOWLS.
G. bankiva, according to Mr. Blyth, the neck-hackles when first
moulted are replaced during two or three months, not by other
hackles, as with our domestic poultry, but by short blackish feath-
ers.21 Mr. Brent, however, has remarked that these black feathers
remain in the wild bird after the development of the lower hackles,
and appear in the domestic bird at the same time with them ; so
that the only difference is that the lower hackles are replaced more
slowly in the wild than in the tame bird; but as confinement is
known sometimes to affect the masculine plumage, this slight dif-
ference cannot be considered of any importance. It is a significant
fact that the voice of both the male and female G. banliva closely re-
sembles, as Mr. Blyth and others have noted, the voice of both sexes
of the common domestic fowl; but the last note of the crow of the
wild bird is rather less prolonged. Captain Hutton, well known for
his researches into the natural history of India, informs me that he
has seen several crossed fowls from the wild species and the Chinese
bantam ; these crossed fowls bred freely with bantams, but unfortu-
nately were not crossed inter se. Captain Hutton reared chickens
from the eggs of the Gallus bankiva ; and these, though at first
very wild, afterwards became so tame that they would crowd round
his feet. He did not succeed in rearing them to maturity; but, as he
remarks, “no wild gallinaceous bird thrives well at first on hard
grain.” Mr. Blyth also found much difficulty in keeping G.bankiva
in confinement. In the Philippine Islands, however, the natives
must succeed better, as they keep wild cocks to fight with their do-
mestic game-birds.22 Sir Walter Elliot informs me that the hen of
a native domestic breed of Pegu is undistinguishable from the hen
of the wild G. bankioa, and the natives constantly catch wild cocks
by taking tame cocks to fight with them in the woods.23 Mr. Craw-
furd remarks that from etymology it might be argued that the fowl
was first domesticated by the Malays and Javanese 24 It is also a
curious fact, of which I have been assured by Mr. Blyth, that wild
specimens of the Gallus bankiva, bronght from the countries east of
the Bay of Bengal, are far more easily tamed than those of India ;
nor is this an unparalleled fact, for, as Humboldt long ago remarked,
the same species sometimes evinces a more tameable disposition in
one country than in another. If we suppose that the G. bankiva
was first tamed in Malaya and afterwards imported into India, we
21 Mr. Blyth, in 'Annals and Mag. of
Nat. Hist.,' 2nd ser., vol. i. (1848), p. 455.
22 Crawfurd. Desc. Dict. of Indian
Islands,' 1856, 112.
23 In Burmah, as I hear from Mr.
Blyth, the wild and tame poultry con-
stantly cross together, and irregular
transitional forms may be seen.
24 Idem, p. 113.

CHAP. VII,
287
THEIR PARENTAGE.
can understand an observation made to me by Mr. Blyth, that the
domestic fowls of India do not resemble the wild G. bankiva more
closely than do those of Europe.
25
From the extremely close resemblance in colour, general
structure, and especially in voice, between Gallus bankiva
and the Game fowl; from their fertility, as far as this
has been ascertained, when crossed; from the possibility
of the wild species being tamed, and from its varying in
the wild state, we may confidently look at it as the pa-
rent of the most typical of all the domestic breeds, namely,
the Game-fowl. It is a significant fact, that almost all
the naturalists in India, namely, Sir W. Elliot, Mr. S. N.
Ward, Mr. Layard, Mr. J. C. Jerdon, and Mr. Blyth,
who are familiar with G. bankiva, believe that it is the
parent of most or all our domestic breeds. But even if
it be admitted that G. bankiva is the parent of the Game
breed, yet it
may
be urged that other wild species have
been the parents of the other domestic breeds; and that
these species still exist, though unknown, in some coun-
try, or have become extinct. The extinction, however,
of several species of fowls, is an improbable hypothesis,
seeing that the four known species have not become ex-
tinct in the most anciently and thickly peopled regions of
the East. There is, in fact, only one kind of domesticated
bird, namely, the Chinese goose or Anser cygnoides; of
which the wild parent-form is said to be still unknown,
or extinct. For the discovery of new, or the rediscovery
of old species of Gallus, we must not look, as fanciers
often look, to the whole world. The larger gallinaceous
birds, as Mr. Blyth has remarked,20 generally have a re-
stricted range; we see this well illustrated in India, where
the
genus
Gallus inhabits the base of the Himalaya, and
25 Mr. Jerdon, in the 'Madras Journ.
of Lit. and Science,' vol. xxii. p. 2,
speaking of G. bankiva, says, "unques-
tionably the origin of most of the va-
rieties of our common fowls." For Mr.
Blyth, see his excellent article in Gar.
dener's Chron. 1851, p. 619; and in
Annals and Mag. of Nat. Hist.,' vol.
xx., 1847, p. 388.
26 Gardener's Chronicle,' 1851, p. 619.

288
CHAP. VII.
FOWLS.
is succeeded higher up by Gallophasis, and still higher
up by Phasianus. Australia, with its islands, is out of
the question as the home for unknown species of the
genus. It is, also, as improbable that Gallus should in-
habit South America 27 as that a humming-bird should be
found in the old world. From the character of the other
gallinaceous birds of Africa, it is not probable that Gal-
lus is an African genus. We need not look to the west-
ern parts of Asia, for Messrs. Blyth and Crawfurd, who
have attended to this subject, doubt whether Gallus ever
existed in a wild state even as far west as Persia. Al-
though the earliest Greek writers speak of the fowl as a
Persian bird, this probably merely indicates its line of
importation. For the discovery of unknown species we
must look to India, to the Indo-Chinese countries, and to
the northern parts of the Malay Archipelago. The south-
ern portion of China is the most likely country; but as
Mr. Blyth informs me, skins have been exported from
China during a long period, and living birds are largely
27 I have consulted an eminent autho-
rity, Mr. Sclater, on this subject, and he
thinks that I have not expressed myself
too strongly. I am aware that one an-
cient author, Acosta, speaks of fowls as
having inhabited S. America at the pe-
riod of its discovery, and more recently,
about 1795, Olivier de Serres speaks of
wild fowls in the forests of Guiana;
these were probably feral birds. Dr.
Daniell tells me, he believes that fowls
have become wild on the west coast of
Equatorial Africa; they may, however,
not be true fowls, but gallinaceous birds
belonging to the genus Phasidus. The
old voyager Barbut says that poultry
are not natural to Guinea. Capt. W.
Allen ("Narrative of Niger Expedition,'
1848, vol. ii. p. 42) describes wild fowls
on Ilha dos Rollas, an island near St.
Thomas's, on the west coast of Africa :
the natives informed him that they had
escaped from a vessel wrecked there
many years ago; they were extremely
wild, and had "a cry quite different to
that of the domestic fowl," and their ap-
pearance was somewhat changed. Hence
it is not a little doubtful, notwithstand-
ing the statement of the natives, wheth-
er these birds really were fowls. That
the fowl has become feral on several is-
lands is certain. Mr. Fry, a very capa-
ble judge, informed Mr. Layard, in a let-
ter, that the fowls which have run wild
on Ascension “had nearly all got back
to their primitive colours, red and black
cocks, and smoky-grey hens." But un-
fortunately we do not know the colour
of the poultry which were turned out.
Fowls have become feral on the Nicobar
Islands (Blyth in the Indian Field,'
1858, p. 62), and in the Ladrones (An-
son's Voyage). Those found in the Pel-
lew Islands (Crawfurd) are believed to
be feral; and lastly, it is asserted that
they have become feral in New Zealand,
but whether this is correct I know not.

CHAP. VII.
289
THEIR PARENTAGE.
kept there in aviaries, so that any native species of Gal-
lus would probably have become known. Mr. Birch, of
the British Museum, has translated for me passages from
a Chinese Encyclopædia published in 1609, but compiled
from more ancient documents, in which it is said that
fowls are creatures of the West, and were introduced
into the East (i.e. China) in a dynasty 1400 B.C. What-
ever may be thought of so ancient a date, we see that the
Indo-Chinese and Indian regions were formerly consider-
ed by the Chinese as the source of the domestic fowl.
From these several considerations we must look to the
present metropolis of the genus, namely, to the south-
eastern parts of Asia, for the discovery of species which
were formerly domesticated, but are now unknown in the
wild state; and the most experienced ornithologists do
not consider it probable that such species will be dis-
covered.
In considering whether the domestic breeds are de-
scended from one species, namely, G. bankiva, or from
several, we must not quite overlook, though we must not
exaggerate, the importance of the test of fertility. Most
of our domestic breeds have been so often crossed, and
their mongrels so largely kept, that it is almost certain,
if any degree of infertility had existed between them, it
would have been detected. On the other hand, the four
known species of Gallus when crossed with each other,
or when crossed, with the exception of G. bankiva, with
the domestic fowl, produce infertile hybrids.
Finally, we have not such good evidence with fowls as
with pigeons, of all the breeds having descended from a
single primitive stock. In both cases the argument of
fertility must go for something; in both we have the im-
probability of man having succeeded in ancient times
in thoroughly domesticating several supposed species,-
most of these supposed species being extremely abnormal
as compared with their natural allies--all being now
either unknown or extinct, though the parent-form of
13

290
CHAP. VII
FOWLS.
scarcely any other domesticated bird has been lost. But
in searching for the supposed parent-stocks of the various
breeds of the pigeon, we were enabled to confine our
search to species having peculiar habits of life; whilst
with fowls there is nothing in their habits in any marked
manner distinct from those of other gallinaceous birds.
In the case of pigeons, I have shown that purely-bred
birds of every race and the crossed offspring of distinct
races frequently resemble, or revert to, the wild rock-
pigeon in general colour and in each characteristic mark.
With fowls we have facts of a similar nature, but less
strongly pronounced, which we will now discuss.
Reversion and Analogous Variation. — Purely-bred
Game, Malay, Cochin, Dorking, Bantam, and, as I hear
from Mr. Tegetmeier, Silk fowls, may frequently or occa.
sionally be met with, which are almost identical in plu-
mage with the wild G. bankiva. This is a fact well de-
serving attention, when we reflect that these breeds rank
amongst the most distinct. Fowls thus coloured are
called by amateurs black-breasted reds. Hamburghs
properly have a very different plumage ; nevertheless, as
Mr. Tegetmeier informs me, "the great difficulty in breed-
ing cocks of the golden-spangled variety is their tenden-
cy to have black breasts and red backs." The males
of white Bantams and white Cochins, as they come to
maturity, often assume a yellowish or saffron tinge; and
the longer neck hackles of black bantam cocks, 2 when
two or three years old, not uncommonly become ruddy;
these latter bantams occasionally “even moult brassy
winged, or actually red shouldered.” So that in these
several cases we see a plain tendency to reversion to the
hues of G. bankiva, even during the lifetime of the indi-
vidual bird. With Spanish, Polish, pencilled Hamburgh,
silver-spangled Hamburgh fowls, and with some other
28 Mr. Hewitt, in 'The Poultry Book,' by W. B. Teget meier, 1866, p. 248.

CHAP. VII.
291
REVERSION AND VARIATION.
less common breeds, I have never heard of a black-breast-
ed red bird having appeared.
From my experience with pigeons, I made the follow-
ing crosses. I first killed all my own poultry, no others
living near my house, and then procured, by Mr. Teget-
meier's assistance, a first-rate black Spanish cock, and
hens of the following pure breeds, -white Game, white
Cochin, silver-spangled Polish, silver-spangled Hamburgh,
silver-pencilled Hamburgh, and white Silk. In none of
these breeds is there a trace of red, nor when kept pure
have I ever heard of the appearance of a red feather;
though such an occurrence would perhaps not be very
improbable with white Games and white Cochins. Of
the many chickens reared from the above six crosses the
majority were black, both in the down and in the first
plumage; some were white, and a very few were mottled
black and white. In one lot of eleven mixed eggs from
the white Game and white Cochin by the black Spanish
cock, seven of the chickens were white, and only four
black: I mention this fact to show that whiteness of plu-
mage is strongly inherited, and that the belief in the pre-
potent power in the male to transmit his colour is not
always correct. The chickens were hatched in the spring,
and in the latter part of August several of the young
cocks began to exhibit a change, which with some of
them increased during the following years. Thus a young
male bird from the silver-spangled Polish hen was in its
first plumage coal-black, and combined in its comb, crest,
wattle, and beard, the characters of both parents; but
when two years old the secondary wing-feathers, became
largely and symmetrically marked with white, and
wherever in G. bankiva the hackles are red, they were
in this bird greenish-black along the shaft, narrowly
bordered with brownish-black, and this again broadly
bordered with very pale yellowish-brown; so that in
general appearance the plumage had become pale-colour-
ed instead of black. In this case, with advancing age

292
CHAP. VI.
FOWLS.
there was a great change, but no reversion to the red
colour of G. bankiva.
A cock with a regular rose comb derived either from
the spangled or pencilled silver Hamburgh was likewise
at first quite black; but in less than a year the neck-
hackles, as in the last case, became whitish, whilst those
on the loins assumed a decided reddish-yellow tint; and
here we see the first symptom of reversion; this likewise
occurred with some other young cocks, which need not
here be described. It has also been recorded” by a
breeder, that he crossed two silver-pencilled Hamburgh
hens with a Spanish cock, and reared a number of chick-
ens, all of which were black, the cocks having golden and
the hens brownish hackles; so that in this instance like-
wise there was a clear tendency to reversion.
Two young cocks from my white Game hen were at
first snow white; of these, one subsequently assumed
pale orange-coloured hackles, chiefly on the loins, and
the other an abundance of fine orange-red hackles on the
neck, loins, and upper wing-coverts. Here again we
have a more decided, though partial, reversion to the
colours of G. bankiva. This second cock was in fact
coloured like an inferior “ pile Game cock ;" —now this
sub-breed can be produced, as I am informed by Mr. Te-
getmeier, by crossing a black-breasted red Game cock with
a white Game hen, and the “pile” sub-breed thus pro-
duced can afterwards be truly propagated. So that we
have the curious fact of the glossy-black Spanish cock
and the black-breasted red Game cock when crossed with
white Game-hens producing offspring of nearly the same
colours.
I reared several birds from the white Silk-hen by the
Spanish cock: all were coal-black, and all plainly showed
their parentage in having blackish combs and bones;
none inherited the so-called silky feathers, and the non-
29 Journal of Horticulture," Jan. 14th, 1862, p. 325.

CHAP. VII.
293
REVERSION AND VARIATION.
inheritance of this character bas been observed by others.
The hens never varied in their plumage. As the young
cocks grew old, one of them assumed yellowish-white
hackles, and thus resembled in a considerable degree the
cross from the Hamburgh hen; the other became a gor-
geous bird, so much so that an acquaintance had it pre-
served and stuffed simply from its beauty. When stalk-
ing about it closely resembled the wild Gallus bankiva,
but with the red feathers rather darker.
On close com-
parison one considerable difference presented itself,
namely, that the primary and secondary wing-feathers
were edged with greenish-black, instead of being edged,
as in G.bankiva, with fulvous and red tints. The space,
also, across the back, which bears dark-green feathers,
was broader, and the comb was blackish. In all other
respects, even in trifling details of plumage, there was
the closest accordance. Altogether it was a marvellous
sight to compare this bird first with G. bankiva, and
then with its father, the glossy green-black Spanish cock,
and with its diminutive mother, the white Silk hen. This
case of reversion is the more extraordinary as the Span-
ish breed has long been known to breed true, and no in-
stance is on record of its throwing a single red feather.
The Silk hen likewise breeds true, and is believed to be
ancient, for Aldrovandi, before 1600, alludes probably to
this breed, and describes it as covered with wool. It is
so peculiar in many characters that some writers have
considered it as specifically distinct; yet, as we now see,
when crossed with the Spanish fowl, it yields offspring
closely resembling the wild G. bankiva.
Mr. Tegetmeier has been so kind as to repeat, at my
request, the cross between a Spanish cock and Silk hen,
and he obtained similar results; for he thus raised, be-
sides a black hen, seven cocks, all of which were dark-
bodied with more or less orange-red hackles. In the
ensuing year he paired the black hen with one of her
brothers, and raised three young cocks, all coloured like
their father, and a black hen mottled with white.

294
CAP. VII.
FOWLS.
The hens from the six above-described crosses showed
hardly any tendency to revert to the mottled-brown
plumage of the female G. bankiva: one hen, however,
from the white Cochin, which was at first coal-black, be-
came slightly brown or sooty. Several hens, which were
for a long time snow-white, acquired as they grew old a
few black feathers. A hen from the white Game, which
was for a long time entirely black glossed with green,
when two years old had some of the primary wing-feathers
greyish-white, and a multitude of feathers over her body.
narrowly and symmetrically tipped or laced with white.
I had expected that some of the chickens whilst covered
with down would have assumed the longitudinal stripes
so general with gallinaceous birds; but this did not occur
in a single instance. Two or three alone were reddish-
brown about their heads. I was unfortunate in losing
nearly all the white chickens from the first crosses ; so
that black prevailed with the grandchildren ; but they
were much diversified in colour, some being sooty, others
mottled, and one blackish chicken had its feathers oddly
tipped and barred with brown.
I will here add a few miscellaneous facts connected
with reversion, and with the law of analogous variation.
This law implies, as stated in a previous chapter, that
the varieties of one species frequently mock distinct but
allied species; and this fact is explained, according to
the views which I maintain, on the principle of allied
species having descended from one primitive form. The
white Silk fowl with black skin and bones degenerates,
as has been observed by Mr. Hewitt and Mr. R. Orton,
in our climate; that is, it reverts to the ordinary colour
of the common fowl in its skin and bones, due care
having been taken to prevent any cross.
In Germany
30
30 Die Hühner und Pfauenzucht.'
Ulm, 1827, s. 17. For Mr. Hewitt's state-
ment with respect to the white Silk fowl,
see the Poultry Book,' by W. B. Teget-
meier, 1866, p. 222. I am indebted to
Mr. Orton for a letter on the same sub-
ject.

CE:P. VII.
295
REVERSION AND VARIATION.
a distinct breed with black bones, and with black, not
silky plumage, has likewise been observed to degen-
erate.
Mr. Tegetmeier informs me that, when distinct breeds
are crossed, fowls are frequently produced with their
feathers marked or pencilled by narrow transverse lines
of a darker colour. This may be in part explained by
direct reversion to the parent-form, the Bankiva hen;
for this bird has all its upper plumage finely mottled
with dark and rufous brown, with the mottling partially
and obscurely arranged in transverse lines. But the
tendency to pencilling is probably much strengthened
by the law of analogous variation, for the bens of some
other species of Gallus are much more plainly pencilled,
and the hens of many gallinaceous birds belonging to
other genera, as the partridge, have pencilled feathers.
Mr. Tegetmeier has also remarked to me, that, although
with domestic pigeons we have so great a diversity of
colouring, we never see either pencilled or spangled
feathers; and this fact is intelligible on the law of ana-
logous variation, as neither the wild rock-pigeon nor any
closely-allied species has such feathers. The frequent
appearance of pencilling in crossed birds probably ac-
counts for the existence of " cuckoo” sub-breeds in the
Game, Polish, Dorking, Cochin, Andalusian, and Bantam
breeds. The plumage of these birds is slaty-blue or
grey, with each feather transversely barred with darker
lines, so as to resemble in some degree the plumage of
the cuckoo. It is a singular fact, considering that the
male of no species of Gallus is in the least barred, that
the cuckco-like plumage has often been transferred to
the male, more especially in the cuckoo Dorking; and
the fact is all the more singular, as in gold and silver
pencilled Hamburghs, in which pencilling is character-
istic of the breed, the male is hardly at all pencilled,
this kind of plumage being confined to the female.
Another case of analogous variation is the occurrence

296
CHAP. VIL
FOWLS.
of spangled sub-breeds of Hamburgh, Polish, Malay, and
Bantam fowls. Spangled feathers have a dark mark,
properly crescent-shaped, on their tips; whilst pencilled
feathers have several transverse bars. The spangling
cannot be due to reversion to G. bankiva; nor does it
often follow, as I hear from Mr. Tegetmeier, from cross-
ing distinct breeds; but it is a case of analogous varia-
tion, for many gallinaceous birds have spangled feathers,
-for instance, the common pheasant. Hence spangled
breeds are often called “pheasant”-fowls. Another case
of analogous variation in several domestic breeds is in-
explicable; it is, that the chickens, whilst covered with
down, of the black Spanish, black Game, black Polish,
and black Bantam, all have white throats and breasts,
and often have some white on their wings. The editor
of the ‘Poultry Chronicle' 32 remarks that all the breeds
which properly have red ear-lappets occasionally produce
birds with white ear-lappets. This remark more especial-
ly applies to the Game breed, which of all comes nearest
to the G. bankiva ; and we have seen that with this
species living in a state of nature, the ear-lappets vary
in colour, being red in the Malayan countries, and gen-
erally, but not invariably, white in India.
In concluding this part of my subject I may repeat
that there exists one widely-ranging, varying, and com-
mon species of Gallus, namely G. bankiva, which can be
tamed, produces fertile offspring when crossed with com-
mon fowls, and closely resembles in its whole structure,
plumage, and voice the Game breed; hence it may be
safely ranked as the parent of this, the most typical do-
mesticated breed. We have seen that there is much diffi-
culty in believing that other, now unknown, species have
been the parents of the other domestic breeds. We know
p. 260.
31 Dixon, Ornamental and Domestic
Poultry,' pp. 253, 324, 335. For game
fowls, see Ferguson on Prize Poultry,'
32 Poultry Chronicle,' vol.ii. p. 71.

CHAP. VII.
297
REVERSION AND VARIATION.
that all the breeds are most closely allied, as shown by
their similarity in most points of structure and in habits,
and by the analogous manner in which they vary. We
have also seen that several of the most distinct breeds
occasionally or habitually closely resemble in plumage G.
bankiva, and that the crossed offspring of other breeds,
which are not thus coloured, show a stronger or weaker
tendency to revert to this same plumage. Some of the
breeds, which appear the most distinct and the least likely
to have proceeded from G. bankiva, such as Polish fowls,
with their protuberant and little ossified skulls, and Co-
chins, with their imperfect tail and small wings, bearin
these characters the plain marks of their artificial origin.
We know well that of late years methodical selection has
greatly improved and fixed many characters; and we have
every reason to believe that unconscious selection, carried
on for many generations, will have steadily augmented
each new peculiarity and thus have given rise to new
breeds. As soon as two or three breeds had once been
formed, crossing would come into play in changing their
character and in increasing their number. Brahma Poo-
tras, according to an account lately published in America,
offer a good instance of a breed, lately formed by a cross,
which can be truly propagated. The well-known Sebright
Bantams offer another and similar instance. Hence it
may be concluded that not only the Game-breed but that
all our breeds are probably the descendants of the Ma-
layan or Indian variety of G. bankiva. If so, this species
has varied greatly since it was first domesticated; but
there has been ample time, as we shall now show.
History of the Fowl.-Rütimeyer found no remains of
the fowl in the ancient Swiss lake-dwellings. It is not
mentioned in the Old Testament; nor is it figured on the
ancient Egyptian monuments.*3 It is not referred to by
83 Dr. Pickering, in his ' Races of Man,'
1850, p. 374, says that the head and neck
of a fowl is carried in a Tribute-pro-
cession to thoutmousis III. (1445 B.C.);
but Mr. Birch of the British Museum
doubts whether the figure can be iden-
13*

298
CHAP. VII.
FOWLS,
Homer or Hesiod (about 900 B.C.); but is mentioned by
Theognis and Aristophanes between 400 and 500 B.C. It
is figured on some of the Babylonian cylinders, of which
Mr. Layard sent me an impression, between the sixth and
seventh centuries B.C.; and on the Harpy Tomb in Lycia,
about 600 B.C. : so that we may feel pretty confident that
the fowl reached Europe somewhere near the sixth century
B.C. It had travelled still farther westward by the time
of the Christian era, for it was found in Britain by Julius
Cæsar. In India it must have been domesticated when
the Institutes of Manu were written, that is, according to
Sir W. Jones, 1200 B.C., but, according to the later au-
thority of Mr. H. Wilson, only 800 B.C., for the domestic
fowl is forbidden, whilst the wild is permitted to be eaten.
If, as before remarked, we may trust the old Chinese En-
cyclopædia, the fowl must have been domesticated several
centuries earlier, as it is said to have been introduced
from the West into China 1400 B.C.
Sufficient materials do not exist for tracing the history
of the separate breeds. About the commencement of
the Christian era, Columella mentions a five-toed fighting
breed, and some provincial breeds; but we know nothing
more about them. He also alludes to dwarf fowls; but
these cannot have been the same with our Bantams,
which, as Mr. Crawfurd has shown, were imported from
Japan into Bantam in Java. A dwarf fowl, probably the
tified as the head of a fowl. Some cau-
tion is necessary with reference to the
absence of figures of the fowl on the
ancient Egyptian monuments, on account
of the strong and widely prevalent pre-
judice against this bird. I am informed
by the Rev. S. Erhardt that on the east
coast of Africa, from 4° to 6° south of the
equator, most of the pagan tribes at the
present day hold the fowl in a version.
The natives of the Pellew Islands would
not eat the fowl, nor will the Indians in
some parts of S. America. For the an-
cient history of the fowl, see also Volz,
Beitrage zur Culturgeschichte,' 1852, s.
77; and Isid. Geoffroy St. Hilaire, 'Hist.
Nat. Gen.,' tom. iii. p. 61. Mr. Crawfurd
has given an admirable history of the
fowl in his paper on the Relation of
Domesticated Animals to Civilization,'
read before the Brit. Assoc. at Oxford in
1860, and since printed separately. I
quote from him on the Greek poet The-
ognis, and on the Harpy Tomb described
by Sir. C. Fellowes. I quote from a letter
of Mr. Blyth's with respect to the Insti-
tutes of Manu.

CHAP. VII.
299
THEIR HISTORY.
true Bantam, is referred to in an old Japanese Encyclo-
pædia, as I am informed by Mr. Birch. In the Chinese En-
cyclopædia, published in 1596, but compiled from various
sources, some of high antiquity, seven breeds are mention-
ed, including what we should now call jumpers or creepers,
and likewise fowls with black feathers, bones, and flesh.
In 1600 Aldrovandi describes seven or eight breeds of
fowls, and this is the most ancient record from which the
age of our European breeds can be inferred. The Gallus
Turcicus certainly seems to be a pencilled Hamburgh;
but Mr. Brent, a most capable judge, thinks that Aldro-
vandi “evidently figured what he happened to see, and
not the best of the breed.” Mr. Brent, indeed, considers
all Aldrovandi's fowls as of impure breed; but it is a far
more probable view that all our breeds since his time
have been much improved and modified; for, as he went
to the expense of so many figures, he probably would
have secured characteristic specimens. The Silk fowl,
however, probably then existed in its present state, as did
almost certainly the fowl with frizzled or reversed feath-
ers. Mr. Dixon 34 considers Aldrovandi's Paduan fowl as
a variety of the Polish,” whereas Mr. Brent believes it
to have been more nearly allied to the Malay. The ana-
tomical peculiarities of the skull of the Polish breed were
noticed by P. Borelli in 1656. I may add that in 1737
one Polish sub-breed, viz. the golden spangled, was
known; but judging from Albin's description, the comb
was then larger, the crest of feathers much smaller, the
breast more coarsely spotted, and the stomach and thighs
much blacker: a golden-spangled Polish fowl in this
condition would now be of no value.
Differences in External and Internal Structure between
the Breeds : Individual Variability.-Fowls have been
exposed to diversified conditions of life, and as we have
34 "Ornamental and Domestic Poul-
try,' 1847, p. 185; for passages trans-
lated from Columella, see p. 312. For
Golden Hamburghs, see Albin's 'Natu.
ral History of Birds,'3 vols., with plates,
1731-38.

300
CHAP. VII.
FOWLS.
just seen there has been ample time for much variability
and for the slow action of unconscious selection. As
there are good grounds for believing that all the breeds
are descended from Gallus bankiva, it will be worth
while to describe in some detail the chief points of dif-
ference. Beginning with the eggs and chickens, I will
pass on to the secondary sexual characters, and then to
the differences in external structure and in the skeleton.
I enter on the following details chiefly to show how va-
riable almost every character has become under domesti-
cation.
Eggs.- Mr. Dixon remarks 35 that “to every hen belongs an indi-
vidual peculiarity in the form, colour, and size of her egg, which
never changes during her life-time, so long as she remains in health,
and which is as well known to those who are in the habit of taking
her produce, as the handwriting of their nearest acquaintance." I
believe that this is generally true, and that, if no great number of
hens be kept, the eggs of each can almost always be recognised
The eggs of differently sized breeds naturally differ much in size;
but, apparently, not always in strict relation to the size of the hen :
thus the Malay is a larger bird than the Spanish, but generally she
produces not such large eggs; white Bantams are said to lay
smaller eggs than other Bantams; 36 white Cochins, on the other
hand, as I hear from Mr. Tegetmeier, certainly lay larger eggs
than buff Cochins. The eggs, however, of the different breeds
vary considerably in character; for instance, Mr. Ballance states 37
that his Malay “pullets of last year laid eggs equal in size to those
of any duck, and other Malay hens, two or three years old, laid
eggs very little larger than a good-sized Bantam's egg. Some
were as white as a Spanish hen's egg, and others varied from a
light cream-colour to a deep rich buff, or even to a brown." The
shape also varies, the two ends being much more equally rounded
in Cochins than in Games or Polish. Spanish fowls lay smoother
eggs than Cochins, of which the eggs are generally granulated.
The shell in this latter breed, and more especially in Malays, is apt
to be thicker than in Games or Spanish ; but the Minorcas, a sub-
35
Ornamental and Domestic Poul-
try, 1. 152.
ever, figures and much information on
eggs. See pp: 34 and 235 on the eggs of
the Game fowl.
38 Ferguson on Rare Prize Poultry,'
p. 297. This writer, I am informed, can-
oot generally be trusted. He gives, how-
37 See · Poultry Book,' by Mr. Teget-
meier, 1866, pp. 81 and 78.

CHAP. VII. DIFFERENCES BETWEEN THE BREEDS. 301
breed of Spanish, are said to lay harder eggs than true Spanish.88
The colour differs considerably,--the Cochins laying buff-coloured
eggs; the Malays a paler variable buff; and Games a still paler
buff. It would appear that darker-coloured eggs characterise the
breeds which have lately come from the East, or are still closely
allied to those now living there. The colour of the yolk, according
to Ferguson, as well as of the shell, differs slightly in the sub-
breeds of the Game, and stands in some degree of correlation with
the colour of the plumage. I am also informed by Mr. Brent that
dark partridge-coloured Cochin hens lay darker coloured eggs than
the other Cochin sub-breeds. The flavour and richness of the egg
certainly differ in different breeds. The productiveness of the
several breeds is very different. Spanish, Polish, and Hamburgh
have lost the incubating instinct.
Chickens. - As the young of almost all gallinaceous birds, even of
the black curassow and black grouse, whilst covered with down,
are longitudinally striped on the back, -of which character, when
adult, neither sex retains a trace,-it might have been expected
that the chickens of all our domestic fowls would have been simi-
larly striped.38 This could, however, hardly have been expected,
when the adult plumage in both sexes has undergone so great a
change as to be wholly white or black. In white fowls of various
breeds the chickens are uniformly yellowish white, passing in the
black-boned Silk fowl into bright canary-yellow. This is also
generally the case with the chickens of white Cochins, but I hear
from Mr. Zurhost that they are sometimes of a buff or oak colour,
and that all those of this latter colour, which were watched, turned
out males. The chickens of buff Cochins are of a golden-yellow,
easily distinguishable from the paler tint of the white Cochins, and
are often longitudinally streaked with dark shades: the chickens of
silver-cinnamon Cochins are almost always of a buff colour. The
chickens of the white Game and white Dorking breeds, when held
in particular lights, sometimes exhibit (on the authority of Mr.
Brent) faint traces of longitudinal stripes. Fowls which are entirely
black, namely Spanish, black Game, black Polish, and black Ban-
tams, display a new character, for their chickens have their breasts
and throats more or less white, with sometimes a little white else-
98 The Cottage Gardener,' Oct. 1855,
p. 13. On the thinness of the eggs of
Game-fowls, see Mowbray on Poultry,
7th edit., p. 13.
39 My information, which is very far
from perfect, on chickens in the down,
is derived chiefly from Mr. Dixon's 'Or-
namental and Domestic Poultry.' Mr.
B. P. Brent has also communicated to
me many facts by letter, as has Mr.
Tegetmeier. I will in each case mark my
authority by the name within brackets.
For the chickens of white Silk-fowls, see
Tegetmeier's' Poultry Book,'1566, p. 221.

302
CHAP. VII.
FOWLS.
where. Spanish chickens also, occasionally (Brent), have, where
the down was white, their first true feathers tipped for a time with
white. The primordially striped character is retained by the
chickens of most of the Game sub-breeds (Brent, Dixon); by Dork-
ings; by the partridge and grouse-coloured sub-breeds of Cochins
(Brent), but not, as we have seen, by all the other sub-breeds; by
the pheasant-Malay (Dixon), but apparently not (at which I am
much surprised) by other Malays. The following breeds and sub-
breeds are barely, or not at all, longitudinally striped; viz. gold and
silver pencilled Hamburghs, which can hardly be distinguished
from each other (Brent) in the down, both having a few dark spots
on the head and rump, with occasionally a longitudinal stripe
(Dixon) on the back of the neck. I have seen only one chicken of
the silver-spangled Hamburgh, and this was obscurely striped along
the back. Gold-spangled Polish chickens (Tegetmeier) are of a
warm russet brown; and silver-spangled Polish chickens are grey,
sometimes (Dixon) with dashes of ochre on the head, wings, and
breast. Cuckoo and blue-dun fowls (Dixon) are grey in the down.
The chickens of Sebright Bantams (Dixon) are uniformly dark
brown, whilst those of the brown-breasted red Game Bantam are
black, with some white on the throat and breast. From these facts
we see that the chickens of the different breeds, and even of the
same main breed, differ much in their downy plumage; and,
although longitudinal stripes characterise the young of all wild
gallinaceous birds, they disappear in several domestic breeds.
Perhaps it may be accepted as a general rule that the more the
adult plumage differs from that of the adult G. bankiva, the more
completely the chickens have lost their proper stripes.
With respect to the period of life at which the charac-
ters proper to each breed first appear, it is obvious that
such structures as additional toes must be formed long
before birth. In Polish fowls, the extraordinary protu-
berance of the anterior part of the skull is well developed
before the chickens come out of the egg;40 but the crest,
which is supported on the protuberance, is at first feebly
developed, nor does it attain its full size until the second
year. The Spanish cock is pre-eminent for his magnifi-
cent comb, and this is developed at an unusually early
40 As I hear from Mr. Tegetmeier ; see
also Proc. Zoolog. Soc. 1856, p. 366.
On the late development of the crest, seo
"Poultry Chronicle,' vol. ii. p. 132.

CHAP. VII. DIFFERENCES BETWEEN THE BREEDS. 303
age; so that the young males can be distinguished from
the females when only a few weeks old, and therefore
earlier than in other breeds; they likewise crow very
early, namely, when about six weeks old. In the Dutch
sub-breed of the Spanish fowl the white ear-lappets are
developed earlier than in the common Spanish breed.41
Cochins are characterised by a small tail, and in the
young cocks the tail is developed at an unusually late
period. Game fowls are notorious for their pugnacity;
and the young cocks crow, clap their little wings, and
obstinately fight with each other, even whilst under their
mother's care.43 “I have often had,” says one author, 44
“whole broods, scarcely feathered, stone-blind from fight-
ing; the rival couples moping in corners, and renewing
their battles on obtaining the first ray of light.” With
the males of all gallinaceous birds the use of their
weapons and pugnacity is to fight for the possession of
the females; so that the tendency in our Game chickens
to fight at an extremely early age is not only useless, but
is injurious, as they suffer so much from their wounds.
The training for battle during an early period may be
natural to the wild Gallus bankiva ; but as man during
many generations has gone on selecting the most obsti-
nately pugnacious cocks, it is more probable that their
pugnacity has been unnaturally increased, and unnatu-
rally transferred to the young male chickens. In the
same manner, it is probable that the extraordinary de-
velopment of the comb in the Spanish cock has been
unintentionally transferred to the young cocks; for
fanciers would not care whether their young birds had
large combs, but would select for breeding the adults
which had the finest combs, whether or not developed at
43 Ferguson on Rare and Prize Poultry,
p. 261.
41 On these points, see Poultry Chro-
nicle,' vol. iii. p. 166; and Tegetmeier's
"Poultry Book,' 1866, pp. 105 and 121.
42 Dixon, Ornamental and Domestic
Poultry,' p. 273.
44 Mowbray on Poultry, 7th edit. 1834,
p. 13.

304
CHAP. VII.
FOWLS.
an early period. The last point which need here be
noticed is that, though the chickens of Spanish and Malay
fowls are well covered with down, the true feathers are
acquired at an unusually late age; so that for a time the
young birds are partially naked, and are liable to suffer
from cold.
Secondary Sexual Characters.--The two sexes in the
parent-form, the Gallus bankiva, differ much in colour.
In our domestic breeds the difference is never greater, but
is often less, and varies much in degree even in the sub-
breeds of the same main breed. Thus in certain Game
fowls the difference is as great as in the parent-form,
whilst in the black and white sub-breeds there is no dif-
ference in plumage. Mr. Brent informs me that he has
seen two strains of black-breasted red Games, in which
the cocks could not be distinguished, whilst the hens in
one were partridge-brown and in the other fawn-brown.
A similar case has been observed in the strains of the
brown-breasted red Game. The hen of the 66 duck-
winged Game" is “extremely beautiful," and differs
much from the hens of all the other Game sub-breeds;
but generally, as with the blue and
grey
Game and with
some sub-varieties of the pile-game, a moderately close
relation may be observed between the males and females
in the variation of their plumage. A similar relation is
also evident when we compare the several varieties of
Cochins. In the two sexes of gold and silver-spangled
and of buff Polish fowls, there is much general similarity
in the colouring and marks of the whole plumage, ex-
cepting of course in the hackles, crest, and beard. In
spangled Hamburghs, there is likewise a considerable de-
gree of similarity between the two sexes. In pencilled
Hamburghs, on the other hand, there is much dissimilar-
ity; the pencilling which is characteristic of the hens be-
45 See the full description of the varie-
ties of the Game-breed, in Tegetmeier's
Poultry Book,' 1866, p. 131. For Cuc-
koo Dorkings, p. 97.

CHAP. VII.
305
SEXUAL DIFFERENCES.
ing almost absent in the males of both the golden and
silver varieties. But, as we have already seen, it cannot
be given as a general rule that male fowls never have
pencilled feathers, for Cuckoo Dorkings are “remarkable
from having nearly similar markings in both sexes."
It is a singular fact that the males in certain sub-breeds
have lost some of their secondary masculine characters,
and, from their close resemblance in plumage to the fe-
males, are often called hennies. There is much diversity
of opinion whether these males are in any degree sterile;
that they sometimes are partially sterile seems clear, 46 but
this may have been caused by too close interbreeding.
That they are not quite sterile, and that the whole case
is widely different from that of old females assuming
masculine characters, is evident from several of these hen-
like sub-breeds having been long propagated. The males
and females of gold and silver-laced Sebright Bantams
can be barely distinguished from each other, except by
their combs, wattles, and spurs, for they are coloured
alike, and the males have not hackles, nor the flowiug
sickle-like tail-feathers. A hen-tailed sub-breed of Ham-
burghs was recently much esteemed. There is also a
breed of Game-fowls, in which the males and females re-
semble each other so closely that the cocks have often
mistaken their hen-feathered opponents in the cock-pit
for real hens, and by the mistake have lost their lives. 47
The cocks, though dressed in the feathers of the hen,
“ are high-spirited birds, and their courage has been often
proved :” an engraving even has been published of one
celebrated hen-tailed victor. Mr. Tegetmeier
Mr. Tegetmeier 18 has re-
corded the remarkable case of a brown-breasted red
Game-cock which, after assuming its perfect masculine
46 Mr. Hewitt in Tegetmeier's 'Poultry
Bouk,' 186, pp. 246 and 156. For hen-
tailed game-cocks, see p. 131.
47 The Field,' April 20th, 1861. The
writer says he has seen half-a-dozen
cocks thus sacrificed.
48 Proceedings of Zoolog. Soc.' March,
1861, p. 102. The engraving of the hen-
tailed cock just alluded to was exhibited
at the Society.

306
CHAP. VII.
FOWLS.
plumage, became hen-feathered in the autumn of the fol-
lowing year; but he did not lose voice, spurs, strength,
nor productiveness. This bird has now retained the same
character during five seasons, and has begot both hen-
'feathered and male-feathered offspring. Mr. Grantley F.
Berkeley relates the still more singular case of a cele-
brated strain of “polecat Game-fowls," which produced
in nearly every brood a single hen-cock.
“ The great
peculiarity in one of these birds was that he, as the sea-
sons succeeded each other, was not always a hen-cock,
and not always of the colour called the polecat, which is
black. From the polecat and hen-cock feather in one
season he moulted to a full male-plumaged black-breasted
red, and in the following year he returned to the former
feather." 49
I have remarked in my Origin of Species' that sec-
ondary sexual characters are apt to differ much in the
species of the same genus, and to be unusually variable
in the individuals of the species. So it is with the breeds
of the fowl, as we have already seen, as far as the colour
of plumage is concerned, and so it is with the other
secondary sexual characters. Firstly, the comb differs
much in the various breeds, and its form is eminently
characteristic of each kind, with the exception of the
Dorkings, in which the form has not been as yet deter-
mined on by fanciers, and fixed by selection. A single,
deeply-serrated comb is the typical and most common
form. It differs much in size, being immensely develop-
ed in Spanish fowls; and in a local breed called Red-
caps, it is sometimes "upwards of threc inches in breadth
at the front, and more than four inches in length, meas-
ured to the end of the peak behind.” 51 In some breeds
the comb is double, and when the two ends are cemented
49 The Field,' April 20th, 1861.
50 I am much indebted to Mr. Brent
for an account, with sketches, of all the
variations of the comb known to him,
and likewise with respect to the tail, as
presently to be given.
51 The Poultry Book,' by Tegetmeier,
1866, p. 231.

CHAP. VII.
307
SEXUAL DIFFERENCES.
together it forms a “cup-comb;' in the “rose-comb” it
is depressed, covered with small projections, and pro-
duced backwards; in the horned and crève-cœur fowl it
is produced into two horns; it is triple in the pea-combed
Brahmas, short and truncated in the Malays, and absent
in the Guelderlands. In the tasselled Game a few long
feathers arise from the back of the comb; in
many
breeds
a crest of feathers replaces the comb. The crest, when
little developed, arises from a fleshy mass, but, when
much developed, from a hemispherical protuberance of
the skull. In the best Polish fowls it is so largely de-
veloped, that I have seen birds which could hardly pick
up their food; and a German writer asserts that they
are in consequence liable to be struck by hawks. Mon-
strous structures of this kind would thus be suppressed
in a state of nature. The wattles, also, vary much in
size, being small in Malays and some other breeds; they
are replaced in certain Polish sub-breeds by a great tuft
of feathers called a beard.
The hackles do not differ much in the various breeds,
but are short and stiff in Malays, and absent in Hennies.
As in some orders of birds the males display extraordi-
narily-shaped feathers, such as naked shafts with discs at
the end, &c., the following case may be worth giving.
In the wild Gallus bankiva and in our domestic fowls,
the barbs which arise from each side of the extremities
of the hackles are naked or not clothed with barbules, so
that they resemble bristles; but Mr. Brent sent me some
scapular hackles from a young Birchen Duckwing Game
cock, in which the naked barbs became densely reclothed
with barbules towards their tips; so that these tips,
which were dark coloured with a metallic lustre, were
separated from the lower parts by a symmetrically-shaped
transparent zone formed of the naked portions of the
62 Die Hühner und Pfauenzucht, 1827, s. 11.

308
CHAP. VII.
FOWLS.
barbs. Hence the coloured tips appeared like little sepa-
rate metallic discs.
The sickle-feathers in the tail, of which there are three
pair, and which are eminently characteristic of the male
sex, differ much in the various breeds. They are scimitar-
shaped in some Hamburghs, instead of being long and flow-
ing as in the typical breeds. They are extremely short in
Cochins, and are not at all developed in Hennies. They
are carried, together with the whole tail, erect in Dork-
ings and Games; but droop much in Malays and in some
Cochins. Sultans are characterized by an additional num.
ber of lateral sickle-feathers. The spurs vary much, being
placed higher or lower on the shank; being extremely
long and sharp in Games, and blunt and short in Cochins.
These latter birds seem aware that their spurs are not
efficient weapons; for though they occasionally use them,
they more frequently fight, as I am informed by Mr.
Tegetmeier, by seizing and shaking each other with their
beaks. In some Indian Game-cocks, received by Mr.
Brent from Germany, there are, as he informs me, three,
four, or even five spurs on each leg. Some Dorkings also
have two spurs on each leg; 5% and in birds of this breed
the spur is often placed almost on the outside of the leg.
Double spurs are mentioned in the ancient Chinese Ency-
clopædia. Their occurrence may be considered as a case
of analogous variation, for some wild gallinaceous birds,
for instance, the Polyplectron, have double spurs.
Judging from the differences which generally distin-
guish the sexes in the Gallinaceæ, certain characters in
our domestic fowls appear to have been transferred from
the one sex to the other. In all the species (except in
Turnix), when there is any conspicuous difference in plu-
mage between the male and female, the male is always
53 Poultry Chronicle,' vol. i. p. 595.
Mr. Brent has informed me of the same
fact. With respect to the position of
the spurs in Dorkings, see Cottage
Gardener,' Sept. 18th, 1860, p. 350.

CHAP. VII,
309
SEXUAL DIFFERENCES.
the most beautiful; but in golden-spangled Hamburghs
the hen is equally beautiful with the cock, and incompa-
rably more beautiful than the hen in any natural species
of Gallus; so that here a masculine character has been
transferred to the female. On the other hand, in cuckoo
Dorkings and in other cuckoo breeds the pencilling, which
in Gallus is a female attribute, has been transferred to
the male: nor, on the principle of analogous variation, is
this transference surprising, as the males in many galli-
naceous genera are barred or pencilled. With most of
these birds head ornaments of all kinds are more fully
developed in the male than in the female ; but in Polish
fowls the crest or top-knot, which in the male replaces
the comb, is equally developed in both sexes. In certain
sub-breeds, which from the hen having a small crest, are
called lark-crested," a single upright comb sometimes al-
most entirely takes the place of the crest in the male.” 54
From this latter case, and from some facts presently to
be given with respect to the protuberance of the skull in
Polish fowls, the crest in this breed ought perhaps to be
viewed as a feminine character which has been transferred
to the male. In the Spanish breed the male, as we know,
has an immense comb, and this has been partially transfer-
red to the female, for her comb is unusually large, though
not upright. In Game-fowls the bold and savage disposi-
tion of the male has likewise been largely transferred to the
female; 65 and she sometimes even possesses the eminently
masculine character of spurs. Many cases are on record
of hens being furnished with spurs; and in Germany, ac-
cording to Bechstein, the spurs in the Silk-hen are some-
times very long. He mentions also another breed simi-
larly characterized, in which the hens are excellent.layers,
64 Dixon, Ornamental and Domestic
Poultry,' p. 320.
65 Mr. Tegetmeier informs me that
Game hens have been found so com-
bative, that it is now generally the prac-
tice to exhibit each ben in a separate
pen.
56 "Naturgeschichte Deutschlands,
Band iii. (1793), s. 339, 407.

310
CHAP. VII.
FOWLS.
but are apt to disturb and break their eggs owing to their
spurs.
Mr. Layard 67 has given an account of a breed of fowls
in Ceylon with black skin, bones, and wattle, but with
ordinary feathers, and which cannot be more aptly de-
scribed than by comparing them to a white fowl drawn
down a sooty chimney; it is, however," adds Mr. Layard,
"a remarkable fact that a male bird of the pure sooty
variety is almost as rare as a tortoise-shell tom-cat.” Mr.
Blyth finds that the same rule holds good with this breed
near Calcutta. The males and females, on the other hand,
of the black-boned European breed, with silky feathers,
do not differ from each other; so that in the one breed
black skin and bones, and the same kind of plumage, are
common to both sexes, whilst in the other breed these
characters are confined to the female sex.
At the present day all the breeds of Polish fowls have
the great bony protuberance on their skulls, which in-
cludes part of the brain and supports the crest, equally
developed in both sexes. But formerly in Germany the
skull of the hen alone was protuberant: Blumenbach,
who particularly attended to abnormal peculiarities in
domestic animals, states, in 1813, that this was the case;
and Bechstein had previously, in 1793, observed the same
fact. This latter author has carefully described the effects
of a crest on the skull not only in fowls, but in ducks,
geese, and canaries. He states that with fowls, when the
crest is not much developed, it is supported on a fatty
mass; but when much developed, it is always supported
57 On the Ornithology of Ceylon in
Annals and Mag. of Nat. History,' 2nd
series, vol. xiv. (1854), p. 63.
58 I quote Blumenbach on the autho-
rity of Mr. Tegetmeier, who gives in
Proc. Zoolog. Soc.,' Nov. 25th, 1856, a
very interesting account of the skets
of Polish fowls. Mr. Tegetmeier,
knowing of Bechstein's account, dis-
puted the accuracy of Blumenbach's
statement. For Bechstein, see Natur-
geschichte Deutschlands, Band ii.
(1793), s. 399, note. I may add that at
the first exhibition of poultry at the
Zoological Gardens, in May, 1845, I saw
some fowls, called Friezland fowls, of
which the hens were crested, and the
cocks were furnished with a comb.

CHAP. VII.
311
EXTERNAL DIFFERENCES.
on a bony protuberance of variable size.
He well de-
scribes the peculiarities of this protuberance, and he at-
tended to the effects of the modified shape of the brain on
the intellect of these birds, and disputes Pallas' statement
that they are stupid. He then expressly states that he
never observed this protuberance in male fowls. Hence
there can be no doubt that this remarkable character in
the skulls of Polish fowls was formerly in Germany con-
fined to the female sex, but has now been transferred to
the males, and has thus become common to both sexes.
External Differences, not connected with the sexes, between
the breeds and between individual birds.
The size of the body differs greatly. Mr. Tegetmeier has known
a Brahma to weigh 17 pounds; a fine Malay cock 10 pounds; whilst
a first-rate Sebright Bantam weighs hardly more than 1 pound.
During the last 20 years the size of some of our breeds has been
largely increased by methodical selection, whilst that of other breeds
has been much diminished. We have already seen how greatly
colour varies even within the same breed; we know that the wild
G. bankiva varies slightly in colour ; we know that colour is varia-
ble in all our domestic animals; nevertheless some eminent fanciers
have so little faith in variability, that they have actually argued
that the chief Game sub-breeds, which differ from each other in
nothing but colour, are descended from distinct wild species! Cross-
ing often causes strange modifications of colour. Mr. Tegetmeier
informs me that when buff and white Cochins are crossed, some of
the chickens are almost invariably black. According to Mr. Brent,
black and white Cochins occasionally produce chickens of a slaty-
blue tint; and this same tint appears, as Mr. Tegetmeier tells me,
from crossing white Cochins with black Spanish fowls, or white
Dorkings with black Minorcas.59 A good observer 60 states that a
first-rate silver-spangled Hamburgh hen gradually lost the mast
characteristic qualities of the breed, for the black lacing to her feath-
ers disappeared, and her legs changed from leaden-blue to white;
but what makes the case remarkable is, that this tendency ran in
the blood, for her sister changed in a similar but less strongly
59 Cottage Gardener,' Jan. 3rd, 1860,
fore the Dublin Nat. Hist. Soc., quoted in
Cottage Gardener,' 1856, p. 161.
p. 218.
60 Mr. Williams, in a paper read be-

312
CHAP. VII.
FOWLS.
marked manner; and chickens produced from this latter hen were
at first almost pure white, “but on moulting acquired black collars
and some spangled feathers with almost obliterated markings;" so
that a new variety arose in this singular manner. The skin in the
different breeds differs much in colour, being white in common kinds,
yellow in Malays and Cochins, and black in Silk fowls; thus mock-
ing, as M. Godron 61 remarks, the three principal types of skin in
mankind. The same author adds, that, as different kinds of fowls
living in distant and isolated parts of the world have black skin and
bones, this colour must have appeared at various times and places.
The shape and carriage of the body and the shape of the head dif
fer much. The beak varies slightly in length and curvature, but in
comparably less than with pigeons. In most crested fowls the nos-
trils offer a remarkable peculiarity in being raised with a crescentic
outline. The primary wing-feathers are short in Cochins; in a
male, which must have been more than twice as heavy as G. bankiva,
these feathers were in both birds of the same length. I have count:
ed, with Mr. Tegetmeier's aid, the primary wing-feathers in thirteen
cocks and hens of various breeds; in four of them, namely in two
Hamburghs, a Cochin, and Game Bantam, there were 10, instead of
the normal number 9; but in counting these feathers I have followed
the practice of fanciers, and have not included the first minute pri
mary feather, barely three-quarters of an inch in length. These
feathers differ considerably in relative length, the fourth, or the fifth,
or the sixth, being the longest ; with the third either equal to, or
considerably shorter than the fifth. In wild gallinaceous species the
relative length and number of the main wing and tail-feathers are
extremely constant.
The tail differs much in erectness and size, being small in Malays
and very small in Cochins. In thirteen fowls of various breeds
which I have examined, five had the normal number of 14 feath.
ers, including in this number the two middle sickle-feathers; six
others (viz. a Caffre cock, Gold-spangled Polish cock, Cochin hen,
Sultan hen, Game hen, and Malay hen) had 16; and two (an old
Cochin cock and Malay hen) had 17 feathers. The rumpless fowl
has no tail, and in a bird which I kept alive the oil-gland had abort-
ed; but this bird, though the.os coccygis was extremely imperfect,
had a vestige of a tail with two rather long feathers in the position
of the outer caudals. This bird came from a family where, as I was
81 De l'Espèce,' 1859, 442. For the
occurrence of black-boned fowls in South
America, see Roulin, in Mém. de
l'Acad. des Sciences,' tom. vi. p. 351; and
Azara, 'Quadrupèdes du Paraguay,' tom.
ii. p. 324. A frizzled fowl sent to me
from Madras had black bones.

CHAP. VII.
313
EXTERNAL DIFFERENCES.
told, the breed had kept true for twenty years; but rumpless fowls
often produce chickens with tails.62 An eminent physiologist has
recently spoken of this breed as a distinct species; had he examined
the deformed state of the os coccyx he would never have come to
this conclusion; he was probably misled by the statement, which
may be found in some works, that tailless fowls are wild in Ceylon;
but this statement, as I have been assured by Mr. Layard and Dr.
Kellaert, who have so closely studied the birds of Ceylon, is utterly
false.
The tarsi vary considerably in length, being relatively to the
femur considerably longer in the Spanish and Frizzled, and shorter
in the Silk and Bantam breeds, than in the wild G. bankiva; but
in the latter, as we have seen, the tarsi vary in length. The tarsi
are often feathered. The feet in many breeds are furnished with
additional toes. Golden-spangled Polish fowls are said 64 to have
the skin between their toes much developed ; Mr. Tegetmeier
observed this in one bird, but it was not so in one which I exam-
ined. In Cochins the middle toe is said 65 to be nearly double the
length of the lateral toes, and therefore much longer than in G.
bankiva or in other fowls; but this was not the case in two which I
examined. The nail of the middle toe in this same breed is sur-
prisingly broad and flat, but in a variable degree in two birds
which I examined; of this structure in the nail there is only a
trace in G. bankiva.
The voice differs slightly, as I am informed by Mr. Dixon, in
almost every breed. The Malays 66 have a loud, deep, somewhat
prolonged crow, but with considerable individual differences. Col-
onel Sykes remarks that the domestic Kulm cock in India has not
the shrill clear pipe of the English bird, and "his scale of notes
appears more limited.” Dr. Hooker was struck with the "pro-
longed howling screech" of the cocks in Sikhim.67 The crow of the
Cochin is notoriously and ludicrously different from that of the
common cock. The disposition of the different breeds is widely
different, varying from the savage and defiant temper of the Game-
cock to the extremely peaceable temper of the Cochin. The latter,
it has been asserted, "graze to a much greater extent than any
62 Mr. Hewitt, in Tegetmeier's Poul-
try Book,' 1866, p. 231.
63 Dr. Broca, in Brown-Sequard's
Journal de Phys.,' tom. ii. p. 361.
64 Dixon's Ornamental Poultry,' p.
325.
65 Poultry Chronicle,' vol. i. p. 485.
Tegetmeier's Poultry Book, 1966, p.
41. On Cochins grazing, idem, p. 46.
66 Ferguson on 'Prize Poultry,' p. 187.
67 Col. Sykes in Proc. Zoolog. Soc.,'
1832, p. 151. Dr. Hooker's Himalayan
Journals,' vol. i. p. 314.
14

314
CHAP. VII,
FOWLS.
other varieties.” The Spanish fowls suffer more from frost than
other breeds.
Before we pass on to the skeleton, the degree of dis-
tinctness of the several breeds from G. bankiva ought to
be noticed. Some writers speak of the Spanish as one
of the most distinct breeds, and so it is in general aspect;
but its characteristic differences are not important. The
Malay appears to me more distinct, from its tall stature,
small drooping tail with more than fourteen tail-feathers,
and from its small comb and wattles; nevertheless one
Malay sub-breed is coloured almost exactly like G. ban-
kiva. Some authors consider the Polish fowl as very
distinct; but this is a semi-monstrous breed, as shown
by the protuberant and irregularly perforated skull.
The Cochin, with its deeply furrowed frontal bones,
peculiarly shaped occipital foramen, short wing-feathers,
short tail containing more than fourteen feathers, broad
nail to the middle toe, fluffy plumage, rough and dark-
coloured eggs, and especially from its peculiar voice, is
probably the most distinct of all the breeds. If any one
of our breeds has descended from some unknown species,
distinct from G. bankiva, it is probably the Cochin;
but the balance of evidence does not favour this view.
All the characteristic differences of the Cochin breed are
more or less variable, and may be detected in a greater
or lesser degree in other breeds. One sub-breed is col-
oured closely like G. bankiva. The feathered legs, often
furnished with an additional toe, the wings incapable of
flight, the extremely quiet disposition, indicate a long
course of domestication; and these fowls come from
China, where we know that plants and animals have
been tended from a remote period with extraordinary
care, and where consequently we might expect to find
profoundly modified domestic races.
Osteological Differences. I have examined twenty-
seven skeletons and fifty-three skulls of various breeds,
including three of G. bankiva: nearly half of these

CHAP. VII.
315
OSTEOLOGICAL DIFFERENCES.
skulls I owe to the kindness of Mr. Tegetmeier, and three
of the skeletons to Mr. Eyton.
The Skull differs greatly in size in different breeds, being nearly
twice as long in the largest Cochins, but not nearly twice as broad,
as in Bantams. The bones at the base, from the occipital foramen
to the anterior end (including the quadrates and pterygoids), are
absolutely identical in shape in all the skulls. So is the lower jaw.
In the forehead slight differences are often perceptible between the
males and females, evidently caused by the presence of the comb.
In every case I take the skull of G. banjoiva as the standard of com-
parison. In four Games, in one Malay hen, in an African cock,
in a Frizzled cock from Madras, in two black-boned Silk hens,
no differences occur worth notice. In three Spanish cocks, the
form of the forehead between the orbits differs considerably; in
one it is considerably depressed, whilst in the two others it is rather
prominent, with a deep medial furrow; the skull of the hen is
smooth. In three skulls of Sebright Bantams the crown is more
globular, and slopes more abruptly to the occiput, than in G. ban-
kiva. In a Bantam or Jumper from Burmah these same characters
are more strongly pronounced, and the supra-occiput is more point-
ed. In a black Bantam the skull is not so globular, and the occipital
foramen is very large, and has nearly the same sub-triangular out-
line presently to be described in Cochins; and in this skull the two
ascending branches of the premaxillary are overlapped in a singu-
lar manner by the processes of the nasal bone, but, as I have seen
only one specimen, some of these differences may be individual.
Of Cochins and Brahmas (the latter a crossed race approaching
closely to Cochins) I have examined seven skulls; at the point
where the ascending branches of the premaxillary rest on the
frontal bone the surface is much depressed, and from this depression
a deep medial furrow extends backwards to a variable distance;
the edges of this fissure are rather prominent, as is the top of the
skull behind and over the orbits. These characters are less de-
veloped in the hens. The pterygoids, and the processes of the
lower jaw, relatively to the size of the head, are broader than in
G. bankiva, and this is likewise the case with Dorkings when of
large size. The terminal fork of the hyoid bone in Cochins is
twice as wide as in G. bankiva, whereas the length of the other
hyoid bones is only as three to two. But the most remarkable
character is the shape of the occipital foramen: in G. bankiva (A)
the breadth in a horizontal line exceeds the height in a vertical
line, and the outline is nearly circular; whereas in Cochins (B) the
outline is sub-triangular, and the vertical line exceeds the hori-

316
CHAP. VII.
FOWLS.
zontal line in length. This same form likewise occurs in the black
Bantam above referred to, and an approach to it may be seen in
some Dorkings, and in a slight degree in certain other breeds.
B
A
Fig. 33.-Occipital Foramen, of natural size. A. Wild Gallus bankiva. B. Cochin
Cock.
Of Dorkings I have examined three skulls, one belonging to the
white sub-breed; the one character deserving notice is the breadth
of the frontal bones, which are moderately furrowed in the middle;
thus in a skull which was less than once and a half the length of
B
CUS
A
Fig. 34.-Skulls of natural size, viewed from above, a little obliquely. A. Wild
Gallus bankiva. B. White-crested Polish Cock.

CHAP. VII.
317
OSTEOLOGICAL DIFFERENCES.
that of G. bankiva, the breadth between the orbits was exactly
double. Of Hamburghs I have examined four skulls (male and fe-
male) of the pencilled sub-breed, and one (male) of the spangled
sub-breed ; the nasal bones stand remarkably wide apart, but in a
variable degree; consequently narrow membrane-covered spaces are
left between the tips of the two ascending branches of the premaxil-
lary bones, which are rather short, and between these branches and
the nasal bones. The surface of the frontal bone, on which the
branches of the premaxillary rest, is very little depressed. These
peculiarities no doubt stand in close relation with the broad flat-
tened rose-comb characteristic of the Hamburgh breed.
I have examined fourteen skulls of Polish and other crested breeds.
Their differences are extraordinary. First for nine skulls of differ-
ent sub-breeds of English Polish fowls. The hemispherical protu-
berance of the frontal bones 68 may be seen in the accompanying
drawings, in which (B) the skull of a white-crested Polish fowl is
shown obliquely from above, with the skull (A) of G. bankiva in
the same position. In fig. 35 longitudinal sections are given of the
skulls of a Polish fowl, and, for comparison, of a Cochin of the same
size. The protuberance in all Polish fowls occupies the same posi-
tion, but differs much in size. In one of my nine specimens it was
extremely slight. The degree to which the protuberance is ossified
varies greatly, larger or smaller portions of bone being replaced by
membrane. In one specimen there was only a single open pore;
generally, there are many variously-shaped open spaces, the bone
forming an irregular reticulation. A medial, longitudinal, arched
ribbon of bone is generally retained, but in one specimen there was
no bone whatever over the whole protuberance, and the skull when
cleaned and viewed from above presented the appearance of an open
basin. The change in the whole internal form of the skull is sur-
prisingly great. The brain is modified in a corresponding manner,
as is shown in the two longitudinal sections, which deserve atten-
tive consideration. The upper and anterior cavity of the three into
which the skull may be divided, is the one which is so greatly
modified; it is evidently much larger than in the Cochin skull of
the same size, and extends much further beyond the interorbital
septum, but laterally is less deep. Whether this cavity is entirely
filled by the brain, may be doubted. In the skull of the Cochin
68 See Mr. Tegetmeier's account, with
woodcuts, of the skull of Polish fowls, in
Proc. Zoolog. Soc.,' Nov. 25th, 1856. For
other references, see Isid. Geoffroy Saint
Hilaire, Hist. Gén. des Anomalies,' tom.
i. p. 237. M. C. Dareste suspects (Re-
cherches sur les Conditions de la Vie,
&c., Lille, 1863, p. 36) that the protube-
rance is not formed by the frontal bones,
but by the ossification of the dura mater.

318
CHAP. VII.
FOWLS.
and of all ordinary fowls a strong internal ridge of bone separates
the anterior from the central cavity; but this ridge is entirely ab-
sent in the Polish skull here figured. The shape of the central
cavity is circular in the Polish, and lengthened in the Cochin skull.
А
B
Fig. 35.–Longitudinal sections of Skull, of natural size, viewed laterally. A.
Polish Cock. B. Cochin Cock, selected for comparison with the above
from being of nearly the same size.
The shape of the posterior cavity, together with the position, size,
and number of the pores for the nerves, differ much in these two
skulls. A pit deeply penetrating the occipital bone of the Cochin
is entirely absent in this Polish skull, whilst in another specimen
it was well developed. In this second specimen the whole internal
surface of the posterior cavity likewise differs to a certain extent in
shape. I made sections of two other skulls,-namely, of a Polish
fowl with the protuberance singularly little developed, and of a
Sultan in which it was a little more developed ; and when these
two skulls were placed between the two above figured (fig. 35), a
perfect gradation in the configuration of each part of the internal
surface could be traced. In the Polish skull, with a small protu-
berance, the ridge between the anterior and middle cavities was

CHAP. VII.
319
OSTEOLOGICAL DIFFERENCES.
present, but low; and in the Sultan this ridge was replaced by a
narrow furrow standing on a broad raised eminence.
It may naturally be asked whether these remarkable modifica-
tions in the form of the brain affect the intellect of Polish fowls;
some writers have stated that they are extremely stupid, but Bech-
stein and Mr. Tegetmeier have shown that this is by no means
generally the case. Nevertheless Bechstein 69 states that he had a
Polish hen which "was crazy, and anxiously wandered about all
day long." A hen in my possession was solitary in her habits, and
was often so absorbed in reverie that she could be touched ; she
was also deficient in the most singular manner in the faculty of
finding her way, so that, if she strayed a hundred yards from her
feeding-place, she was completely lost, and would then obstinately
try to proceed in a wrong direction. I have received other and
similar accounts of Polish fowls appearing stupid or half-idiotic.ro
To return to the skull. The posterior part, viewed externally,
differs little from that of G. bankiva. In most fowls the posterior-
lateral process of the frontal bone and the process of the squamosal
bone run together and are ossified near their extremities: this
union of the two bones, however, is not constant in any breed ;
and in eleven out of fourteen skulls of crested breeds, these pro-
cesses were quite distinct. These processes, when not united,
instead of being inclined anteriorly as in all common breeds,
descend at right angles to the lower jaw; and in this case the
longer axis of the bony cavity of the ear is likewise more perpen-
dicular than in other breeds. When the squamosal process is free,
instead of expanding at the tip, it is reduced to an extremely fine
and pointed style, of variable length. The pterygoid and quadrate
bones present no difference. The palatine bones are a little more
curved upwards at their posterior ends. The frontal bones, an-
teriorly to the protuberance, are, as in Dorkings, very broad, but in
a variable degree. The nasal bones either stand far apart, as in
Hamburghs, or almost touch each other, and in one instance were
ossified together. Each nasal bone properly sends out in front two
long processes of equal lengths, forming a fork; but in all the
Polish skulls, except one, the inner process was considerably, but
in a variable degree, shortened and somewhat upturned. In all the
skulls, except one, the two ascending branches of the premaxillary,
instead of running up between the processes of the nasal bones
and resting on the ethmoid bone, are much shortened and terminate
69 Naturgeschichte Deutschlands, have received communications to a si-
Band iii. (1793), s. 400.
milar effect from Messrs. Brent and
70 The Field,' May 11th, 1861. I Tegetmeier.

320
CHAP. VII.
FOWLS.
in a blunt, somewhat upturned point. In those skulls in which
the nasal bones approach quite close to each other or are ossified
together, it would be impossible for the ascending branches of the
premaxillary to reach the ethmoid and frontal bones; hence we see
that even the relative connection of the bones has been changed.
Apparently in consequence of the branches of the premaxillary and
of the inner processes of the nasal bones being somewhat upturned,
the external orifices of the nostrils are upraised and assume a cre-
scentic outline.
Fig. 36.-Skull of Horned Fowl, of natural size, viewed from above, a little
obliquely. (In the possession of Mr. Tegetmeier.)
I must still say a few words on some of the foreign Crested
breeds. The skull of a crested, rumpless, white Turkish fowl is
very slightly protuberant, and but little perforated; the ascending
branches of the premaxillary are well developed. In another Tur-
kish breed, called Ghoondooks, the skull is considerably protuberant
and perforated; the ascending branches of the premaxillary are so
much aborted that they project only isth of an inch; and the inner
processes of the nasal bone are so completely aborted, that the sur-
face where they should have projected is quite smooth. Here then
we see these two bones modified to an extreme degree. Of Sultans
(another Turkish breed) I examined two skulls; in that of the
female the protuberance was much larger than in the male. In
both skulls the ascending branches of the premaxillary were very
short, and in both the basal portion of the inner processes of the
nasal bones were ossified together. These Sultan skulls differed
from those of English Polish fowls in the frontal bones, anteriorly
to the protuberance, not being broad.
The last skull which I need describe is a unique one, lent to me

CHAP. VII.
321
OSTEOLOGICAL DIFFERENCES.
A series may
by Mr. Tegetmeier: it resembles a Polish skull in most of its cha-
racters, but has not the great frontal protuberance; it has, however,
two rounded knobs of a different nature, which stand more in front,
above the lachrymal bones. These curious knobs, into which the
brain does not enter, are separated from each other by a deep me-
dial furrow; and this is perforated by a few minute pores. The
nasal bones stand rather wide apart, with their inner processes, and
the ascending branches of the premaxillary, upturned and shortened.
The two knobs no doubt supported the two great horn-like pro-
jections of the comb.
From the foregoing facts we see in how astonishing a manner
some of the bones of the skull vary in Crested fowls. The protube-
rance may certainly be called in one sense a monstrosity, as being
wholly unlike anything observed in nature : but as in ordinary
cases it is not injurious to the bird, and as it is strictly inherited, it
can hardly in another sense be called a monstrosity.
be formed commencing with the black-boned Silk fowl, which has
a very small crest with the skull beneath penetrated only by a few
minute orifices, but with no other change in its structure, and from
this first stage we may proceed to fowls with a moderately large
crest, which rests, according to Bechstein, on a fleshy mass, but
without any protuberance in the skull. I may add that I have seen
a similar fleshy or fibrous mass beneath the tuft of feathers on the
head of the Tufted duck; and in this case there was no actual pro-
tuberance in the skull, but it had become a little more globular.
Lastly, when we come to fowls with a largely developed crest, the
skull becomes largely protuberant and is perforated by a multitude
of irregular open spaces.
The close relation between the crest and
the size of the bony protuberance is shown in another way; for
Mr. Tegetmeier informs me that if chickens lately hatched be select-
ed with a large bony protuberance, when adult they will have a
large crest. There can be no doubt that in former times the breed-
er of Polish fowls attended solely to the crest, and not to the skull;
nevertheless, by increasing the crest, in which he has wonderfully
succeeded, he has unintentionally made the skull protuberant to an
astonishing degree; and through correlation of growth, he has at
the same time affected the form and relative connexion of the pre-
maxillary and nasal bones, the shape of the orifice of the nose, the
breadth of the frontal bones, the shape of the post-lateral processes
of the frontal and squamosal bones, the direction of the axis of the
bony cavity of the ear, and lastly the internal configuration of the
whole skull together with the shape of the brain.
Vertebræ.-In G. bankiva there are fourteen cervical, seven dorsal
with ribs, apparently fifteen lumbar and sacral, and six caudal ver-
14*

322
CHAP. VII.
FOWLS.
tebræ ; 71 but the lumbar and sacral are so much anchylosed that I
am not sure of their number, and this makes the comparison of the
total number of vertebræ in the several breeds difficult. I have
spoken of six caudal vertebre, because the basal one is almost com-
pletely anchylosed with the pelvis; but if we consider the number
as seven, the caudal vertebræ agree in all the skeletons.
The cer-
vical vertebræ are, as just stated, in appearance fourteen; but out
of twenty-three skeletons in a fit state for examination, in five of
them, namely, in two Games, in two pencilled Hamburghs, and in
a Polish, the fourteenth vertebra bore ribs, which, though small
were perfectly developed with a double articulation. The presence
of these little ribs cannot be considered as a fact of much impor-
tance, for all the cervical vertebræ bear representatives of ribs; but
their development in the fourteenth vertebra reduces the size of
the passages in the transverse processes, and makes this vertebra
exactly like the first dorsal vertebra. The addition of these little
ribs does not affect the fourteenth cervical alone, for properly the
ribs of the first true dorsal vertebra are destitute of processes; but
in some of the skeletons in which the fourteenth cervical bore little
ribs, the first pair of true ribs had well-developed processes. When
we know that the sparrow has only nine, and the swan twenty-three
cervical vertebræ,72 we need feel no surprise at the number of the
cervical vertebræ in the fowl being, as it appears, variable.
There are seven dorsal vertebræ bearing ribs; the first dorsal is
never anchylosed with the succeeding four, which are generally
anchylosed together. In one Sultan fowl, however, the two first
dorsal vertebræ were free. In two skeletons, the fifth dorsal was
free; generally the sixth is free (as in G. bankiva), but sometimes
only at its posterior end, where in contact with the seventh. The
seventh dorsal vertebra, in every case excepting in one Spanish
cock, was anchylosed with the lumbar vertebræ. So that the degree
to which these middle dorsal vertebræ are anchylosed together is
variable.
Seven is the normal number of true ribs, but in two skeletons of
the Sultan fowl (in which the fourteenth cervical vertebra was not
furnished with little ribs) there were eight pairs; the eighth pair
seemed to be developed on a vertebra corresponding with the first lum-
bar in G.bankiva-, the sternal portion of both the seventh and eighth
ribs did not reach the sternum. In four skeletons in which ribs
71 It appears that I have not correctly
designated the several groups of verte-
bræ, for a great authority, Mr. W. K.
Parker (Transact. Zoolog. Soc.,' vol. v.
p. 198), specifies 16 cervical, 4 dorsal,
15 lumbar, and 6 caudal vertebræ in this
genus. But I have used the same terms
in all the following descriptions.
72 Macgillivray, 'British Birds,' vol.
i. p. 25.

CHAP. VII.
323
OSTEOLOGICAL DIFFERENCES.
were developed on the fourteenth cervical vertebra, there were, when
these cervical ribs are included, eight pairs; but in one Game-cock,
in which the fourteenth cervical was furnished with ribs, there were
only six pairs of true dorsal ribs; the sixth pair in this case did not
have processes, and thus resembled the seventh pair in other skele-
tons; in this game-cock, as far as could be judged from the appear-
ance of the lumbar vertebræ, a whole dorsal vertebra with its ribs
was missing. We thus see that the ribs (whether or not the little
pair attached to the fourteenth cervical vertebra be counted) vary
from six to eight pair. The sixth pair is frequently not furnished
with processes. The sternal portion of the seventh pair is extreme-
ly broad in Cochins, and is completely ossified. As previously stated,
it is scarcely possible to count the lumbo-sacral vertebræ ; but they
certainly do not correspond in shape or number in the several skele-
tons. The caudal vertebræ are closely similar in all the skeletons,
the only difference being, whether or not the basal one is anchylosed
to the pelvis; they hardly vary even in length, not being shorter in
Cochins, with their short tail-feathers, than in other breeds; in a
Spanish cock, however, the caudal vertebræ were a little elongated.
In three rumpless fowls the caudal vertebræ were few in number,
and anchylosed together into a misformed mass.
In the individual vertebræ the differences in structure are very
slight. In the atlas the cavity for the oc-
cipital condyle is either ossified into a ring,
or is, as in Bankiva, open on its upper
B
margin. The upper arc of the spinal ca-
nal is a little more arched in Cochins, in
conformity with the shape of occipital
foramen, than in G. bankiva. In several
skeletons a difference, but not of much
А
importance, may be observed, which com-
mences at the fourth cervical vertebra,
and is greatest at about the sixth, seventh, Fig. 87.—Sixth Cervical Ver-
tebra, of natural size, viewed
or eighth vertebra; this consists in the
laterally. A. Wild Gallus
hæmal descending processes being united vankiva. B. Cochin Cock.
to the body of the vertebra by a sort of
buttress. This structure may be observed in Cochins, Polish, some
Hamburghs, and probably other breeds; but is absent, or barely
developed, in Game, Dorking, Spanish, Bantam, and several other
breeds examined by me. On the dorsal surface of the sixth cervical
vertebra in Cochins three prominent points are more strongly de-
veloped than in the corresponding vertebra of the Game-fowl or G.
bankciva.
Pelvis.-- This differs in some few points in the several skeletons.

324
CHAP. VII.
- FOWLS.
A
The anterior margin of the ilium seems at first to vary much in
outline, but this is chiefly due to the degree to which the margin in
the middle part is ossified to the crest of the spine; the outline,
however, does differ in being more truncated in Bantams, and more
rounded in certain breeds, as in Cochins. The outline of the ischi-
adic foramen differs considerably, being nearly circular in Bantams,
instead of egg-shaped as in the Bankiva, and more regularly oval in
some skeletons, as in the Spanish. The obturator notch is also much
less elongated in some skeletons than in others. The end of the
pubic bone presents the greatest difference; being hardly enlarged
in the Bankiva; considerably and gradually enlarged in Cochins,
and in a lesser degree in some other breeds; and abruptly enlarged
in Bantams. In one Bantam this bone extended very little beyond
the extremity of the ischium. The whole pelvis in this latter bird
differed widely in its proportions, being far broader proportionally to
its length than in Bankiva.
Sternum.—This bone is generally so much deformed that it is
scarcely possible to compare its form strictly in the several breeds.
The shape of the triangular ex-
tremity of the lateral processes
differs considerably, being either
almost equilateral or much elon-
gated. The front margin of the
crest is more or less perpendicu-
lar and varies greatly, as does
the curvature of the posterior
end, and the flatness of the low-
er surface. The outline of the
manubrial process also varies,
being wedge-shaped in the Ban-
kiva, and rounded in the Spanish
breed. The furcula differs in be-
ing more or less arched, and
greatly, as may be seen in the
accompanying outlines, in the
shape of the terminal plate ; but
D the shape of this part differed a
little in two skeletons of the
wild Bankiva. The coracoids
Fig. 38.--Extremity of the Furcula, of
present no difference worth no-
natural size, viewed laterally. A. Wild tice. The scapula varies in shape,
Gallus bankiva. B. Spangled Polish being of nearly uniform breadth
Fowl. C. Spanish Fowl. D. Dorking
in Bankiva, much broader in the
Fowl.
middle in the Polish fowl, and
abruptly narrowed towards the apex in the two Sultan fowls.
B

CHAP. VII.
325
OSTEOLOGICAL DIFFERENCES.
I carefully compared each separate bone of the leg and wing, rela-
tively to the same bones in the wild Bankiva, in the following breeds,
which I thought were the most likely to differ ; namely, in Cochin,
Dorking, Spanish, Polish, Burmese Bantam, Frizzled Indian, and
black-boned Silk fowls; and it was truly surprising to see how abso-
lutely every process, articulation, and pore agreed, though the bones
differed greatly in size. The agreement is far more absolute than
in other parts of the skeleton. In stating this, I do not refer to the
relative thickness and length of the several bones; for the tarsi va-
ried considerably in both these respects. But the other limb-bones
varied little even in relative length.
Finally, I have not examined a sufficient number of
skeletons to say whether any of the foregoing differen-
ces except in the skull, are characteristic of the several
breeds. Apparently some differences are more common
in certain breeds than in others, -as an additional rib
to the fourteenth cervical vertebra in Hamburghs and
Games, and the breadth of the end of the pubic bone in
Cochins. Both skeletons of the Sultan fowl had eight
dorsal vertebræ, and the end of the scapula in both was
somewhat attenuated. In the skull, the deep medial fur-
row in the frontal bones and the vertically elongated oc-
cipital foramen seem to be characteristic of Cochins; as
is the great breadth of the frontal bones in Dorkings;
the separation and open spaces between the tips of the
ascending branches of the premaxillaries and nasal bones,
as well as the front part of the skull being but little de-
pressed, characterise Hamburghs; the globular shape of
the posterior part of the skull seems to be characteristic
of laced Bantams; and lastly, the protuberance of the
skull with the ascending branches of the premaxillaries
partially aborted, together with the other differences be-
fore specified, are eminently characteristic of Polish and
other Crested fowls.
But the most striking result of our examination of the
skeleton is the great variability of all the bones except
those of the extremities. To a certain extent we can
understand why the skeleton fluctuates so much in struc-

326
CHAP. VII.
FOWLS.
ture; fowls have been exposed to unnatural conditions
of life, and their whole organisation has thus been render-
ed variable; but the breeder is quite indifferent to, and
never intentionally selects, any modifications in the ske-
leton. External characters, if not attended to by man,-
.such as the number of the tail and wing feathers and
their relative lengths, which in wild birds are generally
constant points,-fluctuate in our domestic fowls in the
same manner as the several parts of the skeleton. An
additional toe is a "point" in Dorkings, and has become
a fixed character, but is variable in Cochins and Silk-
fowls. The colour of the plumage and the form of the
comb are in most breeds, or even sub-breeds, eminently
fixed characters ; but in Dorkings these points have not
been attended to, and are variable. When any modifica-
tion in the skeleton is related to some external charac-
ter which man values, it has been, unintentionally on his
part, acted on by selection, and has become more or less
fixed. We see this in the wonderful protuberance of the
skull, which supports the crest of feathers in Polish fowls,
and which by correlation has affected other parts of the
skull. We see the same result in the two protuberances
which support the horns in the horned fowl, and in the
flattened shape of the front of the skull in Hamburghs
consequent on their flattened and broad “rose-combs.”
We know not in the least whether additional ribs, or
the changed outline of the occipital foramen, or the
changed form of the scapula, or of the extremity of the
furcula, are in any way correlated with other structures,
or have arisen from the changed conditions and habits
of life to which our fowls have been subjected ; but there
is no reason to doubt that these various modifications in
the skeleton could be rendered, either by direct selection,
or by the selection of correlated structures, as constant
and as characteristic of each breed, as are the size and
shape of the body, the colour of the plumage, and the
form of the comb.

CHAP. VII.
327
THE EFFECTS OF DISUSE.
Effects of the Disuse of Parts.
Judging from the habits of our European gallinaceous birds, Gal-
lus bankiva in its native haunts would use its legs and wings more
than do our domestic fowls, which rarely fly except to their roosts.
The Silk and the Frizzled fowls, from having imperfect wing-feath-
ers, cannot fly at all; and there is reason to believe that both these
breeds are ancient, so that their progenitors during many genera-
tions cannot have flown. The Cochins, also, from their short wings
and heavy bodies, can hardly fly up to a low perch. Therefore in
these breeds, especially in the two first, a considerable diminution
in the wing-bones might have been expected, but this is not the
case. In every specimen, after disarticulating and cleaning the
bones, I carefully compared the relative length of the two main
bones of the wing to each other, and of the two main bones of the
leg to each other, with those of G.bankiva, and it was surprising to
see (except in the case of the tarsi) how exactly the same relative
length had been retained. This fact is curious, from showing how
truly the proportions of an organ may be inherited, although not fully
exercised during many generations. I then compared in several
breeds the length of the femur and tibia with the humerus and ulna
and likewise these same bones with those of G.bankiva ; the result
was that the wing bones in all the breeds (except the Burinese
Jumper, which has unnaturally short legs) are slightly shortened re-
latively to the leg-bones; but the decrease is so slight that it may be
due to the standard specimen of G. bankiva having accidentally had
wings of slightly greater length than usual; so that the measure-
ments are not worth giving. But it deserves notice that the Silk
and Frizzled fowls, which are quite incapable of flight, had their
wings less reduced relatively to their legs than in almost any other
breed! We have seen with domesticated pigeons that the bones of
the wings are somewhat reduced in length, whilst the primary
feathers are rather increased in length, and it is just possible, though
not probable, that in the Silk and Frizzled fowls any tendency to
decrease in the length of the wing-bones from disuse may have
been checked through the law of compensation, by the decreased
growth of the wing-feathers, and consequent increased supply of
nutriment. The wing-bones, however, in both these breeds, are
found to be slightly reduced in length when judged by the standard
of the length of the sternum or head, relatively to these same parts
in G. bankiva.
The actual weight of the main bones of the leg and wing in
twelve breeds is given in the two first columns in the following
table. The calculated weight of the wing-bones relatively to the

328
CHAP. VII.
FOWLS.
leg-bones, in comparison with the leg and wing-bones of G.bankiva,
are given in the third column,—the weight of the wing-bones in
G. bankiva being called a hundred."
73
TABLE I.
Names of Breeds.
Weight of
Wing-
bones rela-
tively to
Actual Actual
the Leg-
Weight of Weight of
Femur and Humerus
bones, in
compari-
Tibia. and Ulna.
son with
these same
bones in G.
bankiva.
Gallus bankiva .. wild male
1
Cochin ..
male
2 Dorking
male
3 Spanish (Minorca) male
4 Gold Spangled Polish male
5 Game, black-breasted male
6 Malay
female
7 Sultan
8 Indian Frizzled .. .. male
9 Burmese Jumper female
10 Hamburgh (pencilled) male
11 Hamburgh (pencilled) female
12 Silk (black-boned) .. female
Grains.
86
311
557
386
306
293
231
189
206
53
157
114
88
Grains.
54
162
248
183
145
143
116
94
88
36
104
77
57
100
83
70
75
75
77
80
79
67
108
106
108
103
.. male
In the eight first birds, belonging to distinct breeds, in this table,
we see a decided reduction in the weight of the bones of the wing.
In the Indian Frizzled fowl, which cannot fly, the reduction is car-
ried to the greatest extent, namely, to thirty-three per cent. of their
proper proportional weight. In the next four birds, including the
Silk-hen, which is incapable of flight, we see that the wings, rela-
tively to the legs, are slightly increased in weight; but it should be
73 It may be well to explain how the
calculation has been made for the third
column. In G. bankiva the leg-bones
are to the wing-bones as 86 : 54, or
as (neglecting decimals) 100: 62 ;-in
Cochins as 311 : 162, or as 100:52 ;-in
Dorkings as 557; 248, or as 100: 44;
and so on for the other breeds. We thus
get the series of 62, 52, 44 for the relative
weights of the wing-bcnes in G. bankiva,
Cochins, Dorkings, &c. And now tak-
ing 100, instead of 62, for the weight of
the wing-bones in G. bankiva, we get,
by another rule of three, 83 as the weight
of the wing-bones in Cochins; 70 in the
Dorkings; and so on for the remainder
of the third column in the table.

CHAP. VII.
329
THE EFFECTS OF DISUSE.
observed that, if in these birds the legs had become from any cause
reduced in weight, this would give the false appearance of the wings
having increased in relative weight. Now a reduction of this na-
ture has certainly occurred with the Burmese Jumper, in which the
legs are abnormally short, and in the two Hamburghs and Silk
fowl, the legs, though not short, are formed of remarkably thin and
light bones. I make these statements, not judging by mere eye-
sight, but after having calculated the weights of the leg-bones rela-
tively to those of G. bankiva, according to the only two standards
of comparison which I could use, namely, the relative lengths of
the head and sternum ; for I do not know the weight of the body in
G. bankiva, which would have been a better standard. According
to these standards, the leg-bones in these four fowls are in a marked
manner far lighter than in any other breed. It may therefore be
concluded that in all cases in which the legs have not been through
some unknown cause much reduced in weight, the wing-bones have
become reduced in weight relatively to the leg-bones, in comparison
with those of G. bankiva. And this reduction of weight may, I ap-
prehend, safely be attributed to disuse.
To make the foregoing table quite satisfactory, it ought to have
been shown that in the eight first birds the leg-bones have not ac-
tually increased in weight out of due proportion with the rest of the
body; this I cannot show, from not knowing, as already remarked,
the weight of the wild Bankiva.74 I am indeed inclined to suspect
that the leg-bones in the Dorking, No. 2 in the table, are proportion-
ally too heavy; but this bird was a very large one, weighing 7 lb.
2 oz., though very thin. Its leg-bones were more than ten times as
heavy as those of the Burmese Jumper! I tried to ascertain the
length both of the leg-bones and wing-bones relatively to other
parts of the body and skeleton ; but the whole organisation in these
birds, which have been so long domesticated, has become so variable,
that no certain conclusions could be reached. For instance, the legs
of the above Dorking cock were nearly three-quarters of an inch too
short relatively to the length of the sternum; and more than three-
quarters of an inch too long relatively to the length of the skull, in
comparison with these same parts in G. bankiva.
In the following Table II. in the two first columns we see in inches
and decimals the length of the sternum, and the extreme depth of
its crest to which the pectoral muscles are attached. In the third
74 Mr. Blyth (in Annals and Mag. of
Nat. Hist.,' 2nd series, vol. i., 1848, p.
456) gives 31 lb. as the weight of a full-
grown male G. bankiva; but from what
I have seen of the skins and skeletons
of various breeds, I cannot believe that
my two specimens of G. bankiva could
have weighed so much,

330
CHAP. VII.
FOWLS.
column we have the calculated depth of the crest, relatively to the
length of the sternum, in comparison with these same parts in G.
bankiva.75
TABLE II.
Names of Breeds.
Length
of
Sternum.
Depth of Crest,
Depth of relatively to the
Crest length of the
of Sternum, in
Sternum. comparison with
G. bankiva.
..
.. male
Gallus bankiva .. male
1 Cochin
.. male
2 Dorking
.. male
3 Spanish
male
4 Polish ..
5 Game ..
.. male
6 Malay..
female
7 Sultan
8 Frizzled hen
9 Burmese Jumper .. female
10 Hamburgh .. male
11 Hamburgh
female
12 Silk fowl ..
.. female
Inches.
4:20
5.83
6.95
6:10
5.07
5:55
5:10
4:47
4.25
3:06
5.08
4.55
4:49
Inches.
1:40
1:55
1.97
1.83
1:50
1:55
1:50
1:36
1.20
0.85
1:40
1.26
1:01
100
78
84
90
87
81
87
90
84
81
81
81
66
.. male
.. male
By looking to the third column we see that in every case the depth
of the crest relatively to the length of the sternum, in comparison
with G. bankiva, is diminished, generally between 10 and 20 per cent.
But the degree of reduction varies much, partly in consequence of
the frequently deformed state of the sternum. In the Silk-fowl,
which cannot fly, the crest is 34 per cent. less deep than what it
ought to have been. This reduction of the crest in all the breeds
probably accounts for the great variability, before referred to in the
curvature of the furcula, and in the shape of its sternal extremity.
Medical men believe that the abnormal form of the spine so com-
monly observed in women of the higher ranks results from the at-
tached muscles not being fully exercised. So it is with our domes-
tic fowls, for they use their pectoral muscles but little, and, out of
twenty-five sternums examined by me, three alone were perfectly
symmetrical, ten were moderately crooked, and twelve were de-
formed to an extreme degree.
Finally, we may conclude with respect to the various
75 The third column is calculated on the same principle as explained in the pre-
vious foot-note, p. 328.

CHAP. VII.
331
CORRELATION OF GROWTH.
breeds of the fowl, that the main bones of the wing have
probably been shortened in a very slight degree; that
they have certainly become lighter relatively to the leg-
bones in all the breeds in which these latter bones are
not unnaturally short or delicate; and that the crest of
the sternum, to which the pectoral muscles are attached,
has invariably become less prominent, the whole sternum
being also extremely liable to deformity. These results
we may attribute to the lessened use of the wings.
Correlation of Growth.—I will here sum up the few
facts which I have collected on this obscure, but impor-
tant, subject. In Cochins and Game-fowls there is some
relation between the colour of the plumage and the dark-
ness of the egg-shell and even of the yolk. In Sultans
the additional sickle-feathers in the tail are apparently
related to the general redundancy of the plumage, as
shown by the feathered legs, large crest, and beard. In
two tailless fowls which I examined the oil-gland was
aborted. A large crest of feathers, as Mr. Tegetmeier
has remarked, seems always accompanied by a great
diminution or almost entire absence of the comb. A
large beard is similarly accompanied by diminished or ab-
sent wattles. These latter cases apparently come under
the law of compensation or balancement of growth. A
large beard beneath the lower jaw and a large top-knot
on the skull often go together. The comb when of any
peculiar shape, as with Horned, Spanish, and Hamburgh
fowls, affects in a corresponding manner the underlying
skull; and we have seen how wonderfully this is the case
with Crested fowls when the crest is largely developed.
With the protuberance of the frontal bones the shape of
the internal surface of the skull and of the brain is great-
ly modified. The presence of a crest influences in some
unknown way the development of the ascending branches
of the premaxillary bone, and of the inner processes of
the nasal bones; and likewise the shape of the external
orifice of the nostrils. There is a plain and curious corre-

332
CHAP. VII.
FOWLS.
lation between a crest of feathers and the imperfectly os-
sified condition of the skull. Not only does this hold
good with nearly all crested fowls, but likewise with tuft-
ed ducks, and as Dr. Günther informs me with tufted
geese in Germany.
Lastly, the feathers composing the crest in male Polish
fowls resemble hackles, and differ greatly in shape from
those in the crest of the female. The neck, wing-coverts,
and loins in the male bird are properly covered with
hackles, and it would appear that feathers of this shape
have spread by correlation to the head of the male. This
little fact is interesting; because, though both sexes of
some wild gallinaceous birds have their heads similarly
ornamented, yet there is often a difference in the size and
shape of feathers forming their crests. Furthermore
there is in some cases, as in the male Gold and in the male
Amherst pheasants (P. pictus and Amherstic), a close
relation in colour, as well as in structure, between the
plumes on the head and on the loins. Hence it would
appear that the same law has regulated the state of the
feathers on the head and body, both with species living
under their natural conditions, and with birds which have
varied under domestication.

CHAP. VIII.
333
DOMESTIC DUCKS.
CHAPTER VIII.
DUCKS-GOOSE - PEACOCK_TURKEY_GUINEA-FOWL-
CANARY-BIRD-GOLD-FISH – HIVE-BEES — SILK-MOTHS.
DUCKS, SEVERAL BREEDS OF — PROGRESS OF DOMESTICATION -
ORIGIN OF, FROM THE COMMON WILD-DUCK — DIFFERENCES IN
THE DIFFERENT BREEDS - OSTEOLOGICAL DIFFERENCES —EF-
FECTS OF USE AND DISUSE ON THE LIMB-BONES.
GOOSE, ANCIENTLY DOMESTICATED — LITTLE VARIATION OT — SE-
BASTOPOL BREED.
PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.
TURKEY, BREEDS OF — CROSSED WITH THE UNITED STATES SPE-
CIES — EFFECTS OF CLIMATE ON.
GUINEA-FOWL, CANARY-BIRD, GOLD-FISH, HIVE-BEES.
SILK-MOTHS, SPECIES AND BREEDS OF — ANCIENTLY DOMESTI-
CATED 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.
I
WILL, as in previous cases, first briefly describe the
chief domestic breeds of the duck:-
BREED 1. Common Domestic Duck.- Varies much in colour and in
proportions, and differs in instincts and disposition from the wild-
duck. There are several sub-breeds :--(1) The Aylesbury, of great
size, white, with pale-yellow beak and legs; abdominal sack largely
developed. (2) The Rouen, of great size, coloured like the wild-
duck, with green or mottled beak; abdominal sack largely devel-
oped. (3) Tufted Duck, with a large top-knot of fine downy
feathers, supported on a fleshy mass, with the skull perforated be-
neath. The top-knot in a duck which I imported from Holland was
two and a half inches in diameter. (4) Labrador (or Canadian, or
Buenos Ayres, or East Indian); plumage entirely black; beak

334
CHLAP. VIII.
DOMESTIC DUCKS.
broader, relatively to its length, than in the wild-duck; eggs
slightly tinted with black. This sub-breed perhaps ought to be
ranked as a breed; it includes two sub-varieties, one as large as the
common domestic duck, which I have kept alive, and the other
smaller and often capable of flight. I presume it is this latter sub-
variety which has been described in France 2 as flying well, being
rather wild, and when cooked having the flavour of the wild-duck ;
nevertheless this sub-variety is polygamous, like other domesticated
ducks and unlike the wild-duck. These black Labrador ducks breed
true; but a case is given by Dr. Turral of the French sub-variety
producing young with some white feathers on the head and neck,
and with an ochre-coloured patch on the breast.
BREED 2. Hook-billed Duck.--This bird presents an extraordinary
appearance from the downward curvature of the beak. The head
is often tufted. The common colour is white, but some are coloured
like wild-ducks. It is an ancient breed, having been noticed in
1676. It shows its prolonged domestication by almost incessantly
laying eggs, like the fowls which are called everlasting layers.
BREED 3. Call-Duck. Remarkable from its small size, and from
the extraordinary loquacity of the female. Beak short. These
birds are either white, or coloured like the wild-duck.
BREED 4. Penguin Duck. This is the most remarkable of all
the breeds, and seems to have originated in the Malayan archipela
go. It walks with its body extremely erect, and with its thin neck
stretched straight upwards. Beak rather short. Tail upturned, in-
cluding only 18 feathers. Femur and metatarsi elongated.
Almost all naturalists admit that the several breeds
are descended from the common wild duck (Anas bos-
chas) ; most fanciers, on the other hand, take as usual a
very different view. Unless we deny that domestica-
tion, prolonged during centuries, can affect even such
unimportant characters as colour, size, and in a slight
degree proportional dimensions and mental disposition,
1 Poultry Chronicle' (1854), vol. ii. p.
91, and vol. i. p. 330.
2 Dr. Turral, in 'Bull. Soc. d'Acclimat.,'
tom. vii., 1860, p. 541.
3 Willughby's Ornithology,' by Ray,
p. 381. This breed is also figured by
Albin, in 1734, in his 'Nat. Hist. of Birds,'
vol. ii. p. 86.
4 F. Cuvier, in ' Annales du Muséum,
tom. ix. p. 128, says that moulting and
incubation alone stop these ducks laying.
Mr. B. P. Brent makes a similar remark
in the Poultry Chronicle, 1855, vol.
iii. p. 512
5 Rev. E. S. Dixon, Ornamental and
Domestic Poultry' (1848), p. 117. Dr. B.
P. Brent, in Poultry Chronicle,' vol.iii.
1855, p. 512.

CHAP. VIII.
335
EXTERNAL DIFFERENCES.
there is no reason whatever to doubt that the domestic
duck is descended from the common wild species, for the
one differs from the other in no important character.
We have some historical evidence with respect to the
period and progress of the domestication of the duck.
It was unknown to the ancient Egyptians, to the Jews
of the Old Testament, and to the Greeks of the Homeric
period. About eighteen centuries ago Columella' and
Varro speak of the necessity of keeping ducks in netted
enclosures like other wild fowl, so that at this period
there was danger of their flying away. Moreover, the
plan recommended by Columella to those who might
wish to increase their stock of ducks, namely, to collect
the eggs of the wild bird and to place them under a hen,
shows, as Mr. Dixon remarks, “that the duck had not at
this time become a naturalised and prolific inmate of the
Roman poultry-yard." The origin of the domestic duck
from the wild species is recognised in nearly every lan-
guage of Europe, as Aldrovandi long ago remarked, by
the same name being applied to both. The wild duck
has a wide range from the Himalayas to North America.
It crosses readily with the domestic bird, and the crossed
offspring are perfectly fertile.
Both in North America and Europe the wild duck has
been found easy to tame and breed. In Sweden this ex-
periment was carefully tried by Tiburtius; he succeeded
in rearing wild ducks for three generations, but, though
they were treated like common ducks, they did not vary
even in a single feather. The young birds suffered from
being allowed to swim about in cold water, as is known
6 Crawfurd on the Relation of Domes-
ticated Animals to Civilisation,' read be-
fore the Brit. Assoc. at Oxford.
7 Dureau de la Malle, in 'Annales des
Sciences Nat.,' tom. xvii. p. 164; and
tom. xxi. p. 55. Rev. E. S. Dixon, 'Or-
namental Poultry,' p. 118. Tame ducks
were not known in Aristotle's time, as re-
marked by Volz, in his Beiträge zur
Kulturgeschichte, 1852, s. 78.
8 I quote this account from 'Die En-
ten, Schwanen-zucht, Ulm, 1828, s. 143.
See Audubon's Ornithological Biogra-
phy,' vol. iii., p. 168, on the taming of
ducks on the Mississippi. For the same
fact in England, see Mr. Waterton, in

336
CITAP. VIII.
DOMESTIC DUCKS.
to be the case, though the fact is a strange one, with the
young of the common domestic duck. An accurate and
well-known observer in England has described in detail
his often repeated and successful experiments in domes-
ticating the wild duck. Young birds are easily reared
from
eggs hatched under a bantam; but to succeed it is
indispensable not to place the eggs of both the wild and
tame duck under the same hen, for in this case “the
young wild ducks die off, leaving their more hardy breth-
ren in undisturbed possession of their foster-mother's
The difference of habit at the onset in the newly-
hatched ducklings almost entails such a result to a cer-
tainty.” The wild ducklings were from the first quite
tame towards those who took care of them as long as
they wore the same clothes, and likewise to the dogs and
cats of the house. They would even snap with their
beaks at the dogs, and drive them away from any spot
which they coveted. But they were much alarmed at
strange men and dogs. Differently from what occurred
in Sweden, Mr. Hewitt found that his young birds al-
ways changed and deteriorated in character in the course
of two or three generations; notwithstanding that great
care was taken to prevent any crossing with tame ducks.
After the third generation his birds lost the elegant car-
riage of the wild species, and began to acquire the gait
of the common duck. They increased in size in each
generation, and their legs became less fine. The white
collar round the neck of the mallard became broader and
less regular, and some of the longer primary wing-feathers
became more or less white. When this occurred, Mr.
Hewitt always destroyed his old stock and procured
fresh eggs from wild nests; so that he never bred the
same family for more than five or six generations. His
care.
Loudon's 'Mag. of Nat. Hist.,' vol. viii.
1835, p. 542; and Mr. St. John, Wild
Sports and Nat. Hist. of the Highlands,
1846, p. 129.
9 Mr. E. Hewitt, in 'Journal of Horti-
culture,' 1862, p. 773; and 1863, p. 39.

CAAP, VIII.
337
EXTERNAL DIFFERENCES.
birds continued to pair together, and never became poly-
gamous like the common domestic duck. I have given
these details, because no other case, as far as I know, has
been so carefully recorded by a competent observer of
the progress of change in wild birds reared for several
generations in a domestic condition.
From these considerations there can hardly be a doubt
that the wild duck is the parent of the common domestic
kind; nor need we look to distinct species for the parent-
age of the more distinct breeds, namely, Penguin, Call,
Hook-billed, Tufted, and Labrador ducks. I will not re-
peat the arguments used in the previous chapters on the
improbability of man having in ancient times domesti-
cated several species since become unknown or extinct,
though ducks are not readily exterminated in the wild
state ;-on some of the supposed parent-species having
had abnormal characters in comparison with all the other
species of the genus, as with hook-billed and penguin
ducks ;-on all the breeds, as far as is known, being fer-
tile together;10—on all the breeds having the same gene-
ral disposition, instinct, &c. But one fact bearing on this
question may be noticed: in the great duck family, one
species alone, namely, the male of A. boschas, has its
four middle tail feathers curled upwardly; now in every
one of the above-named domestic breeds these curled
feathers exist, and on the supposition that they are de-
scended from distinct species, we must assume that man
formerly hit upon species all of which had this now uni-
que character. Moreover, sub-varieties of each breed are
coloured almost exactly like the wild duck, as I have
10 I have met with several statements
on the fertility of the several breeds
when crossed. Mr. Yarrell assured me
that Call and common ducks are per-
fectly fertile together. I crossed Hook-
billed and common ducks, and a Penguin
and Labrador, and the crossed ducks
were quite fertile, though they were not
bred inter se, so that the experiment
was not fully tried. Some half-bred Pen-
guins and Labradors were again crossed
with Penguins, and subsequently bred
by me inter se, and they were extremely
fertile.
15

338
CHAP. VIII.
DOMESTIC DUCKS.
seen with the largest and smallest breeds, namely Rouens
and Call-ducks, and, as Mr. Brent states," is the case
with Hook-billed ducks. This gentleman, as he informs
me, crossed a white Aylesbury drake and a black Labra-
dor duck, and some of the ducklings as they grew up
assumed the plumage of the wild duck.
With respect to Penguins, I have not seen many speci-
mens, and none were coloured precisely like the wild
duck; but Sir James Brooke sent me three skins from
Lombok and Bali, in the Malayan archipelago; the two
females were paler and more rufous than the wild duck,
and the drake differed in having the whole under and
upper 'surface (excepting the neck, tail-coverts, tail, and
wings) silver-grey, finely pencilled with dark lines, closely
like certain parts of the plumage of the wild mallard.
But I found this drake to be identical in every feather
with a variety of the common breed procured from a
farm-yard in Kent, and I have occasionally elsewhere
seen similar specimens. The occurrence of a duck bred
under so peculiar a climate as that of the Malayan archi-
pelago, where the wild species does not exist, with ex-
actly the same plumage as may occasionally be seen in
our farm-yards, is a fact worth notice. Nevertheless the
climate of the Malayan archipelago apparently does tend
to cause the duck to vary much, for Zollinger," speaking
of the Penguin breed, says that in Lombok “there is an
unusual and very wonderful variety of ducks.” One Pen-
guin drake which I kept alive differed from those of
which the skins were sent me from Lombok, in having
its breast and back partially coloured with chesnut-
brown, thus more closely resembling the Mallard.
For these several facts, more especially from the drakes
of all the breeds having curled tail-feathers, and from
certain sub-varieties in each breed occasionally resem-
11 Poultry Chronicle,' 1855, vol. iii.
12 Journal of the Indian Archipelą.
go,' vol. v. p. 334.
p. 512.

CHAP. VIII.
339
EXTERNAL DIFFERENCES.
bling in general plumage the wild duck, we may conclude
with confidence that all the breeds are descended from
A. boschae.
I will now notice some of the peculiarities characteristic of the
several breeds. The eggs vary in colour; some common ducks
laying pale-greenish and others quite white eggs. The eggs which
are first laid during each season by the black Labrador duck, are
tinted black, as if rubbed with ink. So that with ducks, as with
poultry, some degree of correlation exists between the colour of the
plumage and the egg-shell. A good observer assured me that one
year his Labrador ducks laid almost perfectly white eggs, but that
the yolks were this same season dirty olive-green, instead of as
usual of a golden yellow, so that the black tint appeared to have
passed inwards. Another curious case shows what singular varia-
tions sometimes occur and are inherited ; Mr. Hansell 13 relates that
he had a common duck which always laid eggs with the yolk of a
dark brown colour like melted glue; and the young ducks, hatched
from these eggs, laid the same kind of eggs, so that the breed had
to be destroyed.
The hooked-billed duck has a most remarkable appearance (see
fig. of skull, woodcut No. 39); and its peculiar beak has been in-
herited at least since the year 1676. This structure is evidently
analogous with that described in the Bagadotten carrier pigeon.
Mr. Brent 14 says that, when hook-billed ducks are crossed with com-
mon ducks,“ many young ones are produced with the upper man-
dible shorter than the lower, which not unfrequently causes the
death of the bird." A tuft of feathers on the head is by no means
a rare occurrence; namely, in the true tufted breed, the hook-billed,
the common farmyard duck, and in a duck having no other pecu-
liarity which was sent to me from the Malayan archipelago. The
tuft is only so far interesting as it affects the skull, which is thus
rendered slightly more globular, and is perforated by numerous
apertures. Call-ducks are remarkable from their extraordinary
loquacity; the drake only hisses like common drakes; nevertheless,
when paired with the common duck, he transmits to his female
offspring a strong quacking tendency. This loquacity seems at first
a surprising character to have been acquired under domestication.
14 Poultry Chronicle,' 1855, vol. iii,
18 The Zoologist,' vols. vii., vüi.
(1849-1850), p. 2853.
p. 512.

340
CHAP. VIII.
DOMESTIC DUCKS.
But the voice varies in the different breeds; Mr. Brent 15 says that
hook-billed ducks are very loquacious, and that Rouens utter a
dull, loud, and monotonous cry, easily distinguishable by an ex-
perienced ear." As the loquacity of the Call-duck is highly ser-
viceable, these birds being used in decoys, this quality may have
been increased by selection. For instance, Colonel Hawker says, if
young wild-ducks cannot be got for a decoy," by way of make-shift,
select tame birds which are the most clamorous, even if their colour
should not be like that of wild ones. 16 It has been falsely asserted
thai Call-ducks hatch their eggs in less time than common ducks.17
B
A
Fig. 39.-Skulls, viewed laterally, reduced to two-thirds of the natural size.
A. Wild Duck. B. Hook-billed Duck.
The Penguin duck is the most remarkable of all the breeds; the
thin neck and body are carried erect; the wings are small; the tail
is upturned, and the thigh bones and metatarsi are considerably
lengthened in proportion with the same bones in the wild duck. In
five specimens examined by me there were only eighteen tail-feathers
instead of twenty as in the wild duck; but I have also found only
eighteen and nineteen tail-feathers in two Labrador ducks. On the
15 Poultry Chronicle,' vol. iii., 1855,
p. 312. With respect to Rouens, see
ditto, vol. i., 1854, p. 167.
16 Col. Hawker's Instructions to
young Sportsmen,' quoted by Mr. Dixon
in his Ornamental Poultry,' p. 125.
17 Cottage Gardener,' April 9th,
1861.

CHAP. VIII.
DIFFERENCES IN THEIR SKELETONS. 341
middle toe, in three specimens, there were twenty-seven or twenty-
eight scutellæ, whereas in two wild ducks there were thirty-one
and thirty-two. The Penguin when crossed transmits with much
power its peculiar form of body and gait to its offspring; this was
manifest with some hybrids raised in the Zoological Gardens, be-
tween one of these birds and the Egyptian goose 18 (Tadorna Ægyp-
tiaca), and likewise with some mongrels which I raised between
the Penguin and Labrador duck. I am not much surprised that
some writers have maintained that this breed must be descended
from an unknown and distinct species; but from the reasons already
assigned, it seems to me far more probable that it is the descend-
ant, much modified by domestication under an unnatural chimate,
of Anas boschas.
Osteological Characters. The skulls of the several different breeds
differ from each other and from the skull of the wild duck in
very
lit-
tle except in the proportional length and curvature of the premaxil-
laries. These latter bones in the Call-duck are short, and a line
drawn from their extremities to the summit of the skull is nearly
straight, instead of being concave as in the common duck; so that the
skull resembles that of a small goose. In the hook-billed duck (fig. 39)
these same bones as well as the lower jaw curve downwards in a
most remarkable manner, as represented. In the Labrador duck
the premaxillaries are rather broader than in the wild duck ; and
in two skulls of this breed the vertical ridges on each side of the
supra-occipital bone are very prominent. In the Penguin the pre-
maxillaries are relatively shorter than in the wild duck; and the in-
ferior points of the paramastoids more prominent. In a Dutch tufted
duck, the skull under the enormous tuft was slightly more globular
and was perforated by two large apertures; in this skull the lachry-
mal bones were produced much further backwards, so as to have a
different shape and to nearly touch the post. lat. processes of the
frontal bones, thus almost completing the bony orbit of the eye.
As the quadrate and pterygoid bones are of such complex shape
and stand in relation with so many other bones, I carefully com-
pared them in all the principal breeds; but excepting in size they
presented no difference.
Vertebræ and Ribs.In one skeleton of the Labrador duck there
were the usual fifteen cervical vertebræ and the usual nine dorsal
vertebræ bearing ribs; in the other skeleton there were fifteen cervical
and ten dorsal vertebræ with ribs; nor, as far as could be judged, was
18 These hybrids have been described
by M. Selys-Longchamps in the 'Bul-
letins (tom. xii. No. 10) Acad. Roy. de
Bruxelles.'

342
CHAP. VIII.
DOMESTIC DUCKS.
this owing merely to a rib having
been developed on the first lumbar
vertebra ; for in both skeletons the
B
A
lumbar vertebræ agreed perfectly
in number, shape, and size with
those of the wild duck. In two
skeletons of the Call-duck there
were fifteen cervical and nine dor-
sal vertebra; in a third skeleton
D
small ribs were attached to the so- o
called fifteenth cervical vertebra,
making ten pairs of ribs; but these
ten ribs do not correspond, or arise
from the same vertebrae, with the
ten in the above-mentioned Labra-
dor duck. In the Call-duck, which
Fig. 40.--Cervical Vertebræ, of natural
had small ribs attached to the fif- size. A. Eighth cervical vertebra of
teenth cervical vertebra, the hæ-
Wild Duck, viewed on hæmal sur-
mal spines of the thirteenth and
face. B. Eighth cervical vertebra of
Call Duck, viewed as above. C.
fourteenth (cervical) and of the
Twelfth cervical vertebra of Wild
seventeenth (dorsal) vertebræ cor- Duck, viewed laterally. D. Twelfth
responded with the spines on the
cervical vertebra of Aylesbury Duck,
viewed laterally.
fourteenth, fifteenth, and eigh-
teenth vertebræ of the wild duck: so that each of these vertebræ had
acquired a structure proper to one posterior to it in position. In the
twelfth cervical vertebra of this same Call-duck (fig. 40, B), the two
branches of the hæmal spine stand much closer together than in the
wild duck (A), and the descending hæmal processes are much short-
ened. In the Penguin duck the neck from its thinness and erect-
ness falsely appears (as ascertained by measurement) to be much
elongated, but the cervical and dorsal vertebræ present no differ-
ence; the posterior dorsal vertebræ, however, are more completely
anchylosed to the pelvis than in the wild duck. The Aylesbury duck
has fifteen cervical and ten dorsal vertebræ furnished with ribs, but
the same number of lumbar, sacral, and caudal vertebra, as far as
could be traced, as in the wild duck. The cervical vertebræ in this
same duck (fig. 40, D) were much broader and thicker relatively to
their length than in the wild (C); so much so, that I have thought
it worth while to give a sketch of the eighth cervical vertebra in
these two birds. From the foregoing statements we see that the fif-
teenth cervical vertebra occasionally becomes modified into a dorsal
vertebra, and when this occurs all the adjoining vertebræ are modi-
fied. We also see that an additional dorsal vertebra bearing a rib is

CHAP. VIII.
343
EFFECTS OF USE AND DISUSE.
occasionally developed, the number of the cervical and lumbar verte-
bræ apparently remaining the same as usual.
I examined the bony enlargement of the trachea in the males of
the Penguin, Call, Hook-billed, Labrador, and Aylesbury breeds; and
in all it was identical in shape.
The Pelvis is remarkably uniform ; but in the skeleton of the
Hook-billed duck the anterior part is much bowed inwards; in the
Aylesbury and some other breeds the ischiadic foramen is less elon-
gated. In the sternum, furcula, coracoids, and scapula, the differ
ences are so slight and so variable as not to be worth notice, except
that in two skeletons of the Penguin duck the terminal portion of
the scapula was much attenuated.
In the bones of the leg and wing no modification in shape could
be observed. But in Peguin and Hook-billed ducks, the terminal
phalanges of the wing are a little shortened. In the former, the
femur and metatarsus (but not the tibia) are considerably lengthened,
relatively to the same bones in the wild duck, and to the wing-bones
in both birds. This elongation of the leg-bones could be seen whilst
the bird was alive, and is no doubt connected with its peculiar up-
right manner of walking. In a large Aylesbury duck, on the other
hand, the tibia was the only bone of the leg which relatively to the
other bones was slightly lengthened.
On the effects of the increased and decreased Use of the Limbs.-
In all the breeds the bones of the wing (measured separately after
having been cleaned) relatively to those of the leg have become
slightly shortened, in comparison with the same bones in the wild
duck, as may be seen in the following table :-
Name of Breed.
Length of Femur, Length of Humerus,
Tibia, and Meta- Radius, and Meta-
tarsus together.
carpus together.
Or as
Wild mallard
Aylesbury
Tufted (Dutch)
Penguin
Call
Inches.
7:14
8.64
8.25
7.12
6:20
Inches.
9.28
10:43
9.83
8.78
7.77
100 : 129
100 - 120
100 : 119
100 : 123
100 : 125
Length of same
Bones.
Length of all the
Bones of Wing.
Inches.
Inches.
6.85
Wild duck (another
specimen) ..
Common domestic duck
10-07
11.26
100: 147
100 : 138
8:15

344
CHAP. VIII.
DOMESTIC DUCKS.
In the foregoing table we see that, in comparison with the wild
duck, the reduction in the length of the bones of the wing, rela-
tively to those of the legs, though slight, is universal. The reduc-
tion is least in the Call-duck, which has the power and the habit of
frequently flying
In weight there is a greater relative difference between the bones
of the leg and wing, as may be seen in the following table -
Name of Breed.
Weight of Femur,
Tibia, and
Metatarsus.
Weight of
Humerus, Radius,
and Metacarpus.
Or as
Wild mallard
Aylesbury
Hooked-bill
Tufted (Dutch)
Penguin
Labrador
Grains.
54
164
107
111
75
141
Grains.
97
204
160
148
90-5
165
93
100 : 179
100 : 124
100 : 149
100 : 133
100 : 120
100 : 117
100 : 163
Call ..
Weight of all the
Bones of the
Leg and Foot.
Weight of all the
Bones of the
Wing.
Grains,
Grains.
Wild (another speci-
66
115
100 : 173
men)
Common domestic duck
127
158
100 : 124
In these domesticated birds, the considerably lessened weight of
the bones of the wing (i. e. on an average, twenty-five per cent. of
their proper proportional weight), as well as their slightly lessened
length, relatively to the leg-bones, might follow, not from any ac-
tual decrease in the wing-bones, but from the increased weight and
length of the bones of the legs. The first of the two tables on the
next page shows that the leg-bones relatively to the weight of the
entire skeleton have really increased in weight; but the second
table shows that according to the same standard the wing-bones
have also really decreased in weight; so that the relative dispro-
portion shown in the foregoing tables between the wing and leg-
bones, in comparison with those of the wild duck, is partly due to
the increase in weight and length of the leg-bones, and partly to
the decrease in weight and length of the wing-bones.
With respect to the two following tables I may first state that I
tested them by taking another skeleton of a wild duck and of a

CHAP. VIII.
345
EFFECTS OF USE AND DISUSE.
common domestic duck, and by comparing the weight of all the
bones of the leg with all those of the wings, and the result was the
same. In the first of these tables we see that the leg-bones in each
case have increased in actual weight. It might have been expected
that, with the increased or decreased weight of the entire skeleton,
the leg-bones would have become proportionally heavier or lighter;
but their greater weight in all the breeds relatively to the other
bones can be accounted for only by these domestic birds having
used their legs in walking and standing much more than the wild,
for they never fly, and the more artificial breeds rarely swim. In
Name of Breed.
Weight of entire
Skeleton
(N. B. One Meta-
tarsus and Foot
was removed
Weight of
from each skele-
Femur, Tibia,
and Metatarsus.
ton, as it had
been acciden-
tally lost in two
cases.)
Or as
Wild mallard
Aylesbury
Tufted (Dutch)
Penguin
Call (from Mr. Fox)
Grains.
839
1925
1404
871
717
Grains.
54
164
111
75
57
1000 : 64
1000 : 85
1000 : 79
1000 : 86
1000 : 79
Weight of
Weight of Skele- Humerus, Ra-
ton as above.
dius and Ulna,
and Metacarpus.
Wild mallard
Aylesbury
Tufted (Dutch)
Penguin
Call (from Mr. Baker)
Call (from Mr. Fox)
Grains.
839
1925
1404
871
914
717
Grains.
97
204
148
90
100
92
1000 : 115
1000 : 105
1000 : 105
1000 : 103
1000 : 109
1000 : 129
. .
the second table we see, with the exception of one case, a plain re-
duction in the weight of the bones of the wing, and this no doubt
has resulted from their lessened use. The one exceptional case,
namely, in one of the Call-ducks, is in truth no exception, for this
bird was constantly in the habit of flying about; and I have seen
it day after day rise from my grounds, and fly for a long time in
circles of more than a mile in diameter. In this Call-duck there is
not only no decrease, but an actual increase in the weight of the
wing-bones relatively to those of the wild duck; and this probably
15*

346
CHAP. VIII.
DOMESTIC DUCKS.
is consequent on the remarkable lightness and thinness of all the
bones of the skeleton.
Lastly, I weighed the furcula, coracoids, and scapula of a wild
duck and of a common domestic duck, and I found that their
weight, relatively to that of the whole skeleton, was as one hun-
dred in the former to eighty-nine in the latter; this shows that
these bones in the domestic duck have been reduced eleven per
cent. of their due proportional weight. The prominence of the
crest of the sternum, relatively to its length, is also much reduced
in all the domestic breeds. These changes have evidently been
caused by the lessened use of the wings.
It is well known that several birds, belonging to dif-
ferent Orders, and inhabiting oceanic islands, have their
wings greatly reduced in size and are incapable of flight.
I suggested in my Origin of Species' that, as these
birds are not persecuted by any enemies, the reduction
of their wings has probably been caused by gradual dis-
use. Hence, during the earlier stages of the process of
reduction, such birds might be expected to resemble in
the state of their organs of flight our domesticated ducks.
This is the case with the water-hen (Gallinula nesiotis)
of Tristan d'Acunha, which “ can flutter a little, but ob-
viously uses its legs, and not its wings, as a mode of
escape.” Now Mr. Sclater 19 finds in this bird that the
wings, sternum, and coracoids, are all reduced in length,
and the crest of the sternum in depth, in comparison with
the same bones in the European water-hen (G. chloro-
pus). On the other hand, the thigh-bones and pelvis
are increased in length, the former by four lines, rela-
tively to the same bones in the common water-hen.
Hence in the skeleton of this natural species nearly the
same changes have occurred, only carried a little further,
as with our domestic ducks, and in this latter case I
pre-
sume no one will dispute that they have resulted from
the lessened use of the wings and the increased use of
the legs.
19 Proc. Zoolog. Soc.,'1861, p. 261.

CHAP. VIII.
347
DOMESTIC GOOSE. .
THE GOOSE.
This bird deserves some notice, as hardly any other an-
ciently domesticated bird or quadruped has varied so
little. That geese were anciently domesticated we know
from certain verses in Homer; and from these birds hav-
ing been kept (388 B.c.) in the Capitol at Rome as sacred
to Juno, which sacredness implies great antiquity.20 That
the goose has varied in some degree, we may infer from
naturalists not being unanimous with respect to its wild
parent-form; though the difficulty is chiefly due to the
existence of three or four closely allied wild European
species. A large majority of capable judges are con-
vinced that our geese are descended from the wild Grey-
lag goose (A. ferus); the young of which can easily be
tamed, 22 and are domesticated by the Laplanders. This
species, when crossed with the domestic goose, produced
in the Zoological Gardens, as I was assured in 1849, per-
fectly fertile offspring 2 Yarrell 24 has observed that the
lower part of the trachea of the domestic goose is some-
times flattened, and that a ring of white feathers sometimes
surrounds the base of the beak. These characters seem
at first good indications of a cross at some former period
with the white-fronted goose (A. albifrons); but the
white ring is variable in this latter species, and we must
not overlook the law of analogous variation; that is, of
one species assuming some of the characters of allied
species.
As the goose has proved so inflexible in its organization
20 Ceylon,' by Sir J. E. Tennent,
1959, vol. i. p. 485; also J. Crawfurd on
the Relation of Domest. Animals to
Civilisation, read before Brit. Assoc.,
1860. See also Ornamental Poultry,'
by Rev. E. S. Dixon, 1848, p. 132. The
goose figured on the Egyptian monu-
ments seems to have been the Red goose
of Egypt.
21 Macgillivray's 'British Birds,' vol.
iv. p. 593.
22 Mr. A. Strickland (Annals and
Mag. of Nat. Hist.,' 3rd Series, vol. iii.
1859, p. 122) reared some young wild
geese, and found them in habits and in
all characters identical with the domes-
tic goose.
23 See also Hunter's Essays,' edited
by Owen, vol. ii. p. 322.
24 Yarrell's British Birds,' vol. iii. p.
142. He refers to the Laplanders domes.
ticating the goose.

348
CHAP. VIII.
DOMESTIC GOOSE.
under long-continued domestication, the amount of varia-
tion which can be detected is worth giving. It has in-
creased in size and in productiveness; 25 and varies from
white to a dusky colour. Several observers 26 have stated
that the gander is more frequently white than the goose,
and that when old it almost invariably becomes white;
but this is not the case with the parent-form, the A. ferus.
Here, again, the law of analogous variation may have
come into play, as the snow-white male of the Rock-
Goose (Bernicla antarctica) standing on the sea-shore by
his dusky partner is a sight well known to all those wh
have traversed the sounds of Tierra del Fuego and the
Falkland Islands. Some geese have topknots; and the
skull beneath, as before stated, is perforated. A sub-
breed has lately been formed with the feathers reversed
at the back of the head and neck.27 The beak varies a
little in size, and is of a yellower tint than in the wild
species; but its colour and that of the legs are both
slightly variable. This latter fact deserves attention,
because the colour of the legs and beak is highly ser-
viceable in discriminating the several closely allied wild
forms.29 At our Shows two breeds are exhibited; viz.
the Embden and Toulouse; but they differ in nothing ex-
cept colour. Recently a smaller and singular variety
has been imported from Sebastopol, with the scapular
feathers (as I hear from Mr. Tegetmeier, who sent me
specimens) greatly elongated, curled, and even spirally
25 L. Lloyd, Scandinavian Adven-
tures,' 1854, vol. ii. p. 413, says that
the wild goose lays from five to eight
eggs, which is a much fewer number
than that laid by our domestic goose.
26 The Rev. L. Jenyns seems first to
have made this observation in his Bri-
tish Animals.' See also Yarrell, and
Dixon in his Ornamental Poultry' (p.
139), and Gardener's Chronicle,' 1857,
Zoological Soc., Feb. 1860.
28 W. Thompson, 'Natural Hist. of Ire-
land,' 1851, vol. iii. p. 31. The Rev. E. S.
Dixon gave me some information on the
varying colour of the beak and legs.
29 Mr. A. Strickland, in 'Annals and
Mag. of Nat. Hist.,' 3rd series, vol. iii.,
1859, p. 122.
30 Poultry Chronicle,' vol. i., 1854, p.
498; vol. iii. p. 210.
31 The Cottage Gardener,' Sept. 4th,
1860, p. 348.
P. 45.
27 Mr. Bartlett exhibited the head and
neck of a bird thus characterised at the

CHAP. VIII.
DOMESTIC GOOSE.
349
twisted. The margins of these feathers are rendered
plumose by the divergence of the barbs and barbules, so
that they resemble in some degree those on the back of
the black Australian swan. These feathers are likewise
remarkable from the central shaft, which is excessively
thin and transparent, being split into fine filaments, which,
after running for a space free, sometimes coalesce again.
It is a curious fact that these filaments are regularly
clothed on each side with fine down or barbules, precisely
like those on the proper barbs of the feather. This struc-
ture of the feathers is transmitted to half-bred birds.
In Gallus sonneratii the barbs and barbules blend to-
gether, and form thin horny plates of the same nature with
the shaft: in this variety of the goose, the shaft divides
into filaments which acquire barbules, and thus resemble
true barbs.
Although the domestic goose certainly differs some-
what from any known wild species, yet the amount of
variation which it has undergone, as compared with most
domesticated animals, is singularly small. This fact can
be partially accounted for by selection not having come
largely into play. Birds of all kinds which present many
distinct races are valued as pets or ornaments; no one
makes a pet of the goose; the name, indeed, in more lan-
guages than one, is a term of reproach. The goose is
valued for its size and flavour, for the whiteness of its
feathers which adds to their value, and for its prolificness
and tameness. In all these points the goose differs from
the wild parent-form ; and these are the points which
have been selected. Even in ancient times the Roman
.
gourmands valued the liver of the white goose; and
Pierre Belon $2 in 1555 speaks of two varieties, one of
which was larger, more fecund, and of a better colour
than the other; and he expressly states that good mana-
32 L'Hist. de la Nature des Oiseaux,'
par P. Belon, 1555, p. 156. With respect
to the livers of white geese being prefer-
red by the Romans, see Isid. Geoffroy
St. Hilaire, Hist. Nat. Gén.,' tom. iii.
p. 58.

350
CHAP. VIII.
PEACOCK.
gers attended to the colour of their goslings, so that they
might know which to preserve and select for breeding.
THE PEACOCK.
This is another bird which has hardly varied under do-
mestication, except in sometimes being white or piebald.
Mr. Waterhouse carefully compared, as he informs me,
skins of the wild Indian and domestic bird, and they were
identical in every respect, except that the plumage of the
latter was perhaps rather thicker. Whether our birds are
descended from those introduced into Europe in the time
of Alexander, or have been subsequently imported, is
doubtful. They do not breed very freely with us, and
are seldom kept in large numbers,-circumstances which
would greatly interfere with the gradual selection and
formation of new breeds.
There is one strange fact with respect to the peacock,
namely, the occasional appearance in England of the
“japanned” or “ black-shouldered” kind. This form has
lately been named on the high authority of Mr. Sclater
as a distinct species, viz. Pavo nigripennis, which he be-
lieves will hereafter be found wild in some country, but
not in India, where it is certainly unknown. These ja-
panned birds differ conspicuously from the common pea-
cock in the colour of their secondary wing-feathers, sca-
pulars, wing-coverts, and thighs; the females are much
paler, and the young, as I hear from Mr. Bartlett, like-
wise differ. They can be propagated perfectly true.
Although they do not resemble the hybrids which have
been raised between P. cristatus and muticus, neverthe-
less they are in some respects intermediate in character
between these two species; and this fact favours, as Mr.
Sclater believes, the view that they form a distinct and
natural species.
33
33 Mr. Sclater on the black-shouldered peacock of Latham, Proc. Zoolog. Soc.,'
April 24th, 1860.

CHAP. VIII.
351
РЕАСОСК.
On the other hand, Sir R. Heron states 54 that this
breed suddenly appeared within his memory in Lord
Brownlow's large stock of pied, white, and common pea-
cocks. The same thing occurred in Sir J. Trevelyan's
flock composed entirely of the common kind, and in Mr.
Thornton's stock of common and pied peacocks.
It is
remarkable that in these two latter instances the black-
shouldered kind increased,“ to the extinction of the pre-
viously existing breed.” I have also received through
Mr. Sclater a statement from Mr. Hudson Gurney that
he reared many years ago a pair of black-shouldered pea-
cocks from the common kind; and another ornithologist,
Prof. A. Newton, states that, five or six years ago, a
female bird, in all respects similar to the female of the
black-shouldered kind, was produced from a stock of
common peacocks in his possession, which during more
than twenty years had not been crossed with birds of
any
other strain. Here we have five distinct cases of
japanned birds suddenly appearing in flocks of the com-
mon kind kept in England. Better evidence of the first
appearance of a new variety could hardly be desired. If
we reject this evidence, and believe that the japan-
ned peacock is a distinct species, we must suppose in all
these cases that the common breed had at some former
period been crossed with the supposed P. nigripennis,
but had lost every trace of the cross, yet that the birds
occasionally produced offspring which suddenly and com-
pletely reacquired through reversion the characters of P.
nigripennis. I have heard of no other such case in the
animal or vegetable kingdom. To perceive the full im-
probability of such an occurrence, we may suppose that a
breed of dogs had been crossed at some former period
with a wolf, but had lost every trace of the wolf-like
character, yet that the breed gave birth in five instances
in the same country, within no great length of time, to a
34 Proc. Zoolog. Soc.,' April 14th, 1835.

352
CHAP. VIII.
TURKEY.
wolf perfect in every character; and we must further sup-
pose that in two of the cases the newly produced wolves
afterwards spontaneously increased to such an extent as
to lead to the extinction of the parent breed of dogs. So
remarkable a form as the P. nigripennis, when first im-
ported, would have realized a large price; it is therefore
improbable that it should have been silently introduced
and its history subsequently lost. On the whole the evi-
dence seems to me, as it did to Sir R. Heron, to prepon-
derate strongly in favour of the black-shouldered breed
being a variation, induced either by the climate of Eng-
land, or by some unknown cause, such as reversion to a
primordial and extinct condition of the species. On the
view that the black-shouldered peacock is a variety, the
case is the most remarkable ever recorded of the abrupt
appearance of a new form, which so closely resembles a
true species that it has deceived one of the most experi-
enced of living ornithologists.
THE TURKEY.
It seems fairly well established by Mr. Gould, that the
turkey, in accordance with the history of its first intro-
duction, is descended from a wild Mexican species (Me-
leagris Mexicana) which had been already domesticated
by the natives before the discovery of America, and
which differs specifically, as it is generally thought, from
the common wild species of the United States. Some
naturalists, however, think that these two forms should
be ranked only as well-marked geographical races. How-
this may be, the case deserves notice because in the
35 Proc. Zoolog. Soc.,' April 8th,
1856, p. 61. Prof. Baird believes (as
quoted in Tegetmeier's Poultry Book,'
1866, p. 269) that our turkeys are de-
scended from a West Indian species now
extinct. But besides the improbability
of a bird having long ago become extinct
in these large and luxuriant islands, it
appears (as we shall presently see) that
the turkey degenerates in India, and this
fact indicates that it was not aboriginally
an inhabitant of the lowlands of the
tropics.

CHAP. VIII.
353
TURKEY.
United States wild male turkeys sometimes court the
domestic hens, which are descended from the Mexican
form, “and are generally received by them with great
pleasure.” 36 Several accounts bave likewise been pub-
lished of young birds, reared in the United States from
the eggs of the wild species, crossing and commingling
with the common breed. In England, also, this same
species has been kept in several parks; from two of which
the Rev. W. D. Fox procured birds, and they crossed
freely with the common domestic kind, and during many
years afterwards, as he informs me, the turkeys in his
neighbourhood clearly showed traces of their crossed
parentage. We here have an instance of a domestic race
being modified by a cross with a distinct species or wild
race. F. Michaux 37 suspected in 1802 that the common
domestic turkey was not descended from the United
States species alone, but likewise from a southern form,
and he went so far as to believe that English and French
turkeys differed from having different proportions of the
blood of the two parent-forms.
English turkeys are smaller than either wild form.
They have not varied in any great degree; but there are
some breeds which can be distinguished-as Norfolks,
Suffolks, Whites, and Copper-coloured (or Cambridge),
all of which, if precluded from crossing with other breeds,
propagate their kind truly. Of these kinds, the most
distinct is the small, hardy, dull-black Norfolk turkey, of
which the chickens are black, with occasionally white
patches about the head. The other breeds scarcely differ
except in colour, and their chickens are generally mottled
all over with brownish-grey.38 The tuft of hair on the
breast, which is proper to the male alone, occasionally
appears on the breast of the domesticated female.39 The
38 Audubon's Ornithological Bio-
graph.,' vol. i., 1831, pp. 4-13; and
Naturalist's Library,' vol. xiv., Birds,
rica,' 1802, Eng. translat., p. 217.
38. Ornamental Poultry,' by the Rev.
E. S. Dixon, 1848, p. 34.
39 Rev. E. S. Dixon, id., p. 35.
P. 138.
37 F. Michaux, Travels in N. Ame-

354
CHAP. VIII.
TURKEY.
inferior tail-coverts vary in number, and according to a
German superstition the hen lays as many eggs as the
cock has feathers of this kind.40 In Holland there was
formerly, according to Temminck, a beautiful buff-yellow
breed, furnished with an ample white topknot. Mr.
Wilmot has described 41 a white turkey-cock with a crest
formed of " feathers about four inches long, with bare
quills, and a tuft of soft white down growing at the
end." Many of the young
birds whilst
young
inherited
this kind of crest, but afterwards it either fell off or was
pecked out by the other birds. This is an interesting
case, as with care a new breed might probably have been
formed ; and a topknot of this nature would have been to
a certain extent analogous to that borne by the males in
several allied genera, such as Euplocomus, Lophophorus,
and Pavo.
Wild turkeys, believed in every instance to have been
imported from the United States, have been kept in the
parks of Lords Powis, Leicester, Hill, and Derby. The
Rev. W. D. Fox procured birds from the two first-named
parks, and he informs me that they certainly differed a
little from each other in the shape of their bodies and in
the barred plumage on their wings. These birds likewise
differed from Lord Hill's stock. Some of the latter kept
at Oulton by Sir P. Egerton, though precluded from
crossing with common turkeys, occasionally produced
much paler-coloured birds, and one that was almost
white, but not an albino. These half-wild turkeys in thus
slightly differing from each other present an analogous
case with the wild cattle kept in the several British
parks. We must suppose that the differences have re-
sulted from the prevention of free intercrossing between
birds ranging over a wide area, and from the changed
conditions to which they have been exposed in England.
40 Bechstein, Naturgesch. Deutschlands,' B. iii., 1793, s. 309.
41 Gardener's Chronicle,' 1852, p. 699.

CHAP. VIII.
355
GUINEA FOWL---CANARY BIRD.
42
In India the climate has apparently wrought a still greater
change in the turkey, for it is described by Mr. Blyth
as being much degenerated in size, “utterly incapable of
rising on the wing," of a black colour, and “with the
long pendulous appendages over the beak enormously
developed.”
THE GUINEA FOWL.
The domesticated guinea-fowl is now believed by natural-
ists to be descended from the Numida ptilorhynca, which
inhabits very hot, and, in parts, extremely arid districts
in Eastern Africa; consequently it has been exposed in
this country to extremely different conditions of life.
Nevertheless it has hardly varied at all, except in the
plumage being either paler or darker-coloured. It is a
singular fact that this bird varies more in colour in the
West Indies and on the Spanish Main, under a hot though
humid climate, than in Europe. The guinea-fowl has
become thoroughly feral in Jamaica and in St. Domingo,
and has diminished in size; the legs are black, whereas
the legs of the aboriginal African bird are said to be grey.
This small change is worth notice on account of the often-
repeated statement that all feral animals invariably revert
in every character to their original type.
THE CANARY BIRD.
As this bird has been recently domesticated, namely,
within the last 350 years, its variability deserves no-
tice. It has been crossed with nine or ten other species
of Fringillidæ, and some of the hybrids are almost com-
42 E. Blyth, in 'Annals and Mag. of
Nat. Hist.,' 1847, vol. xx. p. 391.
43 Roulin makes this remark in 'Mem.
de divers Savans, l'Acad. des Sciences,
tom. vi., 1835, p. 319. Mr. Hill, of
Spanish Town, in a letter to me, describes
five varieties of the guinea-fowl in Ja-
maica. I have seen singular pale-colour-
ed varieties imported from Barbadoes
and Demerara.
44 For St. Domingo, see M. A. Salle, in
Proc. Soc. Zoolog., ' 1857, p. 236. Mr.
Hill remarks to me, in his letter, on the
colour of the legs of the feral birds in
Jamaica.

356
CHAP. VIIL
CANARY BIRD.
pletely fertile; but we have no evidence that
any
distinct
breed has originated from such crosses. Notwithstand-
ing the modern domestication of the canary, many varie-
ties have been produced; even before the year 1718 a list
of twenty-seven varieties was published in France, 45 and
in 1779 a long schedule of the desired qualities was
printed by the London Canary Society, so that me-
thodical selection has been practised during a consi-
derable period. The greater number of the varieties
differ only in colour and in the markings of their plu-
mage. Some breeds, however, differ in shape, such as
the hooped or bowed canaries, and the Belgian canaries
with their much elongated bodies. Mr. Brent measured
one of the latter and found it eight inches in length,
whilst the wild canary is only five and a quarter inches
long. There are topknotted canaries, and it is a singular
fact, that, if two topknotted birds are matched, the young,
instead of having very fine topknots, are generally bald,
or even have a wound on their heads. It would ap-
pear as if the topknot were due to some morbid con-
dition which is increased to an injurious degree when
two birds in this state are paired. There is a feather-
footed breed, and another with a kind of frill running
down the breast. One other character deserves notice
from being confined to one period of life and from being
strictly inherited at the same period : namely, the wing
and tail feathers in prize canaries being black, “but this
colour is retained only until the first moult; once moult-
ed, the peculiarity ceases.
1948 Canaries differ much in dis-
position and character, and in some small degree in song.
They produce eggs three or four times during the year.
45 Mr. B. P. Brent, The Canary,
British Finches,' &c., pp. 21, 30.
46 Cottage Gardener,' Dec. 11th, 1855,
An account is here given of all
the varieties. For many measurements
of the wild birds, see Mr. E. Vernon Har-
court, id., Dec. 25th, 1855, p. 223.
47 Bechstein, Naturgesch. der Stu-
benvögel,' 1840, s. 243 ; see s. 252, on
the inherited song of Canary-birds.
With respect to their baldness, see also
W. Kidd's Treatise on Song-Birds.'
48 W. Kidd's Treatise on Song-Birds.'
p. 184.
p. 18.

CHAP. VIII.
357
GOLD-FISH.
GOLD-FISH.
BESIDES mammals and birds, few animals belonging to
the other great classes have been domesticated; but to
show that it is an almost universal law that animals,
when removed from their natural conditions of life,
vary, and that races can be formed when selection is
applied, it is necessary to say a few words on gold-fish,
bees, and silk-moths.
Gold-fish (Cyprinus auratus) were introduced into
Europe only two or three centuries ago; but it is be-
lieved that they have been kept in confinement from an
ancient period in China. Mr. Blyth
Mr. Blyth º suspects from the
analogous variation of other fishes that golden-coloured
fish do not occur in a state of nature. These fishes fre-
quently live under the most unnatural conditions, and
their variability in colour, size, and in some important
points of structure is very great. M. Sauvigny has de-
scribed and given coloured drawings of no less than
eighty-nine varieties. Many of the varieties, however,
such as triple tail-fins, &c., ought to be called monstrosi-
ties; but it is difficult to draw any distinct line between
a variation and a monstrosity. As gold-fish are kept for
ornament or curiosity, and as “the Chinese are just the
people to have secluded a chance variety of any kind,
and to have matched and paired from it,” 61 we may feel
nearly confident that selection has been largely practised
in the formation of new breeds. It is however a singular
fact that some of the monstrosities or variations are not
inherited; for Sir R. Heron 52 kept many of these fishes,
and placed all the deformed fishes, namely, those desti-
tute of dorsal fins, and those furnished with a double
anal fin, or triple tail, in a pond by themselves; but
49 The 'Indian Field,' 1858, p. 255.
50 Yarrell's British Fishes,' vol. i. p.
819.
81 Mr. Blyth, in the Indian Field,'
1958, p. 255.
52 Proc. Zoolog. Soc.,' May 25th,
1842.

358
CHAP. VIII.
HIVE-BEES.
they did not produce a greater proportion of deform-
ed offspring than the perfect fishes.”
Passing over an almost infinite diversity of colour, we
meet with the most extraordinary modifications of struc-
ture. Thus, out of about two dozen specimens bought
in London, Mr. Yarrell observed some with the dorsal
fin extending along more than half the length of the
back; others with this fin reduced to only five or six
rays; and one with no dorsal fin. The anal fins are
sometimes double, and the tail is often triple. This lat-
ter deviation of structure seems generally to occur “at
the expense of the whole or part of some other fin;"> 53
but Bory de Saint Vincent 54 saw at Madrid gold-fish
furnished with a dorsal fin and a triple tail. One va-
riety is characterized by a hump on its back near the
head; and the Rev. L. Jenyns has described a most
singular variety, imported from China, almost globular
in form like a Diodon, with “the fleshy part of the tail
as if entirely cut away; the caudal fin being set on a
little behind the dorsal and immediately above the anal.”
In this fish the anal and caudal fins were double; the anal
fin being attached to the body in a vertical line: the eyes
also were enormously large and protuberant.
HIVE-BEES.
BEES have been domesticated from an ancient period ; if
indeed their state can be considered one of domestication,
for they search for their own food, with the exception of
a little generally given to them during the winter. Their
habitation is a hive instead of a hole in a tree. Bees,
however, have been transported into almost every quarter
of the world, so that climate ought to have produced
53 Yarrell's 'British Fishes,' vol. i. p.
319.
64 Dict. Class. d'Hist. Nat.,' tom. v.
p. 211. Dr. Gray has described, in 'An-
nals and Mag. of Nat. Hist.,' 1860, p.
151, a nearly similar variety, but desti-
tute of a dorsal fin.
p. 276.
55 Observations in Nat. Hist.,' 1846,

CHAP. VIII.
359
HIVE-BEES.
whatever direct effect it is capable of producing. It is
frequently asserted that the bees in different parts of
Great Britain differ in size, colour, and temper; and
Godron 6 says that they are generally larger in the south
than in other parts of France; it has also been asserted
that the little brown bees of High Burgundy, when
transported to La Bresse, become large and yellow in
the second generation. But these statements require
confirmation. As far as size is concerned, it is known
that bees produced in very old combs are smaller, owing
to the cells having become smaller from the successive
old cocoons. The best authorities 57
concur that, with
the exception of the Ligurian race or species, presently
to be mentioned, distinct breeds do not exist in Britain
or on the Continent. There is, however, even in the
same stock, some variability in colour. Thus Mr. Wood-
bury states 58 that he has several times seen queen bees of
the common kind annulated with yellow like Ligurian
queens, and the latter dark-coloured like common bees.
He has also observed variations in the colour of the
drones, without any corresponding difference in the
queens or workers of the same hive. The great apiarian
Dzierzon, in answer to my queries on this subject, says
that in Germany bees of some stocks are decidedly dark,
whilst others are remarkable for their yellow colour.
Bees also seem to differ in habits in different districts, for
Dzierzon adds, “If many stocks with their offspring are
more inclined to swarm, whilst others are richer in honey,
so that some bee-keepers even distinguish between swarm-
ing and honey-gathering bees, this is a habit which has
59
56 De l'Espèce,' 1859, p. 459. With
respect to the bees of Burgundy, see M.
Gérard, art. 'Espèce,' in Dict. Univers.
d'Hist. Nat.'
57 See a discussion on this subject, in
answer to a question of mine, in 'Journal
of Horticulture,' 1862, pp. 225-242; also
Mr. Bevan Fox, in ditto, 1862, p. 284.
58 This excellent observer may be im-
plicitly trusted; see Journal of Horti-
culture,' July 14th, 1863, p. 39.
59 Journal of Horticulture,' Sept. 9th,
1862, p. 463; see also Herr Kleine on
same subject (Nov. 11th, p. 643), who
sums up, that, though there is some
variability in colour, no constant or per-
ceptible differences can be detected in
the bees of Germany,

360
CHAP. VIII.
HIVE-BEES.
become second nature, caused by the customary mode of
keeping the bees and the pasturage of the district. For
example; what a difference in this respect one may
perceive to exist between the bees of the Lüneburg heath
and those of this country!"....
......"Removing an old
queen and substituting a young one of the current year
is here an infallible mode of keeping the strongest stock
from swarming and preventing drone-breeding; whilst
the same means if adopted in Hanover would certainly
be of no avail.” I procured a hive full of dead bees
from Jamaica, where they have long been naturalised,
and, on carefully comparing them under the microscope
with my own bees, I could detect not a trace of dif-
ference.
This remarkable uniformity in the hive-bee, wherever
kept, may probably be accounted for by the great diffi-
culty, or rather impossibility, of bringing selection into
play by pairing particular queens and drones, for these
insects unite only during flight. Nor is there any record,
with a single partial exception, of any person having
separated and bred from a hive in which the workers
presented some appreciable difference. In order to form
a new breed, seclusion from other bees would, as we now
know, be indispensable; for since the introduction of the
Ligurian bee into Germany and England, it has been
found that the drones wander at least two miles from
their own hives, and often cross with the queens of the
common bee. The Ligurian bee, although perfectly
fertile when crossed with the common kind, is ranked by
most naturalists as a distinct species, whilst by others it
is ranked as a natural variety: but this form need not
here be noticed, as there is no reason to believe that it is
the product of domestication. The Egyptian and some
other bees are likewise ranked by Dr. Gerstäcker, 1 but
60 Mr. Woodbury has published seve-
ral such accounts in Journal of Horti-
culture,' 1861 and 1862.
61 Annals and Mag. of Nat. Hist.,'
3rd series, vol. xi. p. 389.

CHAP. VIII.
361
SILK-MOTHS.
not by other highly competent judges, as geographical
races; and he grounds his conclusion in chief part on
the fact that in certain districts, as in the Crimea and
Rhodes, the hive-bee varies so much in colour, that the
several geographical races can be closely connected by
intermediate forms.
I have alluded to a single instance of the separation
and preservation of a particular stock of bees. Mr. Lowe 62
procured some bees from a cottager a few miles from
Edinburgh, and perceived that they differed from the com-
mon bee in the hairs on the head and thorax being light-
er coloured and more profuse in quantity. From the date
of the introduction of the Ligurian bee into Great Britain
we may feel sure that these bees had not been crossed
with this form. Mr. Lowe propagated this variety, but
unfortunately did not separate the stock from his other
bees, and after three generations the new character was
almost completely lost. Nevertheless, as he adds, “a
great number of the bees still retain traces, though faint,
of the original colony.” This case shows us what could
probably be effected by careful and long-continued selec-
tion applied exclusively to the workers, for, as we have
seen, queens and drones cannot be selected and paired.
SILK-MOTHS.
THESE insects are in several respects interesting to us,
more especially because they have varied largely at early
periods of life, and the variations have been inherited at
corresponding periods. As the value of the silk-moth
depends entirely on the cocoon, every change in its struc-
ture and qualities has been carefully attended to, and
races differing much in the cocoon, but hardly at all in
the adult state, have been produced. With the races of
82 The Cottage Gardener,' May, 1860, p. 110; and ditto in Journal of Hort..' 1962,
p. 242.
16

362
CHAP. VIII.
SILK-MOTHS.
most other domestic animals, the young resemble each
other closely, whilst the adults differ much.
It would be useless, even if it were possible, to describe
all the many kinds of silk-worms. Several distinct spe-
cies exist in India and China which produce useful silk,
and some of these are capable of freely crossing with the
common silk-moth, as has been recently ascertained in
France. Captain Hutton states that throughout the
world at least six species have been domesticated ; and
he believes that the silk-moths reared in Europe belong
to two or three species. This, however, is not the opi-
nion of several capable judges who have particularly at-
tended to the cultivation of this insect in France; and
hardly accords with some facts presently to be given.
The common silk-moth (Bombyx mori) was brought
to Constantinople in the sixth century, whence it was
carried into Italy, and in 1494 into France. Everything
has been favourable for the variation of this insect. It
is believed to have been domesticated in China as long
ago as 2700 B.C. It has been kept under unnatural and
diversified conditions of life, and has been transported
into many countries. There is reason to believe that the
nature of the food given to the caterpillar influences to a
certain extent the character of the breed.65 Disuse has
apparently aided in checking the development of the
wings. But the most important element in the produc-
tion of the many now existing, much modified races, no
doubt has been the close attention which has long been
applied in many countries to every promising variation.
The care taken in Europe in the selection of the best co-
coons and moths for breeding is notorious, and the pro-
64
69 Transact. Entomolog. Soc.,' 3rd se-
ries, vol. iii. pp. 143-173, and pp. 295-331.
64 Godron, 'De l'Espèce,'1859, tom. i.
p. 460. The antiquity of the silk-worm
in China is given on the authority of
Stanislas Julien,
65 See the remarks of Prof. Westwood,
General Hearsey, and others, at the
meeting of the Entomolog. Soc. of Lon-
don, July, 1861.
66 See, for instance, M. A. de Quatre-
fage's 'Etudes sur les Maladies actuelles
du Ver à Soie,' 1859, p. 101.

CHAP. VIII.
363
THEIR DIFFERENCES.
duction of eggs is followed as a distinct trade in parts of
France. I have made inquiries through Dr. Falconer,
and am assured that in India the natives are equally care-
ful in the process of selection. In China the production
of eggs is confined to certain favourable districts, and the
raisers are precluded by law from producing silk, so that
their whole attention may be necessarily given up to this
one object.º7
The following details on the differences between the several
breeds are taken, when not stated to the contrary, from M. Robinet's
excellent work,68 which bears every sign of care and large ex-
perience. The eggs in the different races vary in colour, in shape
(being round, elliptic, or oval), and in size. The eggs laid in June
in the south of France, and in July in the central provinces, do not
hatch until the following spring; and it is in vain, says M. Robinet,
to expose them to a temperature gradually raised, in order that the
caterpillar may be quickly developed. Yet occasionally, without
any known cause, batches of eggs are produced, which immediately
begin to undergo the proper changes, and are hatched in from
twenty to thirty days. From these and some other analogous facts
it may be concluded that the Trevoltini silkworms of Italy, of
which the caterpillars are hatched in from fifteen to twenty days,
do not necessarily form, as has been maintained, a distinct species.
Although the breeds which live in temperate countries produce
eggs which cannot be immediately hatched by artificial heat, yet
when they are removed to and reared in a hot country they gra-
dually acquire the character of quick development, as in the Trevol-
tini races.
Caterpillars.—These vary greatly in size and colour. The skin is
generally white, sometimes mottled with black or grey, and occasion-
ally quite black. The colour, however, as M. Robinet asserts, is
not constant, even in perfectly pure breeds; except in the race
tigrée, so called from being marked with transverse black stripes.
69
67 My authorities for these statements
will be given in the chapter on Selection.
68 Manuel de l'Educateur de Vers à
Soie,' 1848.
69 Robinet, idem, pp. 12, 318. I may
add that the eggs of N. American silk-
worms taken to the Sandwich Islands
were very irregularly developed ; and
the moths thus raised produced eggs
which were even worse in this respect.
Some were hatched in ten days, and
others not until after the lapse of many
months. No doubt a regular early cha-
racter would ultimately have been ac-
quired. See review in Athenæum,'
1844, p. 329, of J. Jarves' 'Scenes in the
Sandwich Islands,

364
CHAP. VIII.
SILK-MOTHS.
As the general colour of the caterpillar is not correlated with that
of the silk, Tº this character is disregarded by cultivators, and has
not been fixed by selection. Captain Hutton, in the paper before
referred' to, has argued with much force that the dark tiger-
like marks, which so frequently appear during the later moults in
the caterpillars of various breeds, are due to reversion; for the
caterpillars of several allied wild species of Bombyx are marked
and coloured in this manner. He separated some caterpillars with
the tiger-like marks, and in the succeeding spring (pp. 149, 298)
nearly all the caterpillars reared from them were dark-brindled, and
the tints became still darker in the third generation. The moths
reared from these caterpillars 71 also became darker, and resembled
in colouring the wild B. Huttoni. On this view of the tiger like
marks being due to reversion, the persistency with which they are
transmitted is intelligible.
Several years ago Mrs. Whitby took great pains in breeding silk-
worms on a large scale, and she informed me that some of her
caterpillars had dark eyebrows. This is probably the first step in
reversion towards the tiger-like marks, and I was curious to know
whether so trifling a character would be inherited ; at my request
she separated in 1848 twenty of these caterpillars, and having kept
the moths separate, bred from them. Of the many caterpillars thus
reared, every one without exception had eyebrows, some darker
and more decidedly marked than the others, but all had eyebrows
more or less plainly visible.” Black caterpillars occasionally appear
amongst those of the common kind, but in so variable a manner,
that according to M. Robinet the same race will one year exclusively
produce white caterpillars, and the next year many black ones;
nevertheless, I have been informed by M. A. Bossi of Geneva, that,
if these black caterpillars are separately bred from, they reproduce
the same colour ; but the cocoons and moths reared from them do
not present any difference.
The caterpillar in Europe ordinarily moults four times before
passing into the cocoon stage ; but there are races" à trois mues,'
and the Trevoltini race likewise moults only thrice. It might have
been thought that so important a physiological difference would
not have arisen under domestication ; but M. Robinet 72 states that,
on the one hand, ordinary caterpillars occasionally spin their cocoons
after only three moults, and, on the other hand," presque toutes les
races à trois mues, que nous avons expérimentées, ont fait quatre
70 The Art of rearing Silk-worms,'
translated from Count Dandolo, 1825, p.
23.
71 Transact. Ent. Soc.,' ut supra, pp.
153, 308.
72 Robinet, idem, p. 817.

CHAP. VIII.
365
THEIR DIFFERENCES.
mues à la seconde ou à la troisième année, ce qui semble prouver
qu'il a suffi de les placer dans des conditions favorables pour leur
rendre une faculté qu'elles avaient perdue sous des influences moins
favorables."
Cocoons.—The caterpillar in changing into the cocoon loses about
50 per cent. of its weight; but the amount of loss differs in differ-
ent breeds, and this is of importance to the cultivator. The cocoon
in the different races presents characteristic differences; being
large or small;-nearly spherical with no constriction, as in the
Race de Loriol, or cylindrical with either a deep or slight constric-
tion in the middle with the two ends, or with one end alone,
more or less pointed. The silk varies in fineness and quality, and
in being nearly white, of two tints, or yellow. Generally the
colour of the silk is not strictly inherited : but in the chapter on
Selection I shall give a curious account how, in the course of sixty-
five generations, the number of yellow cocoons in one breed has
been reduced in France from one hundred to thirty-five in the
thousand. According to Robinet, the white race, called Sina, by
careful selection during the last seventy-five years, " est arrivée à
un tel état de pureté, qu'on ne voit pas un seul cocon jaune dans
des millions de cocons blancs.” 73 Cocoons are sometimes formed,
as is well known, entirely destitute of silk, which yet produce
moths; unfortunately Mrs. Whitby was prevented by an accident
from ascertaining whether this character would prove hereditary.
Adult stage.--I can find no account of any constant difference in
the moths of the most distinct races. Mrs. Whitby assured me that
there was none in the several kinds bred by her; and I have re-
ceived a similar statement from the eminent naturalist M. de Qua-
trefages. Captain Hutton also says 74 that the moths of all kinds
vary much in colour, but in nearly the same inconstant manner.
Considering how much the cocoons in the several races differ, this
fact is of interest, and may probably be accounted for on the same
principle as the fluctuating variability of colour in the caterpillar,
namely, that there has been no motive for selecting and perpetuat-
ing any particular variation.
The males of the wild Bombycidæ "fly swiftly in the day-time
and evening, but the females are usually very sluggish and inac-
tive." 75 In several moths of this family the females have abortive
wings, but no instance is known of the males being incapable of
flight, for in this case the species could hardly have been perpetu-
73 Robinet, idem, pp. 306-317.
74 Transact. Ent. Soc. ut supra, p.
817.
75 Stephens' Illustrations, Haustel-
lala,' vol.ii. p. 35. See also Capt. Hut-
ton, Transact. Ent. Soc.' idem, p. 152.

366
CHAP. VIII.
SILK-MOTHS.
ated. In the silk-moth both sexes have imperfect, crumpled wings,
and are incapable of flight; but still there is a trace of the charac-
teristic difference in the two sexes; for though, on comparing a
number of males and females, I could detect no difference in the de-
velopment of their wings, yet I was assured by Mrs. Whitby that
the males of the moths bred by her used their wings more than the
females, and could flutter downwards, though never upwards. She
also states that, when the females first emerge from the cocoon,
their wings are less expanded than those of the male. The degree
of imperfection, however, in the wings varies much in different
races and under different circumstances; M. Quatrefages76 says that
he has seen a number of moths with their wings reduced to a third,
fourth, or tenth part of their normal dimensions, and even to mere
short straight stumps : "il me semble qu'il y a là un véritable arrêt
de développement partiel.” On the other hand, he describes the fe-
male moths of the André Jean breed as having "leurs ailes larges
et étalées. Un seul présente quelques courbures irrégulières et des
plis anomaux." As moths and butterflies of all kinds reared from
wild caterpillars under confinement often have crippled wings, the
same cause, whatever it may be, has probably acted on silk-moths,
but the disuse of their wings during so many generations has, it
may be suspected, likewise come into play.
The moths of many breeds fail to glue their eggs to the surface
on which they are laid," but this proceeds, according to Capt. Hut-
ton,78 merely from the glands of the ovipositor being weakened.
As with other long-domesticated animals, the instincts of the silk-
moth have suffered. The caterpillars, when placed on a mulberry-
tree, often commit the strange mistake of devouring the base of the
leaf on which they are feeding, and consequently fall down ; but
they are capable, according to M. Robinet, 79 of again crawling up
the trunk. Even this capacity sometimes fails, for M. Martins 60
placed some caterpillars on a tree, and those which fell were not
able to remount and perished of hunger, they were even inca-
pable of passing from leaf to leaf.
Some of the modifications which the silk-moth has undergone
stand in correlation with each other. Thus the eggs of the moths
which produce white cocoons and of those which produce yellow co-
coons differ slightly in tint. The abdominal feet also of the cater-
pillars which yield white cocoons are always white, whilst those
78 Etudes sur les Maladies du Ver à
Soie,' 1859, pp. 304, 209.
77 Quatrefages, Etudes,' &c., p.
214
78 Transact. Ent. Soc.,' ut supra, p.
151.
79 Manuel de l'Educateur,' &c., p. 26.
80 Godron, 'De l'Espèce,' p. 462.
6

CHAP. VIII.
367
THEIR DIFFERENCES.
which give yellow cocoons are invariably yellow. We have seen
that the caterpillars with dark tiger-like stripes produce moths
which are more darkly shaded than other moths. It seems well
established "2 that in France the caterpillars of the races which pro-
duce white silk, and certain black caterpillars, have resisted, better
than other races, the disease which has recently devastated the silk-
districts. Lastly, the races differ constitutionally, for some do not
succeed so well under a temperate climate as others; and a damp
soil does not equally injure all the races.83
From these various facts we learn that silk-moths, like
the higher animals, vary greatly under long-continued
domestication. We learn also the more important fact
that variations may occur at various periods of life, and
be inherited at corresponding periods. And finally we
see that insects are amenable to the great principle of
Selection.
81 Quatrefages, 'Etudes,' &c., pp. 12,
209, 214,
82 Robinet, Manuel,' &c., p. 308.
83 Robinet, idem, p. 15.

368
CHAP, IX,
PRELIMINARY REMARKS,
CHAPTER IX.
CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.
PRELIMINARY REMARKS ON THE NUMBER AND PARENTAGE
OF CULTIVATED PLANTS — FIRST, STEPS IN CULTIVATION — GEO-
GRAPHICAL DISTRIBUTION OF CULTIVATED PLANTS.
CEREALIA.— DOUBTS ON THE NUMBER OF SPECIES. WHEAT:
VARJETIES 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 INTER-
CROSS. BEANS. -POTATOES : NUMEROUS VARIETIES OF-
DIFFERING LITTLE, EXCEPT IN THE TUBERS - CHARACTERS IN-
HERITED.
I SHALL not enter into so much detail on the variability of
cultivated plants, as in the case of domesticated animals.
The subject is involved in much difficulty. Botanists
have generally neglected cultivated varieties, as beneath
their notice. In several cases the wild prototype is un-
known or doubtfully known; and in other cases it is
hardly possible to distinguish between escaped seedlings
and truly wild plants, so that there is no safe standard
of comparison by which to judge of any supposed amount
of change. Not a few botanists believe that several of
our anciently cultivated plants have become so profoundly
modified that it is not possible now to recognise their
aboriginal parent-forms. Equally perplexing are the

CHAP. IX.
369
ON CULTIVATED PLANTS.
doubts whether some of them are descended from one
species, or from several inextricably commingled by
crossing and variation. Variations often pass into, and
cannot be distinguished from, monstrosities; and mon-
strosities are of little significance for our purpose. Many
varieties are propagated solely by grafts, buds, layers,
bulbs, &c., and frequently it is not known how far
their peculiarities can be transmitted by seminal genera-
tion. Nevertheless some facts of value can be gleaned;
and other facts will hereafter be incidentally given. One
chief object in the two following chapters is to show how
generally almost every character in our cultivated plants
has become variable.
Before entering on details a few general remarks on
the origin of cultivated plants may be introduced. M.
Alph. de Candolle' in an admirable discussion on this
subject, in which he displays a wonderful amount of
knowledge, gives a list of 157 of the most useful cultivat-
ed plants.
Of these he believes that 85 are almost cer-
tainly known in their wild state; but on this head other
competent judges' entertain great doubts. Of 40 of them,
the origin is admitted by M. De Candolle to be doubtful,
either from a certain amount of dissimilarity which they
present when compared with their nearest allies in a wild
state, or from the probability of the latter not being truly
wild plants, but seedlings escaped from culture. Of
the entire 157, 32 alone are ranked by M. De Candolle
as quite unknown in their aboriginal condition. But it
should be observed that he does not include in his list
several plants which present ill-defined characters, namely,
the various forms of pumpkins, millet, sorghum, kidney-
bean, dolichos, capsicum, and indigo. Nor does he in-
clude flowers; and several of the more anciently culti-
1 Géographie Botanique Raisonnée,'
1855, pp. $10 to 991.
2 Review by Mr. Bentham in 'Hort.
Journal,' vol. ix. 1855, p. 133, entitled
Historical Notes on Cultivated Plants,'
by Dr. A. Targioni-Tozzetti. See also
Edinburgh Review,' 1866, p. 510.
16*

370
CHAP. IX.
PRELIMINARY REMARKS,
vated flowers, such as certain roses, the common Imperial
lily, the tuberose, and even the lilac, are said not to be
known in the wild state.
From the relative numbers above given, and from other
arguments of much weight, M. De Candolle concludes
that plants have rarely been so much modified by culture
that they cannot be identified with their wild prototypes.
But on this view, considering that savages probably
would not have chosen rare plants for cultivation, that
useful plants are generally conspicuous, and that they
could not have been the inhabitants of deserts or of re-
mote and recently discovered islands, it appears strange
to me that so many of our cultivated plants should be
still unknown or only doubtfully known in the wild state.
If, on the other hand, many of these plants have been
profoundly modified by culture, the difficulty disappears.
Their extermination during the progress of civilisation
would likewise remove the difficulty; but M. De Candolle
has shown that this probably has seldom occurred. As
soon as a plant became cultivated in any country, the
half-civilised inhabitants would no longer have need
to search the whole surface of the land for it, and thus
lead to its extirpation; and even if this did occur during
a famine, dormant seeds would be left in the ground. In
tropical countries the wild luxuriance of nature, as was
long ago remarked by Humboldt, overpowers the feeble
efforts of man. In anciently civilised temperate coun-
tries, where the whole face of the land has been greatly
changed, it can hardly be doubted that some plants have
been exterminated; nevertheless De Candolle has shown
that all the plants historically known to have been first
cultivated in Europe still exist here in the wild state.
MM. Loiseleur-Deslongchamps and De Candolle have
3 Hist. Notes,' as above, by Targioni-
Tozzetti.
4 Considérations sur les Céréales,'
1842, p. 87. 'Géographle Bot.,' 1855,
p. 930. "Plus on suppose l'agriculture
ancienne et remontant à une époque
d'ignorance, plus il est probable que les
cultivateurs avaient choisi des espèces

CHAP. IX.
371
ON CULTIVATED PLANTS.
remarked that our cultivated plants, more especially the
cereals, must originally have existed in nearly their
present state; for otherwise they would not have been
noticed and valued as objects of food. But these au-
thors apparently have not considered the many accounts
given by travellers of the wretched food collected by
savages. I have read an account of the savages of Aus-
tralia cooking, during a dearth, many vegetables in va-
rious ways, in the hopes of rendering them innocuous
and more nutritious. Dr. Hooker found the half-starved
inhabitants of a village in Sikhim suffering greatly from
having eaten arum-roots, which they had pounded and
left for several days to ferment, so as partially to destroy
their poisonous nature; and he adds that they cooked
and ate many other deleterious plants. Sir Andrew
Smith informs me that in South Africa a large number
of fruits and succulent leaves, and especially roots, are
used in times of scarcity. The natives, indeed, know
the properties of a long catalogue of plants, some having
been found during famines to be eatable, others injurious
to health, or even destructive to life. He met a party of
Baquanas who, having been expelled by the conquering
Zulus, had lived for years on any roots or leaves which
afforded some little nutriment, and distended their stom-
achs, so as to relieve the pangs of hunger. They looked
like walking skeletons, and suffered fearfully from con-
stipation. Sir Andrew Smith also informs me that on
such occasions the natives observe as a guide for them-
selves, what the wild animals, especially baboons and
monkeys, eat.
From innumerable experiments made through dire
necessity by the savages of every land, with the results
handed down by tradition, the nutritious, stimulating,
and medicinal properties of the most unpromising plants
offrant à l'origine même un avantage in-
contestable."
6 Dr. Hooker has given me this in-
formation. See, also, his Himalayan
Journals,' 1854, vol. ii. p. 49.

372
CHAP. IX,
PRELIMINARY REMARKS,
were probably first discovered. It appears, for instance,
at first an inexplicable fact that untutored man, in three
distant quarters of the world, should have discovered
amongst a host of native plants that the leaves of the
tea-plant and mattee, and the berries of the coffee, all
included a stimulating and nutritious essence, now known
to be chemically the same. We can also see that sav-
ages suffering from severe constipation would naturally
observe whether any of the roots which they devoured
acted as aperients. We probably owe our knowledge
of the uses of almost all plants to man having originally
existed in a barbarous state, and having been often com-
pelied by severe want to try as food almost everything
which he could chew and swallow.
From what we know of the habits of savages in
many
quarters of the world, there is no reason to suppose
that
our cereal plants originally existed in their present state
so valuable to man. Let us look to one continent alone,
namely, Africa: Barth® states that the slaves over a
large part of the central region regularly collect the
seeds of a wild grass, the Pennisetum distichum ; in
another district he saw women collecting the seeds of a
Poa by swinging a sort of basket through the rich mea-
dow-land. Near Tete Livingstone observed the natives
collecting the seeds of a wild grass; and farther south, as
Andersson informs me, the natives largely use the seeds
of a grass of about the size of canary-seed, which they
boil in water. They eat also the roots of certain reeds,
and every one has read of the Bushmen prowling about
and digging up with a fire-hardened stake various roots.
Similar facts with respect to the collection of seeds of
wild grasses in other parts of the world could be given."
& Travels in Central Africa,' Eng.
translat., vol. i. pp. 529 and 390; vol. ii.
pp. 29, 265, 270. Livingstone's Tra-
vels,' p. 551.
As in both North and South America.
Mr. Edgeworth ("Journal Proc. Linn.
Soc.,' vol. vi. Bot., 1862, p. 181) states
that in the deserts of the Punjab poor
women sweep up," by a whisk into straw
baskets," the seeds of four genera of

CHAP. IX.
373
ON CULTIVATED PLANTS.
Accustomed as we are to our excellent vegetables and
luscious fruits, we can hardly persuade ourselves that the
stringy roots of the wild carrot and parsnip, or the little
shoots of the wild asparagus, or crabs, sloes, &c., should
ever have been valued; yet, from what we know of the
habits of Australian and South African savages, we need
feel no doubt on this head. The inhabitants of Switzer-
land during the Stone-period largely collected wild crabs,
sloes, bullaces, hips of roses, elderberries, beech-mast,
and other wild berries and fruit. Jemmy Button, a
Fuegian on board the Beagle, remarked to me that the
poor and acid black-currants of Tierra del Fuego were too
sweet for his taste.
The savage inhabitants of each land, having found out
by many and hard trials what plants were useful, or
could be rendered useful by various cooking processes,
would after a time take the first step in cultivation by
planting them near their usual abodes. Livingstone
states that the savage Batokas sometimes left wild fruit-
trees standing in their gardens, and occasionally even
planted them, “a practice seen nowhere else amongst
the natives.” But Du Chaillu saw a palm and some
other wild fruit-trees which had been planted; and these
trees were considered private property. The next step
in cultivation, and this would require but little fore-
thought, would be to sow the seeds of useful plants; and
as the soil near the hovels of the natives 10 would often be
in some degree manured, improved varieties would sooner
or later arise. Or a wild and unusually good variety of
a native plant might attract the attention of some wise
grasses, namely, of Agrostis, Panicum,
Cenchrus, and Pennisetum, as well as
the seeds of four other genera belonging
to distinct families.
8 Prof. O. Heer, Die Pflanzen der
Pfahlbauten, 1865, aus dem Neujahr.
Naturforsc. Gesellschaft,' 1866; and Dr.
H. Christ, in Rütimeyer's Die Fauna
der Pfahlbauten,' 1861, s. 226.
9 Travels,' p. 535. Du Chaillu, 'Ad-
ventures in Equatorial Africa,' 1861, p.
445.
10 In Tierra del Fuego the spot where
wigwams had formerly stood could be dis-
tinguished at a great distance by the
bright green tint of the native vegeta-
tion.

374
CHAP. IX.
PRELIMINARY REMARKS,
old savage; and he would transplant it, or sow its seed.
That superior varieties of wild fruit-trees occasionally are
found is certain, as in the case of the American species of
hawthorns, plums, cherries, grapes, and hickories, specified
by Professor Asa Gray.1 Downing also refers to certain
wild varieties of the hickory, as being “of much larger
size and finer flavour than the common species.” I have
referred to American fruit-trees, because we are not
in this case troubled with doubts whether or not the
varieties are seedlings which have escaped from cultiva-
tion. Transplanting any superior variety, or sowing its
seeds, hardly implies more forethought than might be
expected at an early and rude period of civilisation.
Even the Australian barbarians “have a law that no plant
bearing seeds is to be dug up after it has flowered ;” and
Sir G. Grey 12 never saw this law, evidently framed for
the preservation of the plant, violated. We see the same
spirit in the superstitious belief of the Fuegians, that kill-
ing water-fowl whilst very young will be followed by
"much rain, snow, blow much.” 13 I
may add, as show-
ing forethought in the lowest barbarians, that the Fuegians
when they find a stranded whale bury large portions in
the sand, and during the often-recurrent famines travel
from great distances for the remnants of the half-putrid
mass.
It has often been remarked 14 that we do not owe a
single useful plant to Australia or the Cape of Good
Hope,-countries abounding to an unparalleled degree
with endemic species,—or to New Zealand, or to America
south of the Plata; and, according to some authors, not
to America northward of Mexico. I do not believe that
any edible or valuable plant, except the canary-grass, has
11 American Acad. of Arts and Scien-
ces,' April 10th, 1860, p. 413. Downing,
The Fruits of America,' 1845, p. 261.
12 Journals of Expeditions in Aus-
tralia,' 1841, vol. ii. p. 292.
13 Darwin's Journal of Researches,
1845, p. 215.
14 De Candolle has tabulated the facts
in the most interesting manner in his
Géographie Bot.,' p. 986.

CHAP. IX.
375
ON CULTIVATED PLANTS.
been derived from an oceanic or uninhabited island. If
nearly all our useful plants, natives of Europe, Asia, and
South America, had originally existed in their present
condition, the complete absence of similarly useful plants
in the great countries just named would indeed be a sur-
prising fact. But if these plants have been so greatly
modified and improved by culture as no longer closely to
resemble any natural species, we can understand why the
above-named countries have given us no useful plants, for
they were either inhabited by men who did not cultivate
the ground at all, as in Australia and the Cape of Good
Hope, or who cultivated it very imperfectly, as in some
parts of America. These countries do yield plants which
are useful to savage man; and Dr. Hooker 15 enumerates
no less than 107 such species in Australia alone; but
these plants have not been improved, and consequently
cannot compete with those which have been cultivated
and improved during thousands of years in the civilised
world.
The case of New Zealand, to which fine island we as
yet owe no widely cultivated plant, may seem opposed to
this view; for, when first discovered, the natives cultiva-
ted several plants; but all inquirers believe, in accord-
ance with the traditions of the natives, that the early
Polynesian colonists brought with them seeds and roots,
as well as the dog, which had all been wisely preserved
during their long voyage. The Polynesians are so fre-
quently lost on the ocean, that this degree of prudence
would occur to any wandering party: hence the early
colonists of New Zealand, like the later European colo-
nists, would not have had any strong inducement to cul-
tivate the aboriginal plants. According to De Candolle
we owe thirty-three useful plants to Mexico, Peru, and
Chile; nor is this surprising when we remember the civil-
ized state of the inhabitants, as shown by the fact of their
15 'Flora of Australia,' Introduction p. ex.

376
CHAP. IX.
CEREAL PLANTS.
having practised artificial irrigation and made tunnels
through hard rocks without the use of iron or gunpow-
der, and who, as we shall see in a future chapter, fully
recognised, as far as animals were concerned, and there-
fore probably in the case of plants, the important princi-
ple of selection. We owe some plants to Brazil; and the
early voyagers, namely Vespucius and Cabral, describe
the country as thickly peopled and cultivated. In North
America the natives cultivated maize, pumpkins, gourds,
beans, and
peas, "all different from ours," and tobacco :
and we are hardly justified in assuming that none of our
present plants are descended from these North American
forms. Had North America been civilized for as long a
period, and as thickly peopled, as Asia or Europe, it is
probable that the native vines, walnuts, mulberries, crabs,
and plums, would have given rise, after a long course of
cultivation, to a multitude of varieties, some extremely
different from their parent-stocks; and escaped seedlings
would have caused in the New, as in the Old World,
much perplexity with respect to their specific distinctness
and parentage."
Cerealia.-I will now enter on details. The cereals cultivated in
Europe consist of four genera-wheat, rye, barley, and oats. Of
wheat the best modern authorities 18 make four or five, or even seven
distinct species; of rye, one; of barley, three; and of oats, two,
three, or four species. So that altogether our cereals are ranked by
different authors under from ten to fifteen distinct species. These
16 For Canada, see J. Cartier's Voyage
in 1534; for Florida, see Narvaez and
Ferdinand de Soto's Voyages. As I have
consulted these and other old Voyages
in more than one general collection of
Voyages, I do not give precise references
to the pages. See also, for several refer"
ences, Asa Gray, in the 'American Jour-
nal of Science, vol. xxiv., Nov. 1857, p.
441. For the traditions of the natives of
New Zealand, see Crawfurd's 'Grammar
and Dict. of the Malay Language,' 1552,
17 See, for example, Mr. Hewett C.
Watson's remarks on our wild plums and
cherries and crabs: Cybele Britannica,'
vol. i. pp. 330, 334, &c. Van Mons (in
his 'Arbres Fruitiers,'1835, tom. i. p. 444)
declares that he has found the types of
all our cultivated varieties in wild seed-
lîngs, but then he looks on these seed-
lings as so many aboriginal stocks.
18 See A. De Candolle, 'Géograph.
Bot.,' 1855, p. 928 et seq. Godron, 'De
l'Espèce,' 1859, tom, ii. p. 70; and Metz-
ger, Die Getreidearten,' &c., 1841.
p. cclx,

Chap. IX.
377
WHEAT.
have given rise to a multitude of varieties. It is a remarkable fact
that botanists are not universally agreed on the aboriginal parent-
form of any one cereal plant. For instance, a high authority writes
in 1855,19 “We ourselves have no hesitation in stating our convic-
tion, as the result of all the most reliable evidence, that none of
these Cerealia exist, or have existed, truly wild in their present state,
but that all are cultivated varieties of species now growing in great
abundance in S. Europe or W. Asia.” On the other hand, Alph. De
Candolle 20 has adduced abundantevidence that common wheat (Tri-
ticum vulgare) has been found wild in various parts of Asia, where
it is not likely to have escaped from cultivation; and there is force
in M. Godron's remark, that, supposing these plants to be escaped
seedlings, 21 if they have propagated themselves in a wild state for
several generations, their continued resemblance to cultivated wheat
renders it probable that the latter has retained its aboriginal cha-
racter. M. de Candolle insists strongly on the frequent occurrence
in the Austrian dominions of rye and of one kind of oats in an ap-
parently wild condition. With the exception of these two cases,
which however are rather doubtful, and with the exception of two
forms of wheat and one of barley, which he believes to have been
found truly wild, M. de Candolle does not seem fully satisfied with
the other reported discoveries of the parent-forms of our other ce-
reals. With respect to oats, according to Mr. Buckman,22 the wild
English Avena fatua can be converted by a few years of careful cul-
tivation and selection into forms almost identical with two very dis-
tinct cultivated races. The whole subject of the origin and specific
distinctness of the various cereal plants is a most difficult one; but
we shall perhaps be able to judge a little better after considering
the amount of variation which wheat has undergone.
Metzger describes seven species of wheat, Godron refers to five,
and De Candolle to only four. It is not improbable that, besides the
kinds known in Europe, other strongly characterised forms exist in
19 Mr. Bentham, in his review, entitled
Hist. Notes on cultivated Plants,' by
Dr. A. Targioni-Tozzetti, in 'Journal of
Hort. Soc.,' vol. ix. (1855), p. 133.
20 'Géograph. Bot.,'p. 928. The whole
subject is discussed with admirable full-
ness and knowledge.
21 Godron, 'De l'Espèce,' tom. ii. p. 72.
A few years ago the excellent, though
misinterpreted, observations of M. Fabre
led many persons to believe that wheat
was a modified descendant of Ægilops ;
but M. Godron (tom. i. p. 165) has shown
by careful experiments that the first step
in the series, viz. Ægilops triticoides,
is a hybrid between wheat and Æ. ovata.
The frequency with which these hybrids
spontaneously arise, and the gradual
manner in which the Æ. triticoides, be-
comes converted into true wheat, alone
leave any doubt on the subject.
22 Report to British Association for
1857, p. 207.

378
CHAP. IX.
CEREAL PLANTS.
the more distant parts of the world; for Loiseleur-Deslongchamps 23
speaks of three new species or varieties, sent to Europe in 1822 from
Chinese Mongolia, which he considers as being there indigenous.
Moorcroft 24 also speaks of Hasora wheat in Ladakh as very peculiar.
If those botanists are right who believe that at least seven species
of wheat originally existed, then the amount of variation in any im-
portant character which wheat has undergone under cultivation has
been slight; but if only four or a lesser number of species originally
existed, then it is evident that varieties so strongly marked have
arisen, that they have been considered by capable judges as specifi-
cally distinct. But the impossibility of deciding which forms ought
to be ranked as species and which as varieties, makes it useless to
specify in detail the differences between the various kinds of wheat.
Speaking generally, the organs of vegetation differ little ; 25 but some
kinds grow close and upright, whilst others spread and trail along
the ground. The straw differs in being more or less hollow, and in
quality. The ears 28 differ in colour and in shape, being quadrangu-
lar, compressed, or nearly cylindrical; and the florets differ in their
approximation to each other, in their pubescence, and in being more
or less elongated. The presence or absence of barbs is a conspicuous
difference, and in certain Gramineæ serves even as a generic cha-
racter; 27 although, as remarked by Godron,28 the presence of barbs
is variable in certain wild grasses, and especially in those, such as
Bromus secalinus and Lolium temulentum, which habitually grow
mingled with our cereal crops, and which have thus unintentionally
been exposed to culture. The grains differ in size, weight, and col-
our; in being more or less downy at one end, in being smooth or
wrinkled, in being either nearly globular, oval, or elongated ; and
finally in internal texture, being tender or hard, or even almost
horny, and in the proportion of gluten which they contain.
Nearly all the races or species of wheat vary, as Godron 29 has re-
marked, in an exactly parallel manner,—in the seed being downy
or glabrous, and in colour,—and in the florets being barbed or not
barbed, &c. Those who believe that all the kinds are descended
from a single wild species may account for this parallel variation by
the inheritance of a similar constitution, and a consequent tendency
to vary in the same manner; and those who believe in the general
theory of descent with modification may extend this view to the
several species of wheat, if such ever existed in a state of nature.
23 Considérations sur les Céréales,
1842-43, p. 29.
24 Travels in the Himalayan Pro-
vinces,' &c., 1841, vol. i. p. 224.
25 Col. J. Le Couteur on the Varieties
of Wheat,' pp. 23, 79.
26 Loiseleur-Deslongchamps, Consid.
sur les Céréales,' p. 11.
27 See an excellent review in Hooker's
Journ. of Botany,' vol. viii. p. 82, note.
28 De l'Espèce,' tom. ii. p. 73.
29 Idem, tom. ii. p. 75.

CHAP. IX.
379
WHEAT.
Although few of the varieties of wheat present any conspicuous
difference, their number is great, Dalbret cultivated during thirty
years from 150 to 160 kinds, and excepting in the quality of the
grain they all kept true: Colonel Le Couteur possessed upwards of
150, and Philippar 322 varieties.30 As wheat is an annual, we thus
see how strictly many trifling differences in character are inherited
through many generations. Colonel Le Couteur insists strongly on
this same fact: in his persevering and successful attempts to raise
new varieties by selection, he began by choosing the best ears, but
soon found that the grains in the same ear differed so that he was
compelled to select them separately; and each grain generally trans-
mitted its own character. The great amount of variability in the
plants of the same variety is another interesting point, which would
never have been detected except by an eye long practised to the
work; thus Colonel Le Couteur relates 31 that in a field of his own
wheat, which he considered at least as pure as that of any of his
neighbours, Professor La Gasca found twenty-three sorts; and Pro-
fessor Henslow has observed similar facts. Besides such individual
variations, forms sufficiently well marked to be valued and to become
widely cultivated sometimes suddenly appear: thus Mr. Sheriff has
had the good fortune to raise in his lifetime seven new varieties,
which are now extensively grown in many parts of Britain.
As in the case of many other plants, some varieties, both old and
new, are far more constant in character than others. Colonel Le
Couteur was forced to reject some of his new sub-varieties, which he
suspected had been produced from a cross, as incorrigibly sportive.
With respect to the tendency to vary, Metzger 33 gives from his own
experience some interesting facts: he describes three Spanish sub-
varieties, more especially one known to be constant in Spain, which
in Germany assumed their proper character only during hot sum-
mers; another variety kept true only in good land, but after having
been cultivated for twenty-five years became more constant. He
mentions two other sub-varieties which were at first inconstant, but
subsequently became, apparently without any selection, accustomed
to their new homes, and retained their proper character. These
facts show what small changes in the conditions of life cause varia-
30 For Dalbret and Philippar, see
Loiseleur-Deslongchamps, Consid. sur
les Céréales,' pp. 45,70. Le Couteur on
Wheat, p. 6.
31 Varieties of Wheat,' Introduction,
p. vi. Marshall, in his 'Rural Economy
of Yorkshire,' vol. ii. p. 9, remarks that
"in every field of corn there is as much
variety as in a herd of cattle."
32 Gardener's Chron. and Agricult.
Gazette,' 1862, p. 963.
33 Getreidearten,' 1811, s. 66, 91, 92,
116, 117.

380
CHAP. IX.
CEREAL PLANTS.
bility, and they further show that a variety may become habituated
to new conditions. One is at first inclined to conclude with Loise.
leur-Deslongchamps, that wheat cultivated in the same country is
exposed to remarkably uniform conditions ; but manures differ, seed
is taken from one soil to another, and what is far more important
the plants are exposed as little as possible to struggle with other
plants, and are thus enabled to exist under diversified conditions.
In a state of nature each plant is confined to that particular station
and kind of nutriment which it can seize from the other plants by
which it is surrounded.
Wheat quickly assumes new habits of life. The summer and
winter kinds were classed by Linnæus as distinct species; but M.
Monnier 34 has proved that the difference between them is only tem-
porary. He sowed winter-wheat in spring, and out of one hundred
plants four alone produced ripe seeds; these were sown and resown,
and in three years plants were reared which ripened all their seed.
Conversely, nearly all the plants raised from summer-wheat, which
was sown in autumn, perished from frost; but a few were saved
and produced seed, and in three years this summer-variety was con-
verted into a winter-variety. Hence it is not surprising that wheat
soon becomes to a certain extent acclimatised, and that seed brought
from distant countries and sown in Europe vegetates at first, or even
for a considerable period,35 differently from our European varieties.
In Canada the first settlers, according to Kalm, 36 found their winters
too severe for winter-wheat brought from France, and their summers
often too short for summer-wheat ; and until they procured summer-
wheat from the northern parts of Europe, which succeeded well,
they thought that their country was useless for corn crops. It is
notorious that the proportion of gluten differs much under different
climates. The weight of the grain is also quickly affected by cli-
mate: Loiseleur-Deslongchamps 37 sowed near Paris 54 varieties,
obtained from the South of France and from the Black Sea, and 52
of these yielded seed from 10 to 40 per cent. heavier than the parent-
seed. He then sent these heavier grains back to the South of
France, but there they immediately yielded lighter seed.
All those who have closely attended to the subject insist on the
close adaptation of numerous varieties of wheat to various soils and
climates even within the same country ; thus Colonel Le Couteur 38
34 Quoted by Godron, 'De l'Espèce,'
vol. ii. p. 74. So it is, according to
Metzger ("Getreidearten,' s. 18), with
summer and winter barley.
36 Loiseleur-Deslongchamps, Céré-
ales,' part ii, p. 224. Le Couteur, p. 70.
Many others accounts could be added.
36 Travels in North America,' 1753-
1761, Eng. translat., vol. iii. p. 165.
97 Céréales,' part. ii, pp. 179-183.
38 On the Varieties of Wheat,' Intro-
duct., p. vii. See Marshall,
See Marshall, Rural

CHAP. IX.
381
WHEAT.
says, " It is the suitableness of each sort to each soil that will enable
the farmer to pay his rent by sowing one variety, where he would
be unable to do so by attempting to grow another of a seemingly
better sort." This may be in part due to each kind becoming habi-
tuated to its conditions of life, as Metzger has shown certainly oc-
curs, but it is probably, in main part due to innate differences be-
tween the several varieties.
Much has been written on the deterioration of wheat; that the qua-
lity of the flour, size of grain, time of flowering, and hardiness may
be modified by climate and soil, seems nearly certain; but that the
whole body of any one sub-variety ever becomes changed into an-
other and distinct sub-variety, there is no reason to believe. What
apparently does take place, according Le Couteur, 39 is, that some
one sub-variety out of the many which may always be detected in
the same field is more prolific than the others, and gradually sup-
plants the variety which was first sown.
With respect to the natural crossing of distinct varieties the evi-
dence is conflicting, but preponderates against its frequent occur-
ence. Many authors maintain that impregnation takes place in the
closed flower, but I am sure from my own observations that this is
not the case, at least with those varieties to which I have attended.
But as I shall have to discuss this subject in another work, it may
be here passed over.
In conclusion, all authors admit that numerous varie-
ties of wheat have arisen ; but their differences are unim-
portant, unless, indeed, some of the so-called species are
ranked as varieties. Those who believe that from four to
seven wild species of Triticum originally existed in nearly
the same condition as at present, rest their belief chiefly
on the great antiquity of the several forms. It is an
important fact, which we have recently learnt from the
admirable researches of Heer, 1 that the inhabitants of
Switzerland, even so early as the Neolithic period, culti-
Econ. of Yorkshire,' vol. ii. p. 9. With
respect to similar cases of adaptation in
the varieties of oats, see some interesting
papers in the
Gardener's Chron. and
Agricult. Gazette,' 1850, pp. 204, 219.
99 On the Varieties of Wheat,' p. 59.
Mr. Sheriff, and a higher authority can-
not be given ("Gard. Chron, and Agri-
cult. Gazette,' 1862, p. 963), says, "I
have never seen grain which has either
been improved or degenerated by cul-
tivation, so as to convey the change to
the succeeding crop."
40 Alph. De Candolle, 'Géograph.
Bot.,' p. 930.
41 'Planzen der Pfahlbauten,' 1866.

382
CHAP. IX.
CEREAL PLANTS.
vated no less than ten cereal plants, namely, five kinds of
wheat, of which at least four are commonly looked at as
distinct species, three kinds of barley, a panicum, and a
setaria. If it could be shown that at the earliest dawn
of agriculture five kinds of wheat and three of barley had
been cultivated, we should of course be compelled to look
at these forms as distinct species. But, as Heer has re-
marked, agriculture even at the period of the lake-habita-
tions had already made considerable progress; for, be-
sides the ten cereals, peas, poppies, flax, and apparently
apples, were cultivated. It may also be inferred, from
one variety of wheat being the so-called Egyptian, and
from what is known of the native country of the panicum
and setaria, as well as from the nature of the weeds which
then grew mingled with the crops, that the lake-inhabi-
tants either still kept up commercial intercourse with
some southern people or had originally proceeded as co-
lonists from the South.
Loiseleur-Deslongchamps 49 has argued that, if our ce-
real plants had been greatly modified by cultivation, the
weeds which habitually grow mingled with them would
have been equally modified. But this argument shows
how completely the principle of selection has been over-
looked. That such weeds have not varied, or at least do
not vary now in any extreme degree, is the opinion of
Mr. H. C. Watson and Professor Asa Gray, as they inform
me; but who will pretend to say that they do not vary as
much as the individual plants of the same sub-variety of
wheat ? We have already seen that pure varieties of
wheat, cultivated in the same field, offer many slight va-
riations, which can be selected and separately propaga-
ted; and that occasionally more strongly pronounced va-
riations appear, which, as Mr. Sheriff has proved, are weil
worthy of extensive cultivation. Not until equal atten-
tion be paid to the variability and selection of weeds, can
42 Les Céréales,' p. 94.

CHAP. IX.
383
WHEAT.
43
the argument from their constancy under unintentional
culture be of any value. In accordance with the princi-
ples of selection we can understand how it is that in the
several cultivated varieties of wheat the organs of vege-
tation differ so little ; for if a plant with peculiar leaves
appeared, it would be neglected unless the grains of corn
were at the same time superior in quality or size. The
selection of seed-corn was strongly recommended in
ancient times by Columella and Celsus; and as Virgil
says,
"I've seen the largest seeds, tho'view'd with care,
Degenerate, unless th' industrious hand
Did yearly cull the largest."
But whether in ancient times selection was methodically
pursued we may well doubt, when we hear how labori-
ous the work was found by Le Couteur. Although the
principle of selection is so important, yet the little which
man has effected, by incessant efforts 44 during thousands
of
years, in rendering the plants more productive or the
grains more nutritious than they were in the time of the
old Egyptians, would seem to speak strongly against its
efficacy. But we must not forget that at each successive
period the state of agriculture and the quantity of manure
supplied to the land will have determined the maximum
degree of productiveness; for it would be impossible to
cultivate a highly productive variety, unless the land con-
tained a sufficient supply of the necessary chemical ele-
ments.
We now know that man was sufficiently civilized to
cultivate the ground at an immensely remote period; so
that wheat might have been improved long ago up
that standard of excellence which was possible under the
then existing state of agriculture. One small class of
facts supports this view of the slow and gradual improve-
ments of our cereals. In the most ancient lake-habita-
to
43 Quoted by Le Couteur, p. 16.
44 A, De Candolle, Géograph. Bot.,' p. 982.

384
CHAP. IX.
CEREAL PLANTS.
45
tions of Switzerland, when men employed only flint-tools,
the most extensively cultivated wheat was a peculiar kind,
with remarkably small ears and grains. " Whilst the
grains of the modern forms are in section from seven to
eight millimètres in length, the larger grains from the
lake-habitations are six, seldom seven, and the smaller
ones only four. The ear is thus much narrower, and the
spikelets stand out more horizontally, than in our present
forms." So again with barley, the most ancient and
most extensively cultivated kind had small ears, and the
grains were “smaller, shorter, and nearer to each other,
than in that now grown; without the husk they were 2}
lines long, and scarcely 1} broad, whilst those now grown
have a length of three lines, and almost the same in
breadth." 46 These small-grained varieties of wheat and
barley are believed by Heer to be the parent-forms of
certain existing allied varieties, which have supplanted
their early progenitors.
Heer gives an interesting account of the first appear-
ance and final disappearance of the several plants which
were cultivated in greater or less abundance in Switzer-
land during former successive periods, and which gene-
rally differed more or less from our existing varieties.
The peculiar small-eared and small-grained wheat, al-
ready alluded to, was the commonest kind during the
Stone period; it lasted down to the Helvetico-Roman
age, and then became extinct. A second kind was rare
at first, but afterwards became more frequent. A third,
the Egyptian wheat (T. turgidum), does not agree exact-
ly with any existing variety, and was rare during the
Stone period. A fourth kind (T. dicoccum) differs from
all known varieties of this form. • A fifth kind (T. mono-
coccum) is known to have existed during the Stone pe-
45 O. Heer, Die Pflanzen der Pfahl-
bauten,' 1866. The following passage is
quoted from Dr. Christ, in 'Die Fauna
der Pfahlbauten von Dr. Rütimeyer,'
1861, s. 225.
46 Heer, as quoted by Carl Vogt, 'Lec-
tures on Man,' Eng. translat., p. 355.

CHAP. IX,
385
MAIZE.
riod only by the presence of a single ear. A sixth kind,
the common T. spelta, was not introduced into Switzer-
land until the Bronze age. Of barley, besides the short-
eared and small-grained kind, two others were cultiva-
ted, one of which was very scarce, and resembled our
present common H. distichum. During the Bronze age
rye and oats were introduced; the oat-grains being some-
what smaller than those produced by our existing varie-
ties. The poppy was largely cultivated during the Stone
period, probably for its oil; but the variety which then
existed is not now known. A peculiar pea with small
seeds lasted from the Stone to the Bronze age, and then be-
came extinct; whilst a peculiar bean, likewise having
small seeds, came in at the Bronze period and lasted to
the time of the Romans. These details sound like the
description given by a paleontologist of the mutations in
form, of the first appearance, the increasing rarity, and
final extinction of fossil species, embedded in the succes-
sive stages of a geological formation.
Finally, every one must judge for himself whether it is
more probable that the several forms of wheat, barley,
rye, and oats are descended from between ten and fifteen
species, most of which are now either unknown or extinct,
or whether they are descended froin between four and
eight species, which may have either closely resembled
our present cultivated forms, or have been so widely dif-
ferent as to escape identification. In this latter case, we
must conclude that man cultivated the cereals at an
enormously remote period, and that he formerly practised
some degree of selection, which in itself is not improbable..
We may, perhaps, further believe that, when wheat was
first cultivated, the ears and grains increased quickly in
size, in the same manner as the roots of the wild carrot
and parsnip are known to increase quickly in bulk under
cultivation.
Maize : Zea Mays. Botanists are nearly unanimous that all the
17

386
Chap. IX.
CEREAL PLANTS.
cultivated kinds belong to the same species. It is undoubtedly 47 of
American origin, and was grown by the aborigines throughout the
continent from New England to Chili. Its cultivation must have
been extremely ancient, for Tschudi 18 describes two kinds, now ex-
tinct or not known in Peru, which were taken from tombs apparent-
ly prior to the dynasty of the Incas. But there is even stronger evi-
dence of antiquity, for I found on the coast of Peru 19 heads of maize,
together with eighteen species of recent sea-shell, embedded in a
beach which had been upraised at least 85 feet above the level of
the sea. In accordance with this ancient cultivation, numerous
American varieties have arisen. The aboriginal form has not as yet
been discovered in the wild state. A peculiar kind,50 in which the
grains, instead of being naked, are concealed by husks as much as
eleven lines in length, has been stated on insufficient evidence to
grow wild in Brazil. It is almost certain that the aboriginal form
would have had its grains thus protected ; 51 but the seeds of the
Brazilian variety produce, as I hear from Professor Asa Gray, and
as is stated in two published accounts, either common or husked
maize; and it is not credible that a wild species, when first cultiva-
ted, should vary so quickly and in so great a degree.
Maize has varied in an extraordinary and conspicuous manner.
Metzger,52 who paid particular attention to the cultivation of this
plant, makes twelve races (unter-art) with numerous sub-varieties;
of the latter some are tolerably constant, others quite inconstant.
The different races vary in height from 15-18 feet to only 16-18
inches, as in a dwarf variety described by Bonafous. The whole
ear is variable in shape, being long and narrow, or short and thick,
or branched. The ear in one variety is more than four times as
long as in a dwarf kind. The seeds are arranged in the ear in
from six to even twenty rows, or are placed irregularly. The seeds
47 See Alph. De Candolle's long dis-
cussion in his 'Géograph. Bot.,' p. 942.
With respect to New England, see Silli-
man's 'American Journal,' vol. xliv. p.
99.
48 "Travels in Peru,' Eng. translat.,
p. 177.
49 Geolog. Observ. on S. America,
1846, p. 49.
50 This maize is figured in Bonafous'
magnificent work, Hist. Nat. du Mais,'
1836, Pl. v. bis, and in the 'Journal of
Hort. Soc.,' vol i., 1846, p. 115, where
an account is given of the result of sow-
ing the seed. A young Guarany Indian,
on seeing this kind of maize, told Au-
guste St. Hilaire (see De Candolle, 'Géo-
graph. Bot.,' p. 951) that it grew wild in
the humid forests of his native land. Mr.
Teschemacher, in Proc. ton Soe.
Nat. Hist,' Oct. 19th, 1842, gives an ac-
count of sowing the seed.
51 Moquin-Tandon, 'Eléments de Té-
ratologie,' 1811, p. 126.
52 Die Getreidearten,' 1941, s. 208.
I have modified a few of Metzger's state-
ments in accordance with those made by
Bonafous in his great work, Hist. Nat.
du Maïs,' 1836.

CHAP. IX.
387
MAIZE.
53
are coloured—white, pale-yellow, orange, red, violet, or elegantly
streaked with black; and in the same ear there are sometimes
seeds of two colours. In a small collection I found that a single
grain of one variety nearly equalled in weight seven grains of
another variety. The shape of the seed varies greatly, being very
flat, or nearly globular, or oval; broader than long, or longer than
broad; without any point, or produced into a sharp tooth, and this
tooth is sometimes recurved. One variety (the rugosa of Bonafous)
has its seeds curiously wrinkled, giving to the whole ear a singular
appearance. Another variety (the cymosa of Bon.) carries its ears
so crowded together that it is called maïs à bouquet. The seeds of
some varieties contain much glucose instead of starch. Male
flowers sometimes appear amongst the female flowers, and Mr. J.
Scott has lately observed the rarer case of female flowers on a true
male panicle, and likewise hermaphrodite flowers. 54 Azara de-
scribes 65 a variety in Paraguay the grains of which are very tender,
and he states that several varieties are fitted for being cooked in
various ways. The varieties also differ greatly in precocity, and
have different powers of resisting dryness and the action of violent
wind.58 Some of the foregoing differences would certainly be con-
sidered of specific value with plants in a state of nature.
Le Comte Ré states that the grains of all the varieties which he
cultivated ultimately assumed a yellow colour. But Bonafous 57
found that most of those which he sowed for ten consecutive years
kept true to their proper tints; and he adds that in the valleys of
the Pyrenees and on the plains of Piedmont a white maize has been
cultivated for more than a century, and has undergone no change.
The tall kinds grown in southern latitudes, and therefore exposed
to great heat, require from six to seven months to ripen their seed;
whereas the dwarf kinds, grown in northern and colder climates,
require only from three to four months.58 Peter Kalm,59 who par-
ticularly attended to this plant says that in the United States, in
proceeding from south to north, the plants steadily diminish in
bulk. Seeds brought from lat. 37° in Virginia, and sown in lat.
43°-44° in New England, produce plants which will not ripen their
seed, or ripen them with the utmost difficulty. So it is with seed
carried from New England to lat. 45°-47° in Canada. By taking
p. 31.
53 Godron, 'De l'Espèce,' tom. ii. p.
80; Al. De Candolle, idem, p. 951.
B4 Transact. Bot. Soc. of Edinburgh,
vol. viii. p. 60.
55 Voyages dans l'Amérique Méri-
dionale,' tom. i. p. 147.
68 Bonafous' Hist. Nat. du Mais,'
57 Idem, p. 31.
58 Metzger, Getreidearten,'s. 206.
59 Description of Maize,' by P. Kalm,
1752, in Swedish Acts,' vol. iy. I
have consulted an old English MS. trang-
lation,

388
CHAP. IX.
CEREAL PLANTS.
great care at first, the southern kinds after some years' culture ripen
their seed perfectly in their northern homes, so that this is an ana-
logous case with that of the conversion of summer into winter
wheat, and conversely. When tall and dwarf maize are planted
together, the dwarf kinds are in full flower before the others have
produced a single flower; and in Pennsylvania they ripen their
seed six weeks earlier than the tall maize. Metzger also mentions a
European maize which ripens its seed four weeks earlier than another
European kind. With these facts, so plainly showing inherited
acclimatisation, we may readily believe Kalm, who states that in
North America maize and some other plants have gradually been
cultivated further and further northward. All writers agree that
to keep the varieties of maize pure they must be planted separately
so that they shall not cross.
The effects of the climate of Europe on the American varieties is
highly remarkable. Metzger obtained seed from various parts of
America, and cultivated several kinds in Germany. I will give an
abstract of the changes observed 60 in one case, namely, with a tall
kind (Breit-korniger mays, Zea altissima) brought from the warmer
parts of America. During the first year the plants were twelve
feet high, and few seeds were perfected; the lower seeds in the ear
kept true to their proper form, but the upper seeds became slightly
changed. In the second generation the plants were from nine to
ten feet in height, and ripened their seed better; the depression on
the outer side of the seed had almost disappeared, and the original
beautiful white colour had become duskier. Some of the seeds had
even become yellow, and in their now rounded form they approached
common European maize. In the third generation nearly all re-
semblance to the original and very distinct American parent-form
was lost. In the sixth generation this maize perfectly resembled a
European variety, described as the second sub-variety of the fifth
race. When Metzger published his book, this variety was still
cultivated near Heidelberg, and could be distinguished from the
common kind only by a somewhat more vigorous growth. Analo-
gous results were obtained by the cultivation of another American
race, the "white-tooth corn," in which the tooth nearly disappeared
even in the second generation. A third race, the “chicken-corn,
did not undergo so great a change, but the seeds became less
polished and pellucid.
These facts afford the most remarkable instance
known to me of the direct and prompt action of climate
60 Getreidearten,' s. 208.

CHAP. IX.
389
CULINARY PLANTS : CABBAGES.
on a plant. It might have been expected that the tall-
ness of the stem, the period of the vegetation, and the
ripening of the seed, would have been thus affected; but
it is a much more surprising fact that the seeds should
have undergone so rapid and great a change. As, how-
ever, flowers, with their product the seed, are formed by
the metamorphosis of the stem and leaves, any modifi-
cation in these latter organs would be apt to extend,
through correlation, to the organs of fructification.
Cabbage (Brassica oleracea).-Every one knows how greatly the
various kinds of cabbage differ in appearance. In the island of Jer-
sey, from the effects of particular culture and of climate, a stalk has
grown to the height of sixteen feet, and “had its spring shoots at
the top occupied by a magpie's nest:" the woody stems are not
unfrequently from ten to twelve feet in height, and are there used
as rafters 61 and as walking-sticks. We are thus reminded that in
certain countries plants belonging to the generally herbaceous order
of the Cruciferæ are developed into trees. Every one can appreciate
the difference between green or red cabbages with great single
heads; Brussel-sprouts with numerous little heads; broccolis and
cauliflowers with the greater number of their flowers in an aborted
condition, incapable of producing seed, and borne in a dense corymb
instead of an open panicle; savoys with their blistered and wrinkled
leaves; and borecoles and kales, which come nearest to the wild pa-
rent-form. There are also various frizzled and laciniated kinds,
some of such beautiful colours that Vilmorin in his Catalogue of
1851 enumerates ten varieties, valued solely for ornament, which
are propagated by seed. Some kinds are less commonly known,
such as the Portuguese Couve Tronchuda, with the ribs of its leaves
greatly thickened ; and the Kohlrabi or choux-raves, with their
stems enlarged into great turnip-like masses above the ground; and
the recently formed new race 62 of choux-raves, already including
nine sub-varieties, in which the enlarged part lies beneath the
ground like a turnip.
Although we see such great differences in the shape, size, colour,
arrangement, and manner of growth of the leaves and stem, and of
61 Cabbage Timber,' 'Gardener's
Chron., '1556, p. 744, quoted from Hook-
er's Journal of Botany,' A walking-
stick made from a cabbage-stalk is ex-
hibited in the Museum at Kew.
62 Journal de la Soc. Imp. d'Horti-
culture,' 1855, p. 254, quoted from "Gar-
tenflora,' Ap. 1855.

390
CHAP. IX
CULINARY PLANTS.
the flower-stems in the broccoli and cauliflower, it is remarkable
that the flowers themselves, the seed-pods, and seeds, present ex-
tremely slight differences or none at all.63 I compared the flowers
of all the principal kinds; those of the Couve Tronchuda are white
and rather smaller than in common cabbages; those of the Ports-
mouth broccoli have narrower sepals, and smaller, less elongated
petals; and in no other cabbage could any difference be detected.
With respect to the seed-pods, in the purple Kohlrabi alone, do they
differ, being a little longer and narrower than usual. I made a col-
lection of the seeds of twenty-eight different kinds, and most of them
were undistinguishable; when there was any difference it was ex-
cessively slight; thus, the seeds of various broccolis and cauliflow-
ers, when seen in mass, are a little redder; those of the early green
Ulm savoy are rather smaller; and those of the Breda kail slightly
larger than usual, but not larger than the seeds of the wild cabbage
from the coast of Wales. What a contrast in the amount of differ-
ence is presented if, on the one hand, we compare the leaves and
stems of the various kinds of cabbage with their flowers, pods, and
seeds, and on the other hand the corresponding parts in the varieties
of maize and wheat ! The explanation is obvious; the seeds alone
are valued in our cereals, and their variations have been selected ;
whereas the seeds, seed-pods, and flowers have been utterly neglect-
ed in the cabbage, whilst many useful variations in their leaves and
stems have been noticed and preserved from an extremely remote
period, for cabbages were cultivated by the old Celts.64
It would be useless to give a classified description 65 of the nume-
rous races, sub-races, and varieties of the cabbage; but it may be
mentioned that Dr. Lindley has lately proposed 66 a system founded
on the state of development of the terminal and lateral leaf-buds,
and of the flower-buds. Thus, I. All the leaf-buds active and open,
as in the wild-cabbage, kail, &c. II. All the leaf-buds active, but
forming heads, as in Brussel-sprouts, &c. III. Terminal leaf-bud
alone active, forming a head as in common cabbages, savoys, &c.
IV. Terminal leaf-bud alone active and open, with most of the
flowers abortive and succulent, as in the cauliflower and broccoli.
V. All the leaf-buds active and open, with most of the flowers abor-
tive and succulent, as in the sprouting-broccoli. This latter vari-
ety is a new one, and bears the same relation to common broccoli, as
63 Godron, De l'Espèce,' tom. ii. p.
52; Metzger, Syst. Beschreibung der
Kult. Kohlarten,' 1833, s. 6.
64 Regnier, De l'Economie Publique
des Celtes,' 1818, p. 433.
65 See the elder De Candolle, in
Transact. of Hort. Soc.,' vol. v.; and
Metzger 'Kohlarten,' &c.
68 Gardener's Chronicle,' 1859, p. 992.

CHAP. IX.
391
CABBAGES.
Brussel-sprouts do to common cabbages; it suddenly appeared in a
bed of common broccoli, and was found faithfully to transmit its
newly-acquired and remarkable characters.
The principal kinds of cabbage existed at least as early as the six-
teenth century,67 so that numerous modifications of structure have
been inherited for a long period. This fact is the more remarkable
as great care must be taken to prevent the crossing of the different
kinds. To give one proof of this : I raised 233 seedlings from cab-
bages of different kinds, which had purposely been planted near each
other, and of the seedlings no less than 155 were plainly deteriora-
ted and mongrelized ; nor were the remaining 78 all perfectly true.
It may be doubted whether many permanent varieties have been
formed by intentional or accidental crosses; for such crossed plants
are found to be very inconstant. One kind, however, called “Cot-
tager's Kale," has lately been produced by crossing common kale
and Brussel-sprouts, recrossed with purple broccoli, 8 and is said to
be true, but plants raised by me were not nearly so constant in
character as any common cabbage.
Although most of the kinds keep true if carefully preserved from
crossing, yet the seed-beds must be yearly examined, and a few
seedlings are generally found false; but even in this case the
force of inheritance is shown, for, as Metzger has remarked 69
when speaking of Brussel-sprouts, the variations generally keep
to their "unter art," or main race. But in order that any kind
may be truly propagated there must be no great change in the
conditions of life; thus cabbages will not form heads in hot coun-
tries, and the same thing has been observed with an English
variety grown during an extremely warm and damp autumn near
Paris.70 Extremely poor soil also affects the characters of certain
varieties.
Most authors believe that all the races are descended from the
wild cabbage found on the western shores of Europe ; but Alph.
De Candolle 71 forcibly argues on historical and other grounds that
it is more probable that two or three closely allied forms, generally
ranked as distinct species, still living in the Mediterranean region,
are the parents, now ali commingled together, of the various culti-
vated kinds. In the same manner as we have often seen with do-
mesticated animals, the supposed multiple origin of the cabbage
67 Alph. De Candolle, 'Géograph. Bot.,
pp. S12 and 989.
88 Gardener's Chron.,' Feb. 1858, p.
129.
69 Kohlarten,' s. 22.
70 Godron, De l'Espèce,' tom. ii. p.
52; Metzger, 'Kohlarten,' s. 22.
71 Géograph. Bot.,' p. 840.

392
CHAP. IX.
CULINARY PLANTS.
throws no light on the characteristic differences between the cul-
tivated forms. If our cabbages are the descendants of three or
four distinct species, every trace of any sterility which may origi-
nally have existed between them is now lost, for none of the varieties
can be kept distinct without scrupulous care to prevent intercross-
ing
The other cultivated forms of the genus Brassica are descended,
according to the view adopted by Godron and Metzger,"2 from two
species, B. napus and rapa; but according to other botanists from
three species; whilst others again strongly suspect that all these
forms, both wild and cultivated, ought to be ranked as a single
species. Brassica napus has given rise to two large groups, name
ly, Swedish turnips (by some believed to be of hybrid origin) 73 and
Colzas, the seeds of which yield oil. Brassica rapa (of Koch) has
also given rise to two races, namely, common turnips and the oil
giving rape. The evidence is unusually clear that these latter
plants, though so different in external appearance, belong to the
same species; for the turnip has been observed by Koch and Go-
dron to lose its thick roots in uncultivated soil, and when rape and
turnips are sown together they cross to such a degree that scarcely
a single plant comes true.74 Metzger by culture converted the bien-
nial or winter rape into the annual or summer rape varieties which
have been thought by some authors to be specifically distinct.15
In the production of large, fleshy, turnip-like stems, we have a
case of analogous variation in three forms which are generally
considered as distinct species. But scarcely any modification seems
so easily acquired as a succulent enlargement of the stem or root-
that is a store of nutriment laid up for the plant's own future use.
We see this in our radishes, beet, and in the less generally known
"turnip-rooted” celery, and in the finocchio or Italian variety of the
common fennel. Mr. Buckman has lately proved by his interesting
experiments how quickly the roots of the wild parsnip can be en-
larged, as Vilmorin formerly proved in the case of the carrot.
76
72
Godron, De l'Espèce,' tom. ii. p.
51; Metzger, Kohlarten,' s. 10.
73 Gardener's Chron. and Agricult.
Gazette,' 1856, p. 729.
74 Gardener's Chron. and Agricult.
Gazette,' 1855, p. 730.
75 Metzger, 'Kohlarten,' s. 51.
76 These experiments by Vilmorin have
been quoted by many writers. An emi-
nent botanist, Prof. Decaisne, has lately
expressed doubts on the subject from his
own negative results, but these cannot
be valued equally with positive results.
On the other hand, M. Carrière has
lately stated (Gard. Chronicle,' 1865,
p. 1154) that he took seed from a wild
carrot, growing far from any cultivated
land, and even in the first generation
the roots of his seedlings differed in
being spindle-shaped, longer, softer and
less fibrous than those of the wild plant.
From these seedlings he raised several
distinct varieties.

CHAP. IX.
393
PEAS.
This latter plant, in its cultivated state, differs in scarcely any
character from the wild English species, except in general luxu-
riance and in the size and quality of its roots; but in the root ten
varieties, differing in colour, shape, and quality, are cultivated in
England, and come true by seed. Hence, with the carrot, as in so
many other cases, for instance with the numerous varieties and
sub-varieties of the radish, that part of the plant which is valued
by man, falsely appears alone to have varied. The truth is that
variations in this part alone have been selected; and the seedlings
inheriting a tendency to vary in the same way, analogous modifica-
tions have been again and again selected, until at last a great amount
of change has been effected.
Pea (Pisum sativum).- Most botanists look at the garden-pea as
specifically distinct from the field-pea (P. arvense). The latter ex-
ists in a wild state in Southern Europe ; but the aboriginal parent
of the garden-pea has been found by one collector alone, as he states,
in the Crimea.78 Andrew Knight crossed, as I am inforned by the
Rev. A. Fitch, the field-pea with a well-known garden variety, the
Prussian pea, and the cross seems to have been perfectly fertile.
Dr. Alefeld has recently studied 19 the genus with care, and, after
having cultivated about fifty varieties, concludes that they all cer-
tainly belong to the same species. It is an interesting fact already
alluded to, that, according to 0. Heer, the peas found in the lake
habitations of Switzerland of the Stone and Bronze ages, belong to
an extinct variety, with exceedingly small seeds, allied to P. ar-
vense, or field-pea. The varieties of the cominon garden-pea are
numerous, and differ considerably from each other. For compari-
son I planted at the same time forty-one English and French varie-
ties, and in this one case I will describe minutely their differences.
The varieties differ greatly in height.-namely from between 6 and
12 inches to 8 feet, 81—in manner of growth, and in period of matu-
rity. Some varieties differ in general aspect even while only two
or three inches in height. The stems of the Prussian pea are
much branched. The tall kinds have larger leaves than the dwarf
kinds, but not in strict proportion to their height:-Hairs' Dwarf
Monmouth has very large leaves, and the Pois nain hatif, and the
77 Loudon's Encyclop. of Gardening,'
p. 835.
78 Alph. De Candolle, 'Géograph. Bot.,'
960. Mr. Bentham (Hort. Journal,' vol.
ix. (1855), p. 141) believes that garden
and field peas belong to the same spe-
cies, and in this respect he differs from
Dr. Targioni.
79 Botanische Zeitung,' 1860, s. 204.
80 Die Pflanzen der Pfahlbauten,'
1866, s. 23.
81 A variety called the Rouncival at-
tains this height, as is stated by Mr.
Gordon in Transact. Hort. Soc.,' (2nd
series), vol. i., 1835, p. 374, from wbich
paper I have taken some facts.

394
CHAP. IX.
CULINARY PLANTS.
moderately tall Blue Prussian, have leaves about two-thirds of the
size of the tallest kind. In the Danecroft the leaflets are rather
small and a little pointed; in the Queen of Dwarfs rather rounded ;
and in the Queen of England broad and large. In these three peas
the slight differences in the shape of the leaves are accompanied by
slight differences in colour. In the Pois géant sans parchemin, which
bears purple flowers, the leaflets in the young plant are edged with
red; and in all the peas with purple flowers the stipules are marked
with red.
In the different varieties, one or two, or several flowers in a small
cluster, are borne on the same peduncle ; and this is a difference
which with some of the Leguminosa is considered of specific value.
In all the varieties the flowers closely resemble each other except in
colour and size. They are generally white, sometimes purple, but
the colour is inconstant even in the same variety. In Warner's
Emperor, which is a tall kind, the flowers are nearly double the size
of those of the Pois nain hatif, but Hairs' Dwarf Monmouth, which
has large leaves, likewise has large flowers. The calyx in the
Victoria Marrow is large, and in Bishop's Long Pod the sepals are
rather narrow. In no other kind is there any difference in the flower.
The pods and seeds, which with natural species afford such con-
stant characters, differ greatly in the cultivated varieties of the pea;
and these are the valuable, and consequently the selected parts.
Sugar peas, or Pois sans parchemin, are remarkable from their
thin pods, which, whilst young, are cooked and eaten whole; and
in this group, whichi according to Mr. Gordon includes eleven sub-
varieties, it is the pod which differs most : thus, Lewis's Negro-
podded pea has a straight, broad, smooth, and dark-purple pod, with
the husk not so thin as in the other kinds; the pod of another
variety is extremely bowed; that of the Pois géant is much pointed
at the extremity; and in the variety " à grands cosses" the peas
are seen through the husk in so conspicuous a manner that the pod,
especially when dry, can hardly at first be recognised as that of a pea.
In the ordinary varieties the pods also differ much in size ;-in
colour, that of Woodford's Green Marrow being bright-green when
dry, instead of pale brown, and that of the purple-podded pea being
expressed by its name;—in smoothness, that of Danecroft being re-
markably glossy, whereas that of the Ne plus ultra is rugged ;-
in being either nearly cylindrical, or broad and flat ;-in being
pointed at the end as in Thurston's Reliance, or much truncated as
in the American Dwarf. In the Auvergne pea the whole end of the
pod is bowed upwards. In the Queen of the Dwarfs and in Scimitar
peas the pod is almost elliptic in shape. I here give drawings of
the four most distinct pods produced by the plants cultivated by me.

CHAP. IX.
395
PEAS.
In the pea itself we have every tint between almost pure white,
brown, yellow, and intense green ; in the varieties of the sugar peas
I.
11.
TIL.
IV.
a
6
с
a
Fig. 41.- Pods and Peas. I. Queen of Dwarfs. II. American Dwarf. III. Thurs-
ton's Reliance. IV. Pois Géant sans parchemin. a. Dan O'Rourke Pea.
6. Queen of Dwarfs Pea. c. Knight's Tall White Marrow. d. Lewis's
Negro Pea.
we have these same tints, together with red passing through fine
purple into a dark chocolate tint. These colours are either uniform

396
CHAP. IX.
CULINARY PLANTS.
or distributed in dots, striæ, or moss-like marks; they depend in
some cases on the colour of the cotyledons seen through the skin,
and in other cases on the outer coats of the pea itself. In the dif-
ferent varieties the pods contain, according to Mr. Gordon, from
eleven or twelve to only four or five peas. The largest peas are
nearly twice as much in diameter as the smallest ; and the latter
are not always borne by the most dwarfed kinds. Peas differ much
in shape, being smooth and spherical, smooth and oblong, nearly
oval in the Queen of Dwarfs, and nearly cubical and crumpled in
many of the larger kinds.
With respect to the value of the differences between the chief
varieties, it cannot be doubted that, if one of the tall Sugar-peas,
with purple flowers, thin-skinned pods of an extraordinary shape,
including large, dark-purple peas, grew wild by the side of the
lowly Queen of the Dwarfs, with white flowers, greyish-green,
rounded leaves, scimitar-like pods, containing oblong, smooth, pale-
coloured peas, which became mature at a different season; or by
the side of one of the gigantic sorts, like the Champion of England,
with leaves of great size, pointed pods, and large, green, crumpled,
almost cubical peas,-all three kinds would be ranked as indispu-
tably distinct species.
Andrew Knight 52 has observed that the varieties of peas keep
very true, because they are not crossed by insects. As far as the
fact of keeping true is concerned, I hear from Mr. Masters of Can-
terbury, well known as the originator of several new kinds, that
certain varieties have remained constant for a considerable time, -
for instance, Knight's Blue Dwarf, which came out about the year
1820.43 But the greater number of varieties have a singularly short
existence: thus Loudon remarks 64 that "sorts which were highly
approved in 1821, are now, in 1833, nowhere to be found," and on
comparing the lists of 1833 with those of 1855, I find that nearly
all the varieties have changed. Mr. Masters informs me that the
nature of the soil causes some varieties to lose their character. As
with other plants, certain varieties can be propagated truly, whilst
others show a determined tendency to vary; thus two peas differing
in shape, one round and the other wrinkled, were found by Mr. Mas-
ters within the same pod, but the plants raised from the wrinkled
kind always evinced a strong tendency to produce round peas. Mr.
Masters also raised from a plant of another variety four distinct
sub-varieties, which bore blue and round, white and round, blue and
82 Phil. Transact.,' 1799, p. 196.
83 Gardener's Magazine,' vol. i., 1826,
84 'Encyclopædia of Gardening,' p.
823.
P. 153.

CHAP. IX.
397
PEAS.
wrinkled, and white and wrinkled peas; and although he sowed
these four varieties separately during several successive years, each
kind always reproduced all four kinds mixed together!
With respect to the varieties not naturally intercrossing, I have
ascertained that the pea, which in this respect differs from some
other Leguminosæ, is perfectly fertile without the aid of insects.
Yet I have seen humble-bees whilst sucking the nectar depress the
keel-petals, and become so thickly dusted with pollen, that some
could hardly fail to be left on the stigma of the next flower which
was visited. I have made inquiries from several great raisers of
seed-peas, and I find that but few sow them separately; the majority
take no precaution; and it is certain, as I have myself found, that
true seed may be saved during at least several generations from
distinct varieties growing close together.86 Under these circum-
stances, Mr. Fitch raised, as he informs me, one variety for twenty
years, which always came true. From the analogy of kidney-
beans I should have expected 86 that occasionally, perhaps at
long intervals of time, when some slight degree of sterility had
supervened from long-continued self-fertilisation, varieties thus
growing near each other would have crossed ; and I shall give
in the eleventh chapter two cases of distinct varieties which sponta-
neously intercrossed, as shown in a manner hereafter to be ex-
plained) by the pollen of the one variety having acted directly on
the seeds of the other. Whether the incessant supply of new
varieties is partly due to such occasional and accidental crosses, and
their fleeting existence to changes of fashion; or again, whether
the varieties which arise after a long course of continued self-ferti-
lisation are weakly and soon perish, I cannot even conjecture. It
may, however, be noticed that several of Andrew Knight's varie-
ties, which have endured longer than most kinds, were raised
towards the close of the last century by artificial crosses; some of
them, I believe, were still, in 1860, vigorous; but now, in 1865, a
writer, speaking of Knight's four kinds of marrows, says, they
have acquired a famous history, but their glory has departed.
With respect to Beans (Faba vulgaris), I will say but little. Dr.
Alefeld has given 68 short diagnostic characters of forty varieties.
Every one who has seen a collection must have been struck with
the great difference in shape, thickness, proportional length and
85 See Dr. Anderson to the same effect
in the Bath Soc. Agricultural Papers,
vol. iv. p. 87.
86 I have published full details of ex-
periments on this subject in the Gar-
dener's Chronicle,' 1857, Oct. 25th.
87 Gardener's Chronicle, 1865, p.
387.
88 Bonplandia,' x., 1862, s. 348.

398
CHAP. IX.
CULINARY PLANTS.
breadth, colour, and size which beans present. What a contrast
between a Windsor and Horse-bean! As in the case of the pea,
our existing varieties were preceded during the Bronze age in
Switzerland by a peculiar and now extinct variety producing very
small beans.89
Potato (Solanum tuberosum).—There is little doubt about the
parentage of this plant; for the cultivated varieties differ extremely
little in general appearance from the wild species, which can be
recognised in its native land at the first glance.90 The varieties
cultivated in Britain are numerous; thus Lawson' gives a descrip-
tion of 175 kinds. I planted eighteen kinds in adjoining rows;
their stems and leaves differed but little, and in several cases there
was as great an amount of difference between the individuals of the
same variety as between the different varieties. The flowers vary
in size, and in colour between white and purple, but in no other
respect, except that in one kind the sepals were somewhat elongated.
One strange variety has been described which always produces two
sorts of flowers, the first double and sterile, the second single and
fertile. The fruit orberries also differ, but only in a slight
degree.93
The tubers, on the other hand, present a wonderful amount of
diversity. This fact accords with the principle that the valuable
and selected parts of all cultivated productions present the greatest
amount of modification. They differ much in size and shape, being
globular, oval, flattened, kidney-like, or cylindrical. One variety
from Peru is described 94 as being quite straight, and at least six
inches in length, though no thicker than a man's finger. The eyes
or buds differ in form, position, and colour. The manner in which
the tubers are arranged on the so-called roots is different; thus in
the gurken-kartoffeln they form a pyramid with the apex down-
wards, and in another variety they bury themselves deep in the
ground. The roots themselves run either near the surface or deep
in the ground. The tubers also differ in smoothness and colour,
being externally white, red, purple, or almost black, and internally
white, yellow, or almost black. They differ in flavour and quality,
Chronicle,' 1845, p. 790.
89 0. Heer, Die Pflanzen der Pfahl-
bauten,' 1866, s. 22.
90 Darwin, 'Journal of Researches,
1945, p. 285.
91 Synopsis of the vegetable products
of Scotland, quoted in Wilson's 'British
Farming,' p. 317.
92 Sir G. Mackenzie, in Gardener's
98 Putsche und Vertuch, Versuch
einer Monographie der Kartoffeln,'1819,
s. 9, 15. See also Dr. Anderson's 'Re-
creations in Agriculture,' vol. iv. p. 325.
94. Gardener's Chronicle,' 1862, p.
1052.

CHAP. IX.
399
POTATOES.
being either waxy or mealy; in their period of maturity, and in
their capacity for long preservation.
As with many other plants which have been long propagated by
bulbs, tubers, cuttings, &c., by which means the same individual is
exposed during a length of time to diversified conditions, seedling
potatoes generally display innumerable slight differences. Several
varieties, even when propagated by tubers, are far from constant,
as will be seen in the chapter on Bud-variation. Dr. Anderson 95
procured seed from an Irish purple potato, which grew far from any
other kind, so that it could not at least in this generation have been
crossed, yet the many seedlings varied in almost every possible
respect, so that “scarcely two plants were exactly alike.” Some of
the plants which closely resembled each other above ground, pro-
duced extremely dissimilar tubers, and some tubers which exter-
nally could hardly be distinguished, differed widely in quality when
cooked. Even in this case of extreme variability, the parent-stock
had some influence on the progeny, for the greater number of the
seedlings resembled in some degree the parent Irish potato. Kid-
ney potatoes must be ranked amongst the most highly cultivated
and artificial races; yet their peculiarities can often be strictly pro-
pagated by seed. A great authority, Mr. Rivers, 96 states that "seed-
lings from the ash-leaved kidney always bear a strong resemblance
to their parent. Seedlings from the fluke-kidney are still more
remarkable for their adherence to their parent-stock, for, on closely
observing a great number during two seasons, I have not been able
to observe the least difference either in earliness, productiveness, or
in the size or shape of their tubers."
05 Bath Society Agricult. Papers,
vol. v. p. 127. And Recreations in
Agriculture,' vol. v. p. 86.
96 Gardener's Chronicle,' 1863, p.
613.

400
CHAP. X.
FRUITS.
CHAPTER X.
PLANTS continued - FRUITS - ORNAMENTAL TREES –
FLOWERS.
FRUITS. — GRAPES — VARY IN ODD AND TRIFLING PARTICULARS.
MULBERRY. THE ORANGE GROUP SINGULAR RESULTS
FROM CROSSING. PEACH AND NECTARINE - BUD-VARIATION
ANALOGOUS VARIATION RELATION TO THE ALMOND.
APRICOT. - PLUMS — VARIATION IN THEIR STONES. CHER-
RIES 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. - DAH-
LIA. - HYACINTI — HISTORY AND VARIATION OF.
The Vine (Vitis vinifera).—THE best authorities consider all our
grapes as the descendants of one species which now grows wild in
western Asia, which grew during the Bronze-age wild in Italy,' and
which has recently been found fossil in a tufaceous deposit in the south
of France. Some authors, however, entertain much doubt about the
single parentage of our cultivated varieties, owing to the number
of semi-wild forms found in Southern Europe, especially as de-
scribed by Clemente, in a forest in Spain ; but as the grape sows
1 Heer, 'Pflanzen der Pfahlbauten,'
1866, s. 28.
2 Alph. De Candolle, 'Géograph. Bot.,'
p. 872; Dr. A. Targioni-Tozzetti, in
Jour. Hort. Soc.,' vol. ix. p. 133. For
the fossil vine found by Dr. G. Planchon,
see Nat. Hist. Review,' 1865, April, p.
224.
3 Godron, 'De l'Espèce,' tom. ii. p.
100.

CHAP. X.
401
VINES.
itself freely in Southern Europe, and as several of the chief kinds
transmit their characters by seed, whilst others are extremely
variable, the existence of many different escaped forms could hardly
fail to occur in countries where this plant has been cultivated from
the remotest antiquity. That the vine varies much when propaga-
ted by seed, we may infer from the largely increased number of
varieties since the earlier historical records. New hot-house varie-
ties are produced almost every year; for instance, a golden-coloured
variety has been recently raised in England from a black grape
without the aid of a cross. Van Mons. reared a multitude of va-
rieties from the seed of one vine, which was completely separated
from all others, so that there could not, at least in this generation,
have been any crossing, and the seedlings presented " les analogues
de toutes les sortes," and differed in almost every possible character,
both in the fruit and foliage.
The cultivated varieties are extremely numerous; Count Odart
says that he will not deny that there may exist throughout the
world 700 or 800, perhaps even 1000 varieties, but not a third of these
have any value. In the Catalogue of fruit cultivated in the Horticul-
tural Gardens of London, published in 1842, 99 varieties are enu-
merated. Wherever the grape is grown many varieties occur: Pal-
las describes 24 in the Crimea, and Burnes mentions 10 in Cabool.
The classification of the varieties has much perplexed writers, and
Count Odart is reduced to a geographical system ; but I will not
enter on this subject, nor on the many and great differences between
the varieties. I will merely specify a few curious and trifling pecu-
liarities, all taken from Odart's highly esteemed work, for the sake
of showing the diversified variability of this plant. Simon has classed
grapes into two main divisions, those with downy leaves and those
with smooth leaves, but he admits that in one variety, namely the
Rebazo, the leaves are either smooth or downy; and Odart (p. 70)
states that some varieties have the nerves alone, and other varieties
their young leaves, downy, whilst the old ones are smooth. The
Pedro-Ximenes grape (Odart, p. 397) presents a peculiarity by which
it can be at once recognised amongst a host of other varieties, name-
ly, that when the fruit is nearly ripe the nerves of the leaves or even
the whole surface becomes yellow. The Barbera d'Asti is well
marked by several characters (p. 426), amongst others, “ by some of
4 See an account of M. Vibert's ex-
periments, by Alex. Jordan, in Mém.
de l'Acad. de Lyon,' tom. ii., 1852, P.
108
6. Gardner's Chronicle,'1864, p. 488.
6 Arbres Fruitiers,' 1936, tom. ii. p.
290.
7 Odart, 'Ampelographie Universelle,
1849.

402
CHAP. X.
FRUITS.
the leaves, and it is always the lowest on the branches, suddenly be-
coming of a dark red colour.” Several authors in classifying grapes
have founded their main divisions on the berries being either round
or oblong; and Odart admits the value of this character; yet there
is one variety, the Maccabeo (p. 71), which often produces small
round, and large oblong, berries in the same bunch. Certain grapes
called Nebbiolo (p. 429) present a constant character, sufficient for
their recognition, namely, “the slight adherence of that part of the
pulp which surrounds the seeds to the rest of the berry, when cut
through transversely." A Rhenish variety is mentioned (p. 228)
which likes a dry soil; the fruit ripens well, but at the moment of
maturity, if much rain falls, the berries are apt to rot; on the other
hand, the fruit of a Swiss variety (p. 243) is valued for well sustain-
ing prolonged humidity. This latter variety sprouts late in the
spring, yet matures its fruit early; other varieties (p. 362) have the
fault of being too much excited by the April sun, and in consequence
suffer from frost. A Styrian variety (p. 254) has brittle foot-stalks,
so that the clusters of fruit are often blown off; this variety is said
to be particularly attractive to wasps and bees. Other varieties have
tough stalks, which resist the wind. Many other variable charac-
ters could be given, but the foregoing facts are sufficient to show
in how many small structural and constitutional details the vine
varies. During the vine disease in France certain whole groups of
varieties have suffered far more from mildew than others. Thus
“the group of the Chasselas, so rich in varieties, did not afford a sin-
gle fortunate exception ;" certain other groups suffered much less;
the true old Burgundy, for instance, was comparatively free from
disease, and the Carminat likewise resisted the attack. The Ameri-
can vines, which belong to a distinct species, entirely escaped the
disease in France; and we thus see that those European varieties
which best resist the disease must have acquired in a slight degree
the same constitutional peculiarities as the American species.
White Mulberry (Morus alba).-I mention this plant because it has
varied in certain characters, namely, in the texture and quality of
the leaves, fitting them to serve as food for the domesticated silk-
worm, in a manner not observed with other plants; but this has
arisen simply from such variations in the mulberry having been at-
tended to, selected, and rendered more or less constant. M. de Qua-
trefages' briefly describes six kinds cultivated in one valley in
France: of these the amourouso produces excellent leaves, but is
8 M. Bouchardat, in Comptes Ren-
dus,' Dec. 1st, 1851, quoted in Garden-
er's Chron.,' 1852, p. 435.
9 Etudes sur les Maladies actuelles
du Ver à Soie,' 1859, p. 321.

CHAP. X.
403
ORANGE GROUP.
rapidly being abandoned because it produces much fruit mingled
with the leaves: the antofino yields deeply cut leaves of the finest
quality, but not in great quantity: the claro is much sought for be-
cause the leaves can be easily collected: lastly, the roso bears strong
hardy leaves, produced in large quantity, but with the one incon-
venience, that they are best adapted for the worms after their fourth
moult. MM. Jacquemet-Bonnefont, of Lyon, however, remark in
their catalogue (1862) that two sub-varieties have been confounded
under the name of the roso, one having leaves too thick for the ca-
terpillars, the other being valuable because the leaves can easily be
gathered from the branches without the bark being torn.
In India the mulberry has also given rise to many varieties. The
Indian form is thought by many botanists to be a distinct species;
but as Royle remarks,19 so many varieties have been produced by
cultivation that it is difficult to ascertain whether they all belong to
one species;" they are, as he adds, nearly as numerous as those of
the silkworm.
The Orange Group.–We here meet with great confusion in the
specific distinction and parentage of the several kinds. Gallesio, 11
who almost devoted his life-time to the subject, considers that there
are four species, namely, sweet and bitter oranges, lemons, and ci-
trons, each of which has given rise to whole groups of varieties,
monsters, and supposed hybrids. One high authority 12 believes that
these four reputed species are all varieties of the wild Citrus medica,
but that the shaddock (Citrus decumana), which is not known in a
wild state, is a distinct species; though its distinctness is doubted by
another writer “of great authority on such matters," namely, Dr.
Buchanan Hamilton. Alph. De Candolle, 18 on the other hand-and
there cannot be a more capable judge-advances what he considers
sufficient evidence of the orange (he doubts whether the bitter and
sweet kinds are specifically distinct), the lemon, and citron, having
been found wild, and consequently that they are distinct. He men-
tions two other forms caltivated in Japan and Java, which he ranks
as undoubted species; he speaks rather more doubtfully about the
shaddock, which varies much, and has not been found wild; and
finally he considers some forms, such as Adam's apple and the ber-
gamotte, as probably hybrids.
10 Productive Resources of India,' p.
130.
11 Traité du Citrus,' 1911. Teoria
della Riproduzione Vegetale,' 1816. I
quote chiefly from this second work. In
1839 Gallesio published in folio Gli
Agrumi dei Giard. Bot. di Firenze,' in
which he gives a curious diagram of the
supposed relationship of all the forms.
12 Mr. Bentham, Review of Dr. A.
Targioni-Tozzetti, 'Journal of Hort. Soc.,'
vol. ix. p. 133.
Géograph. Bot.,' p. 863.
136

104
CHAP. X
FRUITS.
I have briefly abstracted these opinions for the sake of showing
those who have never attended to such subjects, how perplexed with
doubt they are. It would, therefore, be useless for my purpose to
give a sketch of the conspicuous differences between the several
forms. Besides the ever-recurrent difficulty of determining whether
forms found wild are truly aboriginal or are escaped seedlings, many
of the forms, which must be ranked as varieties, transmit their cha-
racters almost perfectly by seed. Sweet and bitter oranges differ in
no important respect except in the flavour of their fruit, but Gallesio 24
is most emphatic that both kinds can be propagated by seed with
absolute certainty. Consequently, in accordance with his simple
rule, he classes them as distinct species; as he does sweet and bitter
almonds, the peach and nectarine, &c. He admits, however, that
the soft-shelled pine-tree produces not only soft-shelled but some
hard-shelled seedlings, so that a little greater force in the power of
inheritance would, according to this rule, raise the soft-shelled pine-
tree into the dignity of an aboriginally created species. The posi-
tive assertion made by Macfayden 15 that the pips of sweet oranges
produce in Jamaica, according to the nature of the soil in which
they are sown, either sweet or bitter oranges, is probably an error ;
for M. Alph. De Candolle informs me that since the publication of his
great work he has received accounts from Guiana, the Antilles, and
Mauritius, that in these countries sweet oranges faithfully transmit
their character. Gallesio found that the willow-leafed and the Little
China oranges reproduced their proper leaves and fruit; but the
seedlings were not quite equal in merit to their parents. The red-
fleshed orange, on the other hand, fails to reproduce itself. Gallesio
also observed that the seeds of several other singular varieties all
reproduced trees having a peculiar physiognomy, but partly resem-
bling their parent-forms. I can adduce another case: the myrtle-
leaved orange is ranked by all authors as a variety, but is very dis-
tinct in general aspect: in my father's greenhouse, during many
years, it rarely yielded any seed, but at last produced one; and a
tree thus raised was identical with the parent-form.
Another and more serious difficulty in determining the rank of the
several forms is that, according to Gallesio,16 they largely intercross
without artificial aid ; thus he positively states that seeds taken from
lemon-trees (C. lemonum) growing mingled with the citron (C. me-
dica), which is generally considered as a distinct species, produced
a graduated series of varieties between these two forms. Again, an
14 Teoria della Riproduzione,' pp.
52-57.
15 Hooker's Bot. Misc.,' vol. i. p.
302 ; vol. ii. p. 111.
16 Teoria della Riproduzione,' p. 53.

CHAP. X.
405
ORANGE GROUP.
Adam's apple was produced from the seed of a sweet orange, which
grew close to lemons and citrons. But such facts hardly aid us in de-
termining whether to rank these forms as species or varieties; for it
is now known that undoubted species of Verbascum, Cistus, Primula,
Salix, &c., frequently cross in a state of nature. If indeed it were
proved that plants of the orange tribe raised from these crosses
were even partially sterile, it would be a strong argument in favour
of their rank as species. Gallesio asserts that this is the case; but
he does not distinguish between sterility from hybridism and from
the effects of culture; and he almost destroys the force of this state-
ment by another, 17 namely, that when he impregnated the flowers
of the common orange with the pollen taken from undoubted varie-
ties of the orange, monstrous fruits were produced, which included
"little pulp, and had no seeds, or imperfect seeds."
In this tribe of plants we meet with instances of two highly re-
markable facts in vegetable physiology: Gallesio 18 impregnated an
orange with pollen from a lemon, and the fruit borne on the mother
tree had a raised stripe of peel like that of a lemon both in colour
and taste, but the pulp was like that of an orange and included only
imperfect seeds. The possibility of pollen from one variety or
species directly affecting the fruit produced by another variety or
species, is a subject which I shall fully discuss in the following
chapter.
The second remarkable fact is that two supposed hybrids 19 (for
their hybrid nature was not ascertained) between an orange and
either a lemon or citron produced, on the same tree, leaves, flowers,
and fruit of both pure parent-forms, as well as of a mixed or crossed
nature. A bud taken from any one of the branches and grafted
on another tree produces either one of the pure kinds or a capricious
tree reproducing the three kinds. Whether the sweet lemon, which
includes within the same fruit segments of differently flavoured
pulp,20 is an analogous case, I know not. But to this subject I shall
have to recur.
I will conclude by giving from A. Risso 21 a short account of a very
singular variety of the common orange. It is the “citrus aurantium
fructu variabili," which on the young shoots produces rounded-oval
leaves spotted with yellow, borne on petioles with heart-shaped
wings; when these leaves fall off, they are succeeded by longer and
narrower leaves, with undulated margins, of a pale-green colour
17 Gallesio, Teoria della Riprodu-
zione,' p. 69.
18 Gallesio, idem, p. 67.
19 Gallesio, idem, pp. 75, 76.
20 Gardener's Chronicle,' 1841, p.
613.
21 Annales du Muséum,' tom. xx. p.
188.

406
CHAP. X.
FRUITS,
embroidered with yellow, borne on foot-stalks without wings. The
fruit whilst young is pear-shaped, yellow, longitudinally striated,
and sweet ; but as it ripens, it becomes spherical, of a reddish-yellow,
and bitter.
1
2
3
5
6
7
8
Fig. 42.-Peach and Almond Stones, of natural size, viewed edgeways. 1. Common
English Peach. 2. Double, crimson-flowered, Chinese Peach. 3. Chinese Honey
Peach. 4. English Almond. 5. Barcelona Almond. 6. Malaga Almond. 7. Soft-
shelled French Almond. S. Smyrna Almond.
Peach and Nectarine (Amygdalus Persica).—The best authorities
are nearly unanimous that the peach has never been found wild,

CHAP. X.
407
PEACH AND NECTARINE.
It was introduced from Persia into Europe a little before the Chris.
tian era, and at this period few varieties existed. Alph. De Can-
dolle, 22 from the fact of the peach not having spread from Persia at
an earlier period, and from its not having pure Sanscrit or Hebrew
names, believes that it is not an aboriginal of Western Asia, but
came from the terra incognita of China. The supposition, however,
that the peach is a modified almond which acquired its present cha-
racter at a comparatively late period, would, I presume, account for
these facts; on the same principle that the nectarine, the offspring
of the peach, has few native names, and became known in Europe
at a still later period.
Andrew Knight,23 from finding that a seedling-tree, raised from a
sweet almond fertilised by the pollen of a peach, yielded fruit quite
like that of a peach, suspected that the peach-tree is a modified al-
mond; and in this he has been followed by various authors.24 A
first-rate peach, almost globular in shape, formed of soft and sweet
pulp, surrounding a hard, much furrowed, and slightly-flattened
stone, certainly differs greatly from an almond, with its soft, slight-
ly furrowed, much flattened, and elongated stone, protected by a
tough, greenish layer of bitter flesh. Mr. Bentham 2 has particular-
ly called attention to the stone of the almond being so much more
flattened than that of the peach. But in the several varieties of the
almond, the stone differs greatly in the degree to which it is com-
pressed, in size, shape, strength, and in the depth of the furrows, as
may be seen in the accompanying drawings (Nos. 4 to 8) of such
kinds as I have been able to collect. With peach-stones also Nos.
1 to 3) the degree of compression and elongation is seen to vary; so
that the stone of the Chinese Honey-peach (fig. 3) is much more
elongated and compressed than that of the (No. 8) Smyrna almond.
Mr. Rivers of Sawbridgeworth, to whom I am indebted for some of
the specimens above figured, and who has had such great horticul-
tural experience, has called my attention to several varieties which
connect the almond and the peach. In France there is a variety
called the Peach-almond, which Mr. Rivers formerly cultivated, and
which is correctly described in a French catalogue as being oval and
22 Géograph. Bot.' p. 882.
23 Transactions of Hort. Soc.,' vol.
iii. p. 1, and vol. iv. p. 369, and note to
p. 370. A coloured drawing is given of
this hybrid
24 Gardener's Chronicle,' 1856, p.
532. A writer, it may be presumed Dr.
Lindley, remarks on the perfect series
which may be formed between the al.
mond and the peach. Another high au-
thority, Mr. Rivers, who has had such
wide experience, strongly suspects ("Gar-
dener's Chronicle,' 1863, p. 27) that
peaches, if left to a state of nature, would
in the course of time retrograde into thick-
fleshed almonds.
25 Journal of Hort. Soc.,' vol. ix. p.
168.

408
CHAP. X.
FRUITS.
swollen, with the aspect of a peach, including a hard stone sur-.
rounded by a fleshy covering, which is sometimes eatable 26 A re-
markable statement by M. Luizet has recently appeared in the 'Re-
vue Horticole,' 27 namely, that a Peach-almond, grafted on a peach,
bore during 1863 and 1864 almonds alone, but in 1865 bore six
peaches and no almonds. M. Carrière, in commenting on this fact,
cites the case of a double-flowered almond which, after producing
during several years almonds, suddenly bore for two years in suc-
cession spherical fleshy peach-like fruits, but in 1865 reverted to its
former state and produced large almonds.
Again, as I hear from Mr. Rivers, the double-flowering Chinese
peaches resemble almonds in their manner of growth and in their
flowers; the fruit is much elongated and flattened, with the flesh
both bitter and sweet, but not uneatable, and it is said to be of
better quality in China. From this stage one small step leads us to
such inferior peaches as are occasionally raised from seed. For in-
stance, Mr. Rivers sowed a number of peach-stones imported from
the United States, where they are collected for raising stocks, and
some of the trees raised by him produced peaches which were very
like almonds in appearance, being small and hard, with the pulp
not softening till very late in the autumn. Van Mons 28 also states
that he once raised from a peach-stone a peach having the aspect of
a wild tree, with fruit like that of the almond. From inferior peaches,
such as these just described, we may pass by small transitions,
through clingstones of poor quality, to our best and more melting
kinds. From this gradation, from the cases of sudden variation
above recorded, and from the fact that the peach has not been found
wild, it seems to me by far the most probable view, that the peach is
the descendant of the almond, improved and modified in a marvel-
lous manner.
One fact, however, is opposed to this conclusion. A hybrid, raised
by Knight from the sweet almond by the pollen of the peach, pro-
duced flowers with little or no pollen, yet bore fruit, having been
apparently fertilised by a neighbouring nectarine. Another hybrid
from a sweet almond by the pollen of a nectarine produced during
the first three years imperfect blossoms, but afterwards perfect
flowers with an abundance of pollen. If this slight degree of ste-
rility cannot be accounted for by the youth of the trees (and this
26 Whether this is the same variety as
one lately mentioned ('Gard. Chron.'
1865, p. 1154) by M. Carrière under the
name of Persica intermedia, I know
not: this var. is said to be intermediate
in nearly all its characters between the
almond and peach; it produces during
successive years very different kinds of
fruit.
27 Quoted in "Gard. Chron.'1866, p.
800.
28 Quoted in Journal de la Soc. Imp.
d'Horticulture,' 1855, p. 238.

CHAP. X.
PEACH AND NECTARINE.
409
often causes lessened fertility), or by the monstrous state of the
flowers, or by the conditions to which the trees were exposed, these
two cases would afford a strong argument against the peach being
the descendant of the almond.
Whether or not the peach has proceeded from the almond, it has
certainly given rise to nectarines, or smooth peaches, as they are
called by the French. Most of the varieties both of the peach and
nectarine reproduce themselves truly by seed. Gallesio 29 says he has
verified this with respect to eight races of the peach. Mr. Rivers 30 has
given some striking instances from his own experience, and it is no-
torious that good peaches are constantly raised in North America
from seed. Many of the American sub-varieties come true or near-
ly true to their kind, such as the white-blossom, several of the yel-
low-fruited freestone peaches, the blood clingstone, the heath, and
the lemon-clingstone. On the other hand, a clingstone peach has
been known to give rise to a freestone. 31 In England it has been
noticed that seedlings inherit from their parents flowers of the same
size and colour. Some characters, however, contrary to what might
have been expected, often are not inherited; such as the presence
and form of the glands on the leaves. With respect to nectarines,
both cling and freestones are known in North America to reproduce
themselves by seed.33 In England the new white nectarine was a
seedling of the old white, and Mr. Rivers 34 has recorded several
similar cases. From this strong tendency to inheritance, which
both peach and nectarine trees exhibit,-from certain slight consti-
tutional differences 35 in their nature,--and from the great difference
in their fruit both in appearance and flavour, it is not surprising,
notwithstanding that the trees differ in no other respects and can-
not even be distinguished, as I am informed by Mr. Rivers, whilst
young, that they have been ranked by some authors as specifically
distinct. Gallesio does not doubt that they are distinct; even Alph.
De Candolle does not appear perfectly assured of their specific iden-
tity; and an eminent botanist has quite recently 36 maintained that
the nectarine "probably constitutes a distinct species.
29 Teoria della Riproduzione Vege-
tale,' 1816, p. 86.
30 Gardener's Chronicle,' 1862, p.
1195.
31 Mr. Rivers, Gardener's Chronicle,'
1859, p. 774.
32 Downing, The Fruits of America,'
1845, pp. 475, 489, 492, 494, 496. See
also F. Michaux, Travels in N. Ameri-
ca' (Eng. translat.), p. 228. For similar
cases in France see Godron, De l'Es-
pèce,' tom. ii. p. 97.
33 Brickell's 'Nat. Hist, of N. Caro-
lina,' p. 102, and Downing's Fruit
Trees,' p. 505.
34 Gardener's Chronicle,' 1862, p.
1196.
35 The peach and nectarine do not suc-
ceed equally well in the same soil : see
Lindley's 'Horticulture,' p. 351.
36 Godron, 'De l'Espèce,' tom. ii. 1859,
p. 97.
18

410
CHAP. X.
FRUITS.
Hence it may be worth while to give all the evidence on the origin
of the nectarine. The facts in themselves are curious, and will
hereafter have to be referred to when the important subject of bud-
variation is discussed. It is asserted 97 that the Boston nectarine was
produced from a peach-stone, and this nectarine reproduced itself by
seed.38 Mr. Rivers states 39 that from stones of three distinct varie-
ties of the peach he raised three varieties of nectarine; and in one
of these cases no nectarine grew near the parent peach-tree. In an-
other instance Mr. Rivers raised a nectarine from a peach, and in the
succeeding generation another nectarine from this nectarine.40 Other
such instances have been communicated to me, but they need not be
given. Of the converse case, namely, of nectarine-stones yielding
peach-trees (both free and clingstones), we have six undoubted in-
stances recorded by Mr. Rivers; and in two of these instances the
parent nectarines had been seedlings from other nectarines.41
With respect to the more curious case of full-grown peach-trees
suddenly producing nectarines by bud-variation (or sports as they are
called by gardeners), the evidence is superabundant; there is also
good evidence of the same tree producing both peaches and necta-
rines, or half and half fruit ;-by this term I mean a fruit with the
one-half a perfect peach, and the other half a perfect nectarine.
Peter Collinson in 1741 recorded the first case of a peach-tree pro-
ducing a nectarine,12 and in 1766 he added two other instances. In
the same work, the editor, Sir J. E. Smith, describes the more re-
markable case of a tree in Norfolk, which usually bore both perfect
nectarines and perfect peaches; but during two seasons some of the
fruit were half-and-half in nature.
Mr. Salisbury in 1808 43 records six other cases of peach-trees pro-
ducing nectarines. Three of the varieties are named; viz., the Al-
berge, Belle Chevreuse, and Royal George. This latter tree seldom
failed to produce both kinds of fruit. He gives another case of a
half-and-half fruit.
At Radford in Devonshire 44 a clingstone peach, purchased as the
Chancellor, was planted in 1815, and in 1824, after having previous-
ly produced peaches alone, bore on one branch twelve nectarines;
in 1825 the same branch yielded twenty-six nectarines, and in 1826
1859, p. 774; 1862, p. 1195; 1865, p.
1059; and Journal of Hort.,' 1866, p.
102.
87 Transact. Hort. Soc.,' vol. vi. p.
394.
38 Downing's 'Fruit Trees,' p. 502.
99 Gardener's Chronicle,' 1862, p.
1195.
40 Journal of Horticulture,' Feb. 6th,
1866, p. 102.
41 Mr. Rivers, in Gardener's Chron.,'
42 Correspondence of Linnæus, '1821,
pp. 7, 8, 70.
43 Transact. Hort. Soc.,' vol. i. p. 103.
44 Loudon's "Gardener's Mag.,' 1826,
vol. i. p. 471.

CHAP. X.
411
PEACH AND NECTARINE.
thirty-six nectarines together with eighteen peaches. One of the
peaches was almost as smooth on one side as a nectarine. The nec-
tarines were as dark as, but smaller than, the Elruge.
At Beccles a Royal George peach 45 produced a fruit," three parts
of it being peach and one part nectarine, quite distinct in appear-
ance as well as in flavour.” The lines of division were longitudinal,
as represented in the engraving. A nectarine-tree grew five yards
from this tree.
Professor Chapman states 16 that he has often seen in Virginia
very old peach-trees bearing nectarines.
A writer in the Gardener's Chronicle' says that a peach-tree
planted fifteen years previously 47 produced this year a nectarine
between two peaches; a nectarine-tree grew close by.
In 1844 48 a Vanguard peach-tree produced, in the midst of its or-
dinary fruit, a single red Roman nectarine.
Mr. Calver is stated 49 to have raised in the United States a seed-
ling peach which produced a mixed crop of both peaches and necta
rines.
Near Dorking 50 a branch of the Têton de Venus peach, which re-
produces itself truly by seed, 51 bore its own fruit "so remarkable for
its prominent point, and a nectarine rather smaller but well formed
and quite round."
The previous cases all refer to peaches suddenly producing nec-
tarines, but at Carclew 52 the unique case occurred, of a nectarine-
tree, raised twenty years before from seed and never grafted, pro-
ducing a fruit half peach and half nectarine; subsequently it bore a
perfect peach.
To sum up the foregoing facts: we have excellent evidence of
peach-stones producing nectarine-trees, and of nectarine-stones pro-
ducing peach-trees,-of the same tree bearing peaches and necta-
rines,—of peach-trees suddenly producing by bud-variation necta-
rines (such nectarines reproducing nectarines by seed), as well as
fruit in part nectarine and in part peach,-and lastly of one nec-
tarine-tree first bearing half-and-half fruit, and subsequently true
peaches. As the peach came into existence before the nectarine, it
might have been expected from the law of reversion that nectarines
would give birth by bud-variation or by seed to peaches, oftener
than peaches to nectarines; but this is by no means the case.
p. 53.
45 Loudon's "Gardener's Mag.,' 1828, 49 Phytologist,' vol. iv. p. 299.
50 Gardener's Chron.,'1856, p. 531.
48 Ibid., 1830, p. 597.
51 Godron, 'De l'Espèce,' tom. ii. p.
47 "Gardener's Chronicle,'1841, p. 617. 97.
48 Gardener's Chronicle,'1814, p. 589. 52 Gardener's Chron.,' 1856, p. 531.

412
CHAP. X.
FRUITS.
Two explanations have been suggested to account for these con-
versions. First, that the parent-trees have been in every case hy-
brids 53 between the peach and nectarine, and have reverted by bud-
variation or by seed to one of their pure parent-forms. This view
in itself is not very improbable; for the Mountaineer peach, which
was raised by Knight from the red nutmeg peach by pollen of the
violette hâtive nectarine, 54 produces peaches, but these are said
sometimes to partake of the smoothness and flavour of the necta-
rine. But let it be observed that in the previous list no less than
six well-known varieties and several other unnamed varieties of the
peach have once suddenly produced perfect nectarines by bud-varia-
tion; and it would be an extremely rash supposition that all these
varieties of the peach, which have been cultivated for years in many
districts, and which show not a vestige of a mixed parentage, are,
nevertheless, hybrids. A second explanation is, that the fruit of the
peach has been directly affected by the pollen of the nectarine: al-
though this certainly is possible, it cannot here apply; for we have
not a shadow of evidence that a branch which has borne fruit di-
resily affected by foreign pollen is so profoundly modified as after-
wards to produce buds which continue to yield fruit of the new and
modified form. Now it is known that when a bud on a peach-tree
has once borne a nectarine the same branch has in several instances
gone on during successive years producing nectarines. The Car-
clew nectarine, on the other hand, first produced half-and-half fruit,
and subsequently pure peaches. Hence we may confidently accept
the common view that the nectarine is a variety of the peach, which
may be produced either by bud-variation or from seed. In the follow-
ing chapter many analogous cases of bud-variation will be given.
The varieties of the peach and nectarine run in parallel lines. In
both classes the kinds differ from each other in the flesh of the fruit
being white, red, or yellow; in being clingstones or freestones; in
the flowers being large or small, with certain other characteristic
differences; and in the leaves being serrated without glands, or
crenated and furnished with globose or reniform glands.55 We can
hardly account for this parallelism by supposing that each variety
of the nectarine is descended from a corresponding variety of the
peach ; for though our nectarines are certainly the descendants of
several kinds of peaches, yet a large number are the descendants
of other nectarines, and they vary so much when thus reproduced
that we can scarcely admit the above explanation.
53 Alph. De Candolle, Géograph.
Bot.,' p. 886.
54 Thompson, in Loudon's Encyclop.
of Gardening,' p. 911.
55 Catalogue of Fruit in Garden of
Hort. Soc.,' 1842, p. 105.

CHAP. X.
413
PEACH AND NECTARINE.
The varieties of the peach have largely increased in number
since the Christian era, when from two to five varieties alone were
known ; 56 and the nectarine was unknown. At the present time, ,
besides many varieties said to exist in China, Downing describes in
the United States seventy-nine native and imported varieties of the
peach ; and a few years ago Lindley 57 enumerated one hundred and
sixty-four varieties of the peach and nectarine grown in England.
I have already indicated the chief points of difference between the
several varieties. Nectarines, even when produced from distinct
kinds of peaches, always possess their own peculiar flavour, and are
smooth and small. Clingstone and freestone peaches, which differ
in the ripe flesh either firmly adhering to the stone, or easily sepa-
rating from it, also differ in the character of the stone itself; that
of the freestones or melters being more deeply fissured, with the
sides of the fissures smoother than in clingstones. In the various
kinds, the flowers differ not only in size, but in the larger flowers
the petals are differently shaped, more imbricated, generally red in
the centre and pale towards the margin; whereas in the smaller
flowers the margins of the petal are usually more darkly coloured.
One variety has nearly white flowers. The leaves are more or less
serrated, and are either destitute of glands, or have globose or reni
form glands; 58 and some few peaches, such as the Brugnon, bear
on the same tree both globular and kidney-shaped glands. Ac-
cording to Robertson 60 the trees with glandular leaves are liable to
blister, but not in any great degree to mildew; whilst the non-
glandular trees are more subject to curl, to mildew, and to the
attacks of aphides. The varieties differ in the period of their ma-
turity, in the fruit keeping well, and in hardiness,—the latter cir-
cumstance being especially attended to in the United States. Cer-
tain varieties, such as the Bellegarde, stand forcing in hot-houses
better than other varieties. The flat-peach of China is the most
remarkable of all the varieties; it is so much depressed towards
the summit, that the stone is here covered only by roughened skin
and not by a fleshy layer.61 Another Chinese variety, called the
Honey-peach, is remarkable from the fruit terminating in a long
58 Dr. A. Targioni-Tozzetti, Journal
Hort. Soc.,' vol. ix. p. 167. Alph. De
Candolle, 'Géograph. Bot.,' p. 885.
57 Transact. Hort. Soc.,' vol. v. p.
554.
58 Loudon's Encyclop, of Gardening,'
P. 907.
09 M. Carrière, in "Gard. Chron.,'
1865, p. 1154.
60 Transact. Hort. Soc.,' vol. iii. p.
332. See also "Gardener's Chronicle,'
1865, p. 271, to same effect. Also
"Journal of Horticulture,' Sept. 26th,
1865, p. 254.
61 Transact. Hort. Soc.,' vol. iv. p.
512.

414
CHAP. X
FRUITS.
sharp point; its leaves are glandless and widely dentate.62 The
Emperor of Russia peach is a third singular variety, having deeply
and doubly serrated leaves; the fruit is deeply cleft with one-half
projecting considerably beyond the other; it originated in America,
and its seedlings inherit similar leaves.63
The peach has also produced in China a small class of trees
valued for ornament, namely the double-flowered; of these five
varieties are now known in England, varying from pure white,
through rose, to intense crimson.64 One of these varieties, called
the camellia-flowered, bears flowers above 21 inches in diameter,
whilst those of the fruit-bearing kinds do not at most exceed 11
inch in diameter. The flowers of the double-flowered peaches have
the singular property 65 of frequently producing double or treble
fruit. Finally, there is good reason to believe that the peach is an
almond profoundly modified; but whatever its origin may have
been, there can be no doubt that it has yielded during the last
eighteen centuries many varieties, some of them strongly character-
ised, belonging both to the nectarine and peach form.
Apricot (Prunus armeniaca).-It is commonly admitted that this
tree is descended from a single species, now found wild in the Cauca-
sian region. On this view the varieties deserve notice, because
they illustrate differences supposed by some botanists to be of spe-
cific value in the almond and plum. The best monograph on the
apricot is by Mr. Thompson,67 who describes seventeen varieties.
We have seen that peaches and nectarines vary in a strictly parallel
manner; and in the apricot, which forms a closely allied genus, we
again meet with variations analogous to those of the peach, as well
as to those of the plum. The varieties differ considerably in the
shape of their leaves, which are either serrated or crenated, some-
times with ear-like appendages at their bases, and sometimes with
glands on the petioles. The flowers are generally alike, but are
small in the Masculine. The fruit varies much in size, shape, and
in having the suture little pronounced or absent; in the skin being
smooth, or downy as in the orange-apricot; and in the flesh cling-
ing to the stone, as in the last-mentioned kind, or in readily sepa-
rating from it, as in the Turkey-apricot. In all these differences we
see the closest analogy with the varieties of the peach and nectarine.
5
62 Journal of Horticulture,' Sept.
8th, 1863, p. 188.
63 Transact. Hort. Soc.,' vol. vi. p.
412.
64 Gardener's Chronicle, 1857, p.
216.
65 Journal of Hort. Soc.,' vol. ii. p.
283.
68 Alph. De Candolle, Géograph.
Bot., p. 879.
67 Transact. Hort. Soc.' (2nd series),
vol. i., 1835, p. 56. See also 'Cat. of
Fruit in Garden of Hort. Soc.,' 3rd
edit., 1842

CHAP. X.
415
APRICOT-PLUMS.
99 68
In the stone we have more important differences, and these in the
case of the plum have been esteemed of specific value: in some
apricots the stone is almost spherical, in others much flattened, being
either sharp in front or blunt at both ends, sometimes channelled
along the back, or with a sharp ridge along both margins. In the
Moor-park, and generally in the Hemskirke, the stone presents a
singular character in being perforated, with a bundle of fibres pass-
ing through the perforation from end to end. The most constant
and important character, according to Thompson, is whether the
kernel is bitter or sweet; yet in this respect we have a graduated
difference, for the kernel is very bitter in Shipley's apricot; in the
Hemskirke less better than in some other kinds: slightly bitter in
the Royal; and “sweet like a hazelaut” in the Breda, Angoumois,
and others. In the case of the almond, bitterness has been thought
by some high authorities to indicate specific difference.
In N. America the Roman apricot endures "cold and unfavourable
situations, where no other sort, except the Masculine, will succeed ;
and its blossoms bear quite a severe frost without injury. Ac-
cording to Mr. Rivers 69 seedling apricots deviate but little from the
character of their race: in France the Alberge is constantly repro-
duced from seed with but little variation. In Ladakh, according to
Moorcroft, 70 ten varieties of the apricot, very different from each
other, are cultivated, and all are raised from seed, excepting one,
which is budded.
Plums (Prunus insititia).–Formerly the sloe, P. spinosa, was
thought to be the parent of all our plums; but now this honour is
very commonly accorded to P. insititia or the bullace, which is
found wild in the Caucasus and N.-Western India, and is natural-
ised in England." It is not at all improbable, in accordance with
some observations made by Mr. Rivers, 72 that both these forms,
which some botanists rank as a single species, may be the parents
of our domesticated plums. Another supposed parent-form, the
P. domestica, is said to be found wild in the region of the Caucasus.
Godron remarks 73 that the cultivated varieties may be divided into
two main groups, which he supposes to be descended from two
68 Downing, The Fruits of America,'
1845, p. 157; with respect to the Alberge
apricot in France, see p. 153.
89 Gardener's Chronicle, 1863, p.
364.
70 Travels in the Himalayan Pro-
ces,' vol. i., 1841, p. 295.
71 See an excellent discussion on this
subject in Hewett C. Watson's Cybele
Britannica,' vol. iv. p. 80.
72 Gardener's Chronicle,' 1865, p. 27.
73 De l'Espèce,' tom. ii. p. 94. On
the parentage of our plums, see also
Alph. De Candolle, 'Géograph Bot.,' p.
878.
Also Targioni-Tozzetti, 'Journal
Hort. Soc.,' vol. ix. p. 164. Also Babing-
ton, Manual of Brit. Botany,' 1851, P.
87.

416
CHAP. X.
FRUITS.
aboriginal stocks; namely, those with oblong fruit and stones
pointed at both ends, having narrow separate petals and upright
branches; and those with rounded fruit, with stones blunt at both
ends, with rounded petals and spreading branches. From what we
know of the variability of the flowers in the peach and of the di
1
2
3
4
5
6
7
Fig. 43.-Plum Stones, of natural size, viewed laterally. 1. Bullace Plum. 2. Shrop-
shire Damson. 3. Blue Gage. 4. Orleans. 5. Elvas. 6. Denyer's Victoria. 7. Dia-
mond.
versified manner of growth in our various fruit-trees, it is difficult
to lay much weight on these latter characters. With respect to the
shape of the fruit, we have conclusive evidence that it is extremely
variable: Downing 74 gives outlines of the plums of two seedlings,
namely, the red and imperial gages, raised from the greengage;
and the fruit of both is more elongated than that of the greengage.
The latter has a very blunt broad stone, whereas the stone of the
imperial gage is "oval and pointed at both ends." These trees also
differ in their manner of growth : "the greengage is a very short-
jointed, slow-growing tree, of spreading and rather dwarfish habit;"
whilst its offspring, the imperial gage, “grows freely and rises
rapidly, and has long dark shoots." The famous Washington
74 "Fruits of America,' pp. 276, 278,
314, 284, 276, 310. Mr. Rivers raised
("Gard. Chron,' 1863, p. 27) from the
Prune-pêche, which bears large, round,
red plums on stout robust shoots, a seed-
ling which bears oval, smaller fruit on
shoots that are so slender as to be almost
pendulous.

CHAP. X.
417
PLUMS.
plum bears a globular fruit, but its offspring, the emerald drop, is
nearly as much elongated as the most elongated plum figured by
Downing, namely, Manning's prune. I have made a small collec-
tion of the stones of twenty-five kinds, and they graduate in shape
from the bluntest into the sharpest kinds. As characters derived
from seeds are generally of high systematic importance, I have
thought it worth while to give drawings of the most distinct kinds
in my small collection; and they may be seen to differ in a surpris-
ing manner in size, outline, thickness, prominence of the ridges, and
state of surface. It deserves notice that the shape of the stone is
not always strictly correlated with that of the fruit: thus the
Washington plum is spherical and depressed at the pole, with a
somewhat elongated stone, whilst the fruit of the Goliath is more
elongated, but the stone less so, than in the Washington. Again,
Denyer's Victoria and Goliath bear fruit closely resembling each
other, but their stones are widely different. On the other hand, the
Harvest and Black Margate plums are very dissimilar, yet include
closely similar stones.
The varieties of the plum are numerous, and differ greatly in
size, shape, quality, and colour, -being bright yellow, green, almost
white, blue, purple, or red. There are some curious varieties, such
as the double or Siamese, and the Stoneless plum : in the latter the
kernel lies in a roomy cavity surrounded only by the pulp. The
climate of North America appears to be singularly favourable for
the production of new and good varieties; Downing describes no
less than forty, seven of which of first-rate quality have been recent-
ly introduced into England.75 Varieties occasionally arise having
an innate adaptation for certain soils, almost as strongly pronounc-
ed as with natural species growing on the most distinct geological
formations; thus in America the imperial gage, differently from al-
most all other kinds, "is peculiarily fitted for dry light soils where
many sorts drop their fruit," whereas on rich heavy soils the fruit is
often insipid.76 My father could never succeed in making the
Wine-Sour yield even a moderate crop in a sandy orchard near
Shrewsbury, whilst in some parts of the same county and in its na-
tive Yorkshire it bears abundantly: one of my relations also re-
peatedly tried in vain to grow this variety in a sandy district in
Staffordshire.
Mr. Rivers has given a number of interesting facts, showing
75 Gardener's Chronicle,'1855, p. 726.
76 Downing's 'Fruit Trees,' p. 278.
77.Gardener's Chronicle,' 1863, p. 27.
Sageret, in his 'Pomologie Phys.,' p. 346,
enumerates five kinds which can be pro-
pagated in France by seed: see also
Downing's 'Fruit Trees of America,' p.
305, 312, &c.

418
CHAP. X.
FRUITS.
how truly many varieties can be propagated by seed. He sowed the
stones of twenty bushels of the greengage for the sake of raising
stocks, and closely observed the seedlings; "all had the smooth
shoots, the prominent buds, and the glossy leaves of the greengage,
but the greater number had smaller leaves and thorns." There are
two kinds of damson, one the Shropshire with downy shoots, and
the other the Kentish with smooth shoots, and these differ but
slightly in any other respect: Mr. Rivers sowed some bushels of the
Kentish damson, and all the seedlings had smooth shoots, but in
some the fruit was oval, in others round or roundish, and in a few
the fruit was small, and, except in being sweet, closely resembled
that of the wild sloe. Mr. Rivers gives several other striking in-
stances of inheritance: thus, he raised eighty thousand seedlings
from the common German Quetsche plum, and “not one could be
found varying in the least, in foliage or habit.” Similar facts were
observed with the Petite Mirabelle plum, yet this latter kind (as
well as the Quetsche) is known to have yielded some well-established
varieties; but, as Mr. Rivers remarks, they all belong to the same
group with the Mirabelle.
Cherries (Prunus cerasus, avium, &c).-Botanists believe that our
cultivated cherries are descended from one, two, four, or even more
wild stocks.78 That there must be at least two parent-species we
may infer from the sterility of twenty hybrids raised by Mr. Knight
from the morello fertilized by pollen of the Elton cherry; for these
hybrids produced in all only five cherries, and one alone of these con-
tained a seed." Mr. Thompson 80 has classified the varieties in an
apparently natural method in two main groups by characters taken
from the flowers, fruit, and leaves; but some varieties which stand
widely separate in this classification are quite fertile when crossed ;
thus Knight's Early Black cherry is the product of a cross between
two such kinds.
Mr. Knight states that seedling cherries are more variable than
those of any other fruit-tree.81 In the Catalogue of the Horticultural
Society for 1842, eighty varieties are enumerated. Some varieties
present singular characters: thus the flower of the Cluster cherry
includes as many as twelve pistils, of which the majority abort; and
they are said generally to produce from two to five or six cherries
aggregated together and borne on a single peduncle. In the Ratafia
79 Transact. Hort. Soc.' vol. v., 1824,
p. 295.
78 Compare Alph. De Candolle, 'Géo-
graph. Bot.,' p. 877; Bentham and Tar-
gioni-Tozzetti, in 'Hort. Journal,' vol.
ix. p. 163; Godron, 'De l'Espèce,' tom.
ii. p. 92.
80 Ibid., second series, vol. i., 1835,
p. 248.
81 Ibid., vol. ii. p. 138.

CHAP. X.
419
APPLES.
cherry several flower-peduncles arise from a common peduncle, up-
wards of an inch in length. The fruit of Gascoigne's Heart has its
apex produced into a globule or drop: that of the white Hungarian
Gean has almost transparent flesh. The Flemish cherry is "a very
odd-looking fruit,” much flattened at the summit and base, with the
latter deeply furrowed, and borne on a stout very short footstalk.
In the Kentish cherry the stone adheres so firmly to the footstalk,
that it can be drawn out of the flesh; and this renders the fruit
well fitted for drying. The Tobacco-leaved cherry, according to
Sageret and Thompson, produces gigantic leaves, more than a foot
and sometimes even eighteen inches in length, and half a foot in
breadth. The Weeping cherry, on the other hand, is valuable only
as an ornament, and, according to Downing, is “a charming little
tree with slender weeping branches, clothed with small almost
myrtle-like foliage.” There is also a peach-leaved variety.
Sageret describes a remarkable variety, le griottier de la Toussaint,
which bears at the same time, even as late as September, flowers
and fruit of all degrees of maturity. The fruit, which is of inferior
quality, is borne on long, very thin footstalks. But the extraordi-
nary statement is made that all the leaf-bearing shoots spring from
old flower-buds. Lastly, there is an important physiological dis-
tinction between those kinds of cherries which bear fruit on young
or on old wood; but Sageret positively asserts that a Bigarreau in
his garden bore fruit on wood of both ages.82
Apple (Pyrus malus).- The one source of doubt felt by botanists
with respect to the parentage of the apple is whether, besides P.
malus, two or three other closely allied wild forms, namely, P.
acerba and præcox or paradisiaca, do not deserve to be ranked as
distinct species. The P. præcox is supposed by some authors 83 to be
the parent of the dwarf paradise stock, which, owing to the fibrous
roots not penetrating deeply into the ground, is so largely used for
grafting ; but the paradise stock, it is asserted, 64 cannot be propa-
gated true by seed. The common wild crab varies considerably in
82 These several statements are taken
from the four following works, which
may, I believe, be trusted. Thompson,
in Hort. Transact.,' see above; Sage-
ret's 'Pomologie Phys.,' 1830, pp. 358,
364, 367, 379; Catalogue of the Fruit in
the Garden of Hort. Soc.,' 1842, pp. 57,
60; Downing, The Fruits of America,
1845, pp. 189, 195, 200.
83 Mr. Lowe states in his Flora of
Madeira' (quoted in Gard. Chron.,
1862, p. 215) that the P. malus, with its
nearly sessile fruit, ranges farther south
than the long-stalked P. acerba, which
is entirely absent in Madeira, the Cana-
ries, and apparently in Portugal. This
fact supports the belief that these two
forms deserve to be called species. But
the characters separating them are of
slight importance, and of a kind known
to vary in other cultivated fruit-trees.
84 See 'Journ. of Hort. Tour.,' by De-
putation of the Caledonian Hort. Soc.,
1823, p. 459.

420
CHAF, X.
FRUITS.
England; but many of the varieties are believed to be escaped seed-
lings. Every one knows the great difference in the manner of
growth, in the foliage, flowers, and especially in the fruit, between
the almost innumerable varieties of the apple. The pips or seeds
(as I know by comparison) likewise differ considerably in shape, size,
and colour. The fruit is adapted for eating or for cooking in differ-
ent ways, and keeps for only a few weeks or for nearly two years.
Some few kinds have the fruit covered with a powdery secretion,
called bloom, like that on plums; and "it is extremely remarkable
that this occurs almost exclusively among varieties cultivated in
Russia.?86 Another Russian apple, the white Astracan, possesses the
singular property of becoming transparent, when ripe, like some
sorts of crabs. The api étoilé has five prominent ridges, hence its
name; the api noir is nearly black: the twin cluster pippin often
bears fruit joined in pairs.87 The trees of the several sorts differ
greatly in their periods of leafing and flowering; in my orchard the
Court Pendu Plat produces its leaves so late, that during several
springs I have thought it dead. The Tiffin apple scarcely bears a
leaf when in full bloom; the Cornish crab, on the other hand, bears
so many leaves at this period that the flowers can hardly be seen.
In some kinds the fruit ripens in midsummer; in others, late in the
autumn. These several differences in leafing, flowering, and fruit-
ing, are not at all necessarily correlated ; for, as Andrew Knight has
remarked, no one can judge from the early flowering of a new
seedling, or from the early shedding or change of colour of the
leaves, whether it will mature its fruit early in the season.
The varieties differ greatly in constitution. It is notorious that
our summers are not hot enough for the Newtown Pippin,90 which
is the glory of the orchards near New York; and so it is with seve-
ral varieties which we have imported from the Continent. On the
other hand, our Court of Wick succeeds well under the severe cli-
mate of Canada. The Calville rouge de Micoud occasionally bears
two crops during the same year. The Burr Knot is covered with
small excrescences, which emit roots so readily that a branch with
blossom-buds may be stuck in the ground, and will root and bear
85 H. O. Watson, Cybele Britannica,'
vol. i. p. 334.
86 Loudon's "Gardener's Mag.,' vol.
vi., 1830, p. 83.
vol. iv., 1828, p. 112.
89 The Culture of the Apple,' p. 43.
Van Mons makes the same remark on
Arbres Fruitiers,' tom. ii.,
1836, p. 414.
the pear,
87 See Catalogue of Fruit in Garden
of Hort. Soc.,' 1812, and Downing's
American Fruit Trees.'
88 Loudon's Gardener's Magazine,
90 Lindley's 'Horticulture,' p. 116.
See also Knight on the Apple-Tree, in
Transact. of Hort. Soc.,' vol. vi. p. 229.

CHAP. X.
421
APPLES.
a few fruit even during the first year.91 Mr. Rivers has recently de-
scribed some seedlings valuable from their roots running near the
surface. One of these seedlings was remarkable from its extremely
dwarfed size, "forming itself into a bush only a few inches in
height." Many varieties are particularly liable to canker in cer-
tain soils. But perhaps the strangest constitutional peculiarity is
that the Winter Majetin is not attacked by the mealy bug or coccus;
Lindley 93 states that in an orchard in Norfolk infested with these
'insects the Majetin was quite free, though the stock on which it
was grafted was affected; Knight makes a similar statement with
respect to a cider apple, and adds that he only once saw these in-
sects just above the stock, but that three days afterwards they en-
tirely disappeared; this apple, however, was raised from a cross be-
tween the Golden Hervey and the Siberian Crab; and the latter, I
believe, is considered by some authors as specifically distinct.
The famous St. Valery apple must not be passed over; the flower
has a double calyx with ten divisions, and fourteen styles surmounted
by conspicuous oblique stigmas, but is destitute of stamens or corolla.
The fruit is constricted round the middle, and is formed of five seed-
cells, surmounted by nine other cells.94 Not being provided with
stamens, the tree requires artificial fertilisation; and the girls of
St. Valery annually go to "faire ses pommes,” each marking her
own fruit with a ribbon ; and as different pollen is used, the fruit
differs, and we here have an instance of the direct action of foreign
pollen on the mother-plant. These monstrous apples include, as we
have seen, fourteen seed-cells; the pigeon-apple, 95 on the other hand,
has only four, instead of, as with all common apples, five cells; and
this certainly is a remarkable difference.
In the catalogue of apples published in 1842 by the Horticultural
Society, 897 varieties are enumerated; but the differences between
most of them are of comparatively little interest, as they are not
strictly inherited. No one can raise, for instance, from the seed of
the Ribston Pippin, a tree of the same kind; and it is said that the
“Sister Ribston Pippin" was a white, semi-transparent, sour-fleshed
apple, or rather large crab.96 Yet it is a mistake to suppose that
with most varieties the characters are not to a certain extent in-
91 Transact. Hort. Soc.,' vol. i., 1812,
p. 120.
92 Journal of Horticulture,' March
13th, 1866, p. 194.
93 Transact. Hort. Soc.,' vol. iv. p.
68. For Knight's case, see vol. vi. p.
547. When the coccus first appeared in
this country, it is said (vol. ii. p. 163)
that it was more injurious to crab-stocks
than to the apples grafted on them.
94 Mém. de la Soc. Linn. de Paris,
tom. iii., 1825, p. 164; and Seringe,
Bulletin Bot.,' 1830, p. 117.
95 Gardener's Chronicle,' 1849, p. 24.
96 R. Thompson, in Gardener's
Chron.,' 1850, p. 788.

422
CHAP. X.
FRUITS.
herited. In two lots of seedlings raised from two well-marked
kinds, many worthless, crab-like seedlings will appear, but it is now
known that the two lots not only usually differ from each other, but
resemble to a certain extent their parents. We see this indeed in
the several sub-groups of Russetts, Sweetings, Codlins, Pearmains,
Reinettes, &c.,97 which are all believed, and many are known, to be
descended from other varieties bearing the same names.
Pears (Pyrus communis).-I need say little on this fruit, which
varies much in the wild state, and to an extraordinary degree when
cultivated, in its fruit, flowers, and foliage. One of the most cele-
brated botanists in Europe, M. Decaisne, has carefully studied the
many varieties; 98 although he formerly believed that they were de-
rived from more than one species, he is now convinced that all be-
Iong to one. He has arrived at this conclusion from finding in the
several varieties a perfect gradation between the most extreme
characters; so perfect is this gradation that he maintains it to be
impossible to classify the varieties by any natural method. M. De-
caisne raised many seedlings from four distinct kinds, and has care-
fully recorded the variations in each. Notwithstanding this ex-
treme degree of variability, it is now positively known that many
kinds reproduce by seed the leading characters of their race.99
Straroberries (Fragaria).—This fruit is remarkable on account of
the number of species which have been cultivated, and from their
rapid improvement within the last fifty or sixty years. Let any
one compare the fruit of one of the largest varieties exhibited at
our Shows with that of the wild wood strawberry, or, which will
be a fairer comparison, with the somewhat larger fruit of the wild
American Virginian Strawberry, and he will see what prodigies
horticulture has effected.100 The number of varieties has likewise
increased in a surprisingly rapid manner. Only three kinds were
known in France, in 1746, where this fruit was early cultivated. In
1766 five species had been introduced, the same which are now cul-
97 Sageret, Pomologie Physiologique,
1830, p. 263. Downing's 'Fruit Trees,
pp. 130, 134, 139, &c. Loudon's Gar-
dener's Mag.,' vol viii. p. 317. Alexis
Jordan, 'De l'Origine des diverses Vari.
étés,' in 'Mém. de l'Acad. Imp. de
Lyon,' tom. ii., 1852, pp. 95, 114, Gar-
dener's Chronicle,' 1850, pp. 774, 788.
98 Comptes Rendus,' July 6th, 1863.
99 Gardener's Chronicle,' 1856, p.
804; 1857, p. 820; 1862, p. 1195.
100 Most of the largest cultivated
strawberries are the descendants of F.
grandiflora or Chiloensis, and I have
seen no account of these forms in their
wild state. Methuen's Scarlet (Down-
ing, Fruits,' p. 527) has immense fruit
of the largest size," and belongs to the
section descended from F. Virginiana;
and the fruit of this species, as I hear
from Prof. A. Gray, is only a little larger
than that of F. vesco, or our common
wood strawberry.

CHAP. X.
423
STRAWBERRIES.
tivated, but only five varieties of Fragaria vesca, with some sub-va-
rieties, had been produced. At the present day the varieties of the
several species are almost innumerable. The species consist of,
firstly, the wood or Alpine cultivated strawberries, descended from
F. vesca, a native of Europe and of North America. There are eight
wild European varieties, as ranked by Duchesne, of F. vesca, but
several of these are considered species by some botanists. Secondly,
the green strawberries, descended from the European F. collina,
and little cultivated in England. Thirdly, the Hautbois, from the
European F. elatior. Fourthly, the Scarlets, descended from F.
Virginiana, a native of the whole breadth of North America.
Fifthly, the Chili, descended from F. Chiloensis, an inhabitant of
the west coast of the temperate parts both of North and South
America. Lastly, the Pines or Carolinas (including the old Blacks),
which have been ranked by most authors under the name of F.
grandiflora as a distinct species, said to inhabit Surinam; but this
is a manifest error. This form is considered by the highest autho-
rity, M. Gay, to be merely a strongly marked race of F. Chiloensis.101
These five or six forms have been ranked by most botanists as spe-
cifically distinct; but this may be doubted, for Andrew Knight,102
who raised no less than 400 crossed strawberries, asserts that the F.
Virginiana, Chiloensis and grandiflora "may be made to breed to-
gether indiscriminately," and he found, in accordance with the prin-
ciple of analogous variation, “that similar varieties could be obtain-
ed from the seeds of any one of them."
Since Knight's time there is abundant and additional evidence 103
of the extent to which the American forms spontaneously cross.
We owe indeed to such crosses most of our choicest existing vari-
eties. Knight did not succeed in crossing the European wood-
strawberry with the American Scarlet or with the Hautbois. Mr.
Williams, of Pitmaston, however, succeeded; but the hybrid off-
spring from the Hautbois, though fruiting well, never produced
seed, with the exception of a single one, which reproduced the pa-
rent hybrid form.104 Major R. Trevor Clarke informs me that he
crossed two members of the Pine class (Myatt's B. Queen and Keen's
Seedling), with the wood and hautbois, and that in each case he
raised only a single seedling; one of these fruited, but was almost
101 Le Fraisier,' par le Comte L. de
Lambertye, 1864, p. 50.
102 Transact. Hort. Soc.,' vol. iii.
1820, p. 207.
103 See an account. by Prof. Decaisne,
and by others in Gardener's Chronicle,
1862, p. 335, and 1858, p. 172; and Mr.
Barnet's paper in 'Hort. Soc. Transact.,'
vol. vi., 1826, p. 170.
104 Transact. Hort. Soc.,' vol. V.,
1824, p. 294.
9

424
CHAP. X.
FRUITS.
barren. Mr. W. Smith, of York, has raised similar hybrids with
equally poor success. 105 We thus see 106 that the European and
American species can with some difficulty be crossed ; but it is
improbable that hybrids sufficiently fertile to be worth cultivation
will ever be thus produced. This fact is surprising, as these forms
structurally are not widely distinct, and are sometimes connected in
the districts where they grow wild, as I hear from Professor Asa
Gray, by puzzling intermediate forms.
The energetic culture of the strawberry is of recent date, and the
cultivated varieties can in most cases still be classed under some one
of the above five native stocks. As the American strawberries cross
so freely and spontaneously, we can hardly doubt that they will
ultimately become inextricably confused. We find, indeed, that
horticulturists at present disagree under which class to rank some
few of the varieties; and a writer in the 'Bon Jardinier' of 1840
remarks that formerly it was possible to class all of them under
some one species, but that now this is quite impossible with the
American forms, the new English varieties having completely filled
up the gaps between them. 107 The blending together of two or
more aboriginal forms, which there is every reason to believe has
occurred with some of our anciently cultivated productions, we now
see actually occurring with our strawberries.
The cultivated species offer some variations worth notice. The
Black Prince, a seedling from Keen's Imperial (this latter being a
seedling of a very white strawberry, the white Carolina), is remark-
able from " its peculiar dark and polished surface, and from present-
ing an appearance entirely unlike that of any other kind." 108 Al-
though the fruit in the different varieties differs so greatly in form,
size, colour, and quality, the so-called seed (which corresponds with
the whole fruit in the plum), with the exception of being more or
less deeply imbedded in the pulp, is, according to De Jonghe,109 ab-
solutely the same in all, and this no doubt may be accounted for
by the seed being of no value, and consequently not having been
subjected to selection. The strawberry is properly three-leaved, but
in 1761 Duchesne raised a single-leaved variety of the European
wood-strawberry, which Linnæus doubtfully raised to the rank of a
species. Seedlings of this variety, like those of most varieties not
fixed by long-continued selection, often revert to the ordinary form,
105 'Journal of Horticulture, Dec.
S0th, 1862, p. 779. Sec also Mr. Prince
to the same effect, idem, 1863, p. 418.
106 For additional evidence see 'Jour-
nal of Horticulture, Dec. 9th, 1862, p.
21.
107 'Le Fraisier, par le Comte L. de
Lambertye, pp. 221, 230.
108 Transact. Hort. Soc.,' vol. vi., p.
200.
109 Gardener's Chron.,'1858, p. 173.

CEAP. X.
425
STRAWBERRIES.
or present intermediate states. 110 A variety raised by Mr. Myatt, 111
apparently belonging to one of the American forms, presents a va-
riation of an opposite nature, for it has five leaves ; Godron and
Lambertye also mention a five-leaved variety of F. collina.
The Red Bush Alpine strawberry (one of the F. vesca section does
not produce stolons or runners, and this remarkable deviation of
structure is reproduced truly by seed. Another sub-variety, the
White Bush Alpine, is similarly characterised, but when propagated
by seed it often degenerates and produces plants with runners. 112
A strawberry of the American Pine section is also said to make but
few runners. 113
Much has been written on the sexes of strawberries; the true
Hautbois properly bears the male and female organs on separate
plants, 114 and was consequently named by Duchesne dioica; but it
frequently produces hermaphrodites; and Lindley, 115 by propagat-
ing such plants by runners, at the same time destroying the males,
soon raised a self-prolific stock. The other species often 'show a
tendency towards an imperfect separation of the sexes, as I have
noticed with plants forced in a hot-house. Several English varie-
ties, which in this country are free from any such tendency, when
cultivated in rich soils under the climate of North America com-
monly produce plants with separate sexes. Thus a whole acre of
Keen's Seedlings in the United States has been observed to be al-
most sterile from the absence of male flowers; but the more general
rule is, that the male plants overrun the females. Some members
of the Cincinnati Horticultural Society, especially appointed to in-
vestigate this subject, report that "few varieties have the flowers
perfect in both sexual organs,” &c. The most successful cultivators
in Ohio, plant for every seven rows of “pistillata," or female plants,
one row of hermaphrodites, which afford pollen for both kinds; but
the hermaphrodites, owing to their expenditure in the production of
pollen, bear less fruit than the female plants.
The varieties differ in constitution. Some of our best English
kinds, such as Keen's Seedlings, are too tender for certain parts of
North America, where other English and many American varieties
succeed perfectly. That splendid fruit, the British Queen, can be
cultivated but in few places either in England or France; but this
116
p. 210.
110 Godron, 'De l'Espèce,' tom. i. p.
161.
111 Gardener's Chron.,'1851, p. 440.
112 F. Gloede, in Gardener's Chron.,'
1862, p. 1053
113 Downing's 'Fruits,' p. 532.
114 Barnet, in 'Hort. Transact.,' vol. vi.
115 Gardener's Chron.,' 1847, p. 539.
116 For the several statements with
respect to the American strawberries, see
Downing, "Fruits,' p. 524; Gardener's
Chronicle,' 1843, p. 158; 1847, p. 539;
1861, p. 717.

426
CHAP, X,
FRUITS.
") 117
apparently depends more on the nature of the soil than on the cli-
mate: a famous gardener says that "no mortal could grow the
British Queen at Shrubland Park unless the whole nature of the soil
was altered.' La Constantina is one of the hardiest kinds, and
can withstand Russian winters, but is easily burnt by the sun, so
that it will not succeed in certain soils either in England or the
United States. 118 The Filbert Pine Strawberry “requires more
water than any other variety; and if the plants once suffer from
drought, they will do little or no good afterwards.” 119 Cuthill's
Black Prince Strawberry evinces a singular tendency to mildew :
no less than six cases have been recorded of this variety suffering
severely, whilst other varieties growing close by, and treated in ex-
actly the same manner, were not at all infested by this fungus. 120
The time of maturity differs much in the different varieties; some
belonging to the wood or alpine section produce a succession of crops
throughout the summer.
Gooseberry (Ribes grossularia).-No one, I believe, has hitherto
doubted that all the cultivated kinds are sprung from the wild plant
bearing this name, which is common in Central and Northern Eu-
rope; therefore it will be desirable briefly to specify all the points,
though not very important, which have varied. If it be admitted
that these differences are due to culture, authors perhaps will not
be so ready to assume the existence of a large number of unknown
wild parent-stocks for our other cultivated plants. The gooseberry
is not alluded to by writers of the classical period. Turner men-
tions it in 1573, and Parkinson, in 1629, specifies eight varieties;
the Catalogue of the Horticultural Society for 1842 gives 149 varie-
ties, and the lists of the Lancashire nurserymen are said to include
above 300 names.121 In the 'Gooseberry Grower's Register for 1862'
I find that 243 distinct varieties have at various periods won prizes ;
so that a vast number must have been exhibited. No doubt the
difference between many of the varieties is very small; but Mr.
Thompson in classifying the fruit for the Horticultural Society found
less confusion in the nomenclature of the gooseberry than of any
other fruit, and he attributes this " to the great interest which the
prize-growers have taken in detecting sorts with wrong names," and
117 Mr. D. Beaton, in Cottage Gar-
dener,' 1860, p. 86, See also Cottage
Gardener,' 1855, p. 88, and many other
authorities. For the Continent, see F.
Gloede, in 'Gardener's Chronicle,' 1862,
119 Mr. H. Doubleday in Gardener's
Chron.,' 1862, p. 1101.
120 Gardener's Chronicle,' 1854, p.
254.
121 Loudon's Encyclop. of Gardening,'
p. 930; and Alph, de Candolle, 'Géo-
graph. Bot.,' p. 910.
p. 1053.
118 Rev. W. F. Radclyffe, in Journal
of Hort.,' March 14, 1865, p. 207.

CHAP. X.
427
THE GOOSEBERRY.
this shows that all the kinds, numerous as they are, can be recog-
nised with certainty.
The bushes differ in their manner of growth, being erect, or spread-
ing, or pendulous. The periods of leafing and flowering differ both ab-
solutely and relatively to each other; thus the Whitesmith produces
early flowers, which from not being protected by the foliage, as it is
believed, continually fail to produce fruit.122 The leaves vary in
size, tint, and in depth of lobes; they are smooth, downy, or hairy
on the upper surface. The branches are more or less downy or
spinose; "the Hedgehog has probably derived its name from the
singular bristly condition of its shoots and fruit." The branches of
the wild gooseberry, I may remark, are smooth, with the exception
of thorns at the bases of the buds. The thorns themselves are
either very small, few and single, or very large and triple; they
are sometimes reflexed and much dilated at their bases. In the dif-
ferent varieties the fruit varies in abundance, in the period of matu-
rity, in hanging until shrivelled, and greatly in size," some sorts
having their fruit large during a very early period of growth,
whilst others are small until nearly ripe.” The fruit varies also
much in colour, being red, yellow, green, and white—the pulp of
one dark-red gooseberry being tinged with yellow; in flavour; in
being smooth or downy,-few, however, of the Red gooseberries,
whilst many of the so-called Whites, are downy; or in being so
spinose that one kind is called Henderson's Porcupine. Two kinds
acquire when mature a powdery bloom on their fruit. The fruit
varies in the thickness and veining of the skin, and, lastly, in shape,
being spherical, oblong, oval, or obovate.123
I cultivated fifty-four varieties, and, considering how greatly the
fruit differs, it was curious how closely similar the flowers were
in all these kinds. In only a few I detected a trace of difference in
the size or colour of the corolla. The calyx differed in a rather
greater degree, for in some kinds it was much redder than in others;
and in one smooth white gooseberry it was unusually red. The
calyx also differed in the basal part being smooth or woolly, or
covered with glandular hairs. It deserves notice, as being contrary
to what might have been expected from the law of correlation, that
a smooth red gooseberry had a remarkably hairy calyx. The
flowers of the Sportsman are furnished with very large coloured
bracteæ; and this is the most singular deviation of structure which
122 Loudon's Gardener's Magazine,
vol. iv. 1828, p. 112.
123 The fullest account of the goose-
berry is given by Mr. Thompson in
Transact. Hort. Soc.,' vol. i., 2nd series,
1835, p. 218, from which most of the fore-
going facts are given.

429
CHAP. X.
FRUITS.
I have observed. These same flowers also varied much in tho
number of the petals, and occasionally in the number of the stamens
and pistils; so that they were semi-monstrous in structure, yet they
produced plenty of fruit. Mr. Thompson remarks that in the Pas-
time gooseberry “extra bracts are often attached to the sides of the
“fruit."124
The most interesting point in the history of the gooseberry is the
steady increase in the size of the fruit. Manchester is the metro-
polis of the fanciers, and prizes from five shillings to five or ten
pounds are yearly given for the heaviest fruit. The 'Gooseberry
Grower's Register' is published annually; the earliest known copy
is dated 1786, but it is certain that meetings for the adjudication of
prizes were held some years previously. 125. The 'Register' for 1845
gives an account of 171 Gooseberry Shows, held in different places
during that year; and this fact shows on how large a scale the cul-
ture has been carried on. The fruit of the wild gooseberry is said 126
to weigh about a quarter of an ounce or 5 dwts., that is, 120 grs. ;
about the year 1786 gooseberries were exhibited weighing 10 dwts.,
so that the weight was then doubled ; in 1817 26 dwts. 17 grs. was
attained; there was no advance till 1825, when 31 dwts. 16 grs. was
reached; in 1830 “ Teazer” weighed 32 dwts. 13 grs.; in 1841
“Wonderful” weighed 32 dwts. 16 grs. ; in 1844 "London" weighed
35 dwts. 12 grs., and in the following year 36 dwts. 16 grs.; and in
1852 in Staffordshire the fruit of this same variety reached the
astonishing weight of 37 dwts. 7 grs., or 895 grs. ; that is, be-
tween seven and eight times the weight of the wild fruit. I find
that a small apple, 64 inches in circumference, has exactly this same
weight. The "London" gooseberry (which in 1862 had altogether
gained 343 prizes) has, up to the present year of 1864, never reached
a greater weight than that attained in 1852. Perhaps the fruit of
the gooseberry has now reached the greatest possible weight, unless
in the course of time some quite new and distinct variety shall
arise.
This gradual, and on the whole steady increase of weight from
the latter part of the last century to the year 1852, is probably in
large part due to improved methods of cultivation, for extreme care
is now taken; the branches and roots are trained, composts are
127
127
6
194 Catalogue of Fruits of Hort. Soc.
Garden,' 3rd edit. 1842.
125 Mr. Clarkson, of Manchester, on
the Culture of the Gooseberry, in Lou-
don's Gardener's Magazine,' vol. iv.
1828, p. 452.
126 Downing's Fruits of Amer.,' p. 213.
Gardener's Chronicle,' 1944, p.
811, where a table is given; and 1845, p.
819. For the extreme weights gained,
see 'Journal of Horticulture,' July 26,
1964, p. 61.

CHAP. X.
429
WALNUTS.
128
made, the soil is mulched, and only a few berries are left on each
bush; but the increase no doubt is in main part due to the con-
tinued selection of seedlings which have been found to be more and
more capable of yielding such extraordinary fruit. Assuredly the
“Highwayman " in 1817 could not have produced fruit like that of
the “Roaring Lion” in 1825; nor could the “Roaring Lion,"
though it was grown by many persons in many places, gain the
supreme triumph achieved in 1852 by the “ London ” Gooseberry.
Walnut (Juglans regia).--This tree and the common nut belong
to a widely different order from the foregoing fruits, and are there-
fore here noticed. The walnut grows wild in the Caucasus and
Himalaya, where Dr. Hooker 120 found the fruit of full size, but "as
hard as a hickory-nut." In England the walnut presents consider-
able differences, in the shape and size of the fruit, in the thickness
of the husk, and in the thinness of the shell; this latter quality
has given rise to a variety called the thin-shelled, which is valuable,
but suffers from the attacks of tom-tits.130 The degree to which
the kernel fills the shell varies much. In France there is a variety
called the Grape or cluster-walnut, in which the nuts grow in
“bunches of ten, fifteen, or even twenty together." There is another
variety which bears on the same tree differently shaped leaves, like
the heterophyllous hornbeam; this tree is also remarkable from
having pendulous branches, and bearing elongated, large, thin-
shelled nuts.131 M. Cardan has minutely described 132 some singu-
lar physiological peculiarities in the June-leafing variety, which
produces its leaves and flowers four or five weeks later, and retains
its leaves and fruit in the autumn much longer, than the common
varieties; but in August is in exactly the same state with them.
These constitutional peculiarities are strictly inherited. Lastly,
walnut-trees, which are properly monoicous, sometimes entirely fail
to produce male flowers. 133
Nuts (Corylus avellana).—Most botanists rank all the varieties
under the same species, the common wild nut.134 The husk, or in-
volucre, differs greatly, being extremely short in Barr's Spanish,
and extremely long in filberts, in which it is contracted so as to
128 Mr. Saul, of Lancaster, in Loudon's "Gardener's Mag.,' 1829, vol. v. p. 202.
Gardener's Mag., vol. iii. 1828, p. 421; 132 Quoted in Gardener's Chronicle,'
and vol. x. 1834, p. 42.
1819, p. 101.
129 'Himalayan Journals,' 1854, vol. 133 Gardener's Chronicle,' 1847, pp.
ii. p. 334. Moorcroft (Travels,' vol. ii. 541 and 558.
p. 146) describes four varieties cultivated 134 The following details are taken
in Kashmir.
from the Catalogue of Fruits, 1842, in
Gardener's Chronicle,' 1850, p. Garden of Hort. Soc., p. 103 ; and from
723.
Loudon's Encyclop. of Gardening,' p.
131 Paper translated in Loudon's 943.
130 4

430
CHAP. X.
CUCURBITACEOUS PLANTS.
prevent the nut falling out. This kind of husk also protects the
nut from birds, for titmice (Parus) have been observed 135 to pass
over filberts, and attack cobs and common nuts growing in the
same orchard. In the purple-filbert the husk is purple, and in
the frizzled-filbert it is curiously laciniated; in the red-filbert
the pellicle of the kernel is red. The shell is thick in some va-
rieties, but is thin in Cosford's-nut, and in one variety is of a blu-
ish colour. The nut itself differs much in size and shape, being
ovate and compressed in filberts, nearly round and of great size in
cobs and Spanish nuts, oblong and longitudinally striated in Cos-
ford's, and obtusely four-sided in the Downton Square nut.
Cucurbitaceous plants. These plants have been for a long period
the opprobrium of botanists; numerous varieties have been ranked
as species, and, what happens more rarely, forms which now must
be considered as species have been classed as varieties. Owing to
the admirable experimental researches of a distinguished botanist,
M. Naudin,136 a flood of light has recently been thrown on this group
of plants. M. Naudin, during many years, observed and experi-
mented on above 1200 living specimens, collected from all quarters
of the world. Six species are now recognised in the genus Cucur-
bita ; but three alone have been cultivated and concern us, namely,
C. maxima and pepo, which include all pumpkins, gourds, squashes,
and vegetable marrow, and C. moschata, the water-melon. These
three species are not known in a wild state; but Asa Gray 137 gives
good reason for believing that some pumpkins are natives of N.
America.
These three species are closely allied, and have the same general
habit, but their innumerable varieties can always be distinguished,
according to Naudin, by certain almost fixed characters; and what
is still more important, when crossed they yield no seed, or only
sterile seed; whilst the varieties spontaneously intercross with the
utmost freedom. Naudin insists strongly (p. 15), that, though these
three species have varied greatly in many characters, yet it has
been in so closely an analogous manner that the varieties can be
arranged in almost parallel series, as we have seen with the forms
of wheat, with the two main races of the peach, and in other cases.
Though some of the varieties are inconstant in character, yet others,
when grown separately under uniform conditions of life, are, as
Naudin repeatedly (pp. 6, 16, 35) urges, “douées d'une stabilité
presque comparable à celle des espèces les mieux caractérisées.”
195 Gardener's Chron., '1860, p. 956.
136 Annales des Sc. Nat. Bot.,' 4th
series, vol. vi. 1856, p. 5.
137 "American Journ. of Science,' 2nd
ser. vol. xxiv. 1857, p. 442.

CHAP. X.
431
CUCURBITACEOUS PLANTS.
One variety, l'Orangin (pp. 43, 63), has such prepotency in trans-
mitting its character that when crossed with other varieties a vast
majority of the seedlings come true. Naudin, referring (p. 47) to
C. pepo, says that its races “ne diffèrent des espèces véritables qu'en
ce qu'elles peuvent s'allier les unes aux autres par voie d'hybridité,
sans que leur descendance perde la faculté de se perpétuer.” If we
were to trust to external differences alone, and give up the test of
sterility, a multitude of species would have to be formed out of the
varieties of these three species of Cucurbita. Many naturalists at
the present day lay far too little stress, in my opinion, on the test
of sterility; yet it is not improbable that distinct species of plants
after a long course of cultivation and variation may have their
mutual sterility eliminated, as we have every reason to believe has
occurred with domesticated animals. Nor, in the case of plants
under cultivation, should we be justified in assuming that varieties
never acquire a slight degree of mutual sterility, as we shall more
fully see in a future chapter when certain facts are given on the
high authority of Gärtner and Kölreuter.138
The forms of C. pepo are classed by Naudin under seven sections,
each including subordinate varieties. He considers this plant as
probably the most variable in the world. The fruit of one variety
(pp. 33, 46) exceeds in volume that of another by more than two
thousand fold! When the fruit is of very large size, the number
produced is few (p. 45); when of small size, many are produced.
No less astonishing (p. 33) is the variation in the shape of the fruit;
the typical form apparently is egg-like, but this becomes either
drawn out into a cylinder, or shortened into a flat disc. We have
also an almost infinite diversity in the colour and state of surface of
the fruit, in the hardness both of the shell and of the flesh, and in
the taste of the flesh, which is either extremely sweet, farinaceous,
or slightly bitter. The seeds also differ in a slight degree in shape,
and wonderfully in size (p. 34), namely, from six or seven to more
than twenty-five millimètres in length.
In the varieties which grow upright or do not run and climb, the
tendrils, though useless (p. 31), are either present or are represented
by various semi-monstrous organs, or are quite absent. The ten-
drils are even absent in some running varieties in which the stems
are much elongated. It is a singular fact that (p. 31), in all the
varieties with dwarfed stems, the leaves closely resemble each other
in shape.
198 Gärtner, 'Bastarderzeugung,'1849,
s. 87, and s. 169 with respect to Maize;
on Verbascum, idem, ss. 92 and 181 ;
also his 'Kenntniss der Befruchtung,'
s. 137. With respect to Nicotiana, see
Kölreuter, Zweite Forts.,' 1764, s. 53
though this is a somewhat different
case.

432
CHAP, X
CUCURBITACEOUS PLANTS.
Those naturalists who believe in the immutability of species often
maintain that, even in the most variable forms, the characters
which they consider of specific value are unchangeable. To give
an example from a conscientious writer, 139 who, relying on the
labours of M. Naudin and referring to the species of Cucurbita,
says, "au milieu de toutes les variations du fruit, les tiges, les
feuilles, les calices, les corolles, les étamines restent invariables dans
chacune d'elles." Yet M. Naudin in describing Cucurbita pepo
(p. 30) says, “Ici, d'ailleurs, ce ne sont pas seulement les fruits qui
varient, c'est aussi le feuillage et tout le port de la plante. Néan-
moins, je crois qu'on la distinguera toujours facilement des deux
autres espèces, si l'on veut ne pas perdre de vue les caractères dif-
férentiels que je m'efforce de faire ressortir. Ces caractères sont
quelquefois peu marqués : il arrive même que plusieurs d'entre eux
s'effacent presque entièrement, mais il en reste toujours quelques-
uns qui remettent l'observateur sur la voie.” Now let it be noted
what a difference, with regard to the immutability of the so-called
specific characters, this paragraph produces on the mind, from that
above quoted from M. Godron.
I will add another remark : naturalists continually assert that no
important organ varies; but in saying this they unconsciously argue
in a vicious circle; for if an organ, let it be what it may, is highly
variable, it is regarded as unimportant, and under a systematic point
of view this is quite correct. But as long as constancy is thus taken
as the criterion of importance, it will indeed be long before an im-
portant organ can be shown to be inconstant. The enlarged form
of the stigmas, and their sessile position on the summit of the ovary,
must be considered as important characters, and were used by Gas-
parini to separate certain pumpkins as a distinct genus; but Naudin
says (p. 20) these parts have no constancy, and in the flowers of the
Turban varieties of C. maxima they sometimes resume their ordi-
nary structure. Again in C. maxima, the carpels (p. 19) which form
the Turban project even as much as two-thirds of their length out
of the receptacle, and this latter part is thus reduced to a sort of
platform ; but this remarkable structure occurs only in certain va-
rieties, and graduates into the common form in which the carpels
are almost entirely enveloped within the receptacle. In C. moschata
the ovarium (p. 50) varies greatly in shape, being oval, nearly sphe-
rical, or cylindrical, more or less swollen in the upper part, or con-
stricted round the middle, and either straight or curved. When the
ovarium is short and oval the interior structure does not differ from
189 De l'Espèce;' par M. Godron, tom. ii. p. 64.

CHAP. X.
433
CUCURBITACLOUS PLANTS.
that of C. maxima and pepo, but when it is elongated the carpels oc-
cupy only the terminal and swollen portion. I may add that in one
variety of the cucumber (Cucumis sativus) the fruit regularly con
tains five carpels instead of three.140 I presume that it will not be
disputed that we here have instances of great variability in organs
of the highest physiological importance, and with most plants of the
highest classificatory importance.
Sageret 141 and Naudin found that the cucumber (C. sativus) could
not be crossed with any other species of the genus; therefore no
doubt it is specifically distinct from the melon. This will appear to
most persons a superfluous statement; yet we hear from Naudin 142
that there is a race of melons, in which the fruit is so like that of
the cucumber, "both externally and internally, that it is hardly
possible to distinguish the one from the other except by the leaves."
The varieties of the melon seem to be endless, for Naudin after six
years' study has not come to the end of them: he divides them into
ten sections, including numerous sub-varieties which all intercross
with perfect ease. 143 Of the forms considered by Naudin to be va-
rieties, botanists have made thirty distinct species ! "and they had
not the slightest acquaintance with the multitude of new forms
which have appeared since their time." Nor is the creation of so
many species at all surprising when we consider how strictly their
characters are transmitted by seed, and how wonderfully they differ
in appearance: "Mira est quidem foliorum et habitus diversitas,
sed multo magis fructuum," says Naudin. The fruit is the valuable
part, and this, in accordance with the common rule, is the most mo-
dified part. Some melons are only as large as small plums, others
weigh as much as sixty-six pounds. One variety has a scarlet
fruit! Another is not more than an inch in diameter, but some-
times more than a yard in length, “twisting about in all directions
like a serpent.” It is a singular fact that in this latter variety many
parts of the plant, namely, the stems, the footstalks of the female
flowers, the middle lobe of the leaves, and especially the ovarium,
as well as the mature fruit, all show a strong tendency to become
elongated. Several varieties of the melon are interesting from as-
suming the characteristic features of distinct species and even of
distinct though allied genera: thus the serpent-melon has some re-
140 Naudin, in 'Annal. des Sci. Nat.,'
4th ser. Bot. tom. xi. 1859, p. 28.
141 Mémoire sur les Cucurbitacées,
1826, pp. 6, 24.
142 Flore des Serres,' Oct. 1861, quot-
ed in Gardener's Chronicle,' 1861, p.
1135. I have also consulted and taken
some facts from M. Naudin's Memoir on
Cucumis in Annal. des Sc. Nat.,' 4th
series, Bot. tom. xi. 1859, p. 5.
143 See also Sageret's "Mémoire,'
p. 7.
19

434
CHAP. X.
TREES.
semblance to the fruit of Trichosanthes anguina; we have seen that
other varieties closely resemble cucumbers; some Egyptian varie-
ties have their seeds attached to a portion of the pulp, and this is
characteristic of certain wild forms. Lastly, a variety of melon from
Algiers is remarkable from announcing its maturity by “a sponta-
neous and almost sudden dislocation," when deep cracks suddenly
appear, and the fruit falls to pieces; and this occurs with the wild
C. momordica. Finally, M. Naudin well remarks that this “extra-
ordinary production of races and varieties by a single species, and
their permanence when not interfered with by crossing, are pheno-
mena well calculated to cause reflection."
USEFUL AND ORNAMENTAL TREES.
TREES deserve a passing notice on account of the numerous varie-
ties which they present, differing in their precocity, in their manner
of growth, foliage, and bark. Thus of the common ash (Fraxinus
excelsior) the catalogue of Messrs. Lawson of Edinburgh includes
twenty-one varieties, some of which differ much in their bark; there
is a yellow, a streaked reddish-white, a purple, a wart-barked and a
fungous-barked variety. 144 Of hollies no less than eighty-four va-
rieties are grown alongside each other in Mr. Paul's nursery,
In
the case of trees, all the recorded varieties, as far as I can find out,
have been suddenly produced by one single act of variation. The
length of time required to raise many generations, and the little
value set on the fanciful varieties, explains how it is that successive
modifications have not been accumulated by selection ; hence, also
it follows that we do not here meet with sub-varieties subordinate
to varieties, and these again subordinate to higher groups. On the
Continent, however, where the forests are more carefully attended
to than in England, Alph. De Candolle 146 says that there is not a
forester who does not search for seeds from that variety which he
esteems the most valuable.
Our useful trees have seldom been exposed to any great change
of conditions; they have not been richly manured, and the English
kinds grow under their proper climate. Yet in examining extensive
beds of seedlings in nursery-gardens considerable differences may
be generally observed in them; and whilst touring in England I
have been surprised at the amount of difference in the appearance
of the same species in our hedgerows and woods. But as plants
(144 Loudon's Arboretum et Frutice-
tum,' vol. ii. p. 1217.
145 Gardener's Chron., ' 1866, p. 1096.
148 "Géograph. Bot.,' p. 1096.

CHAP. X.
435
TREES.
vary so much in a truly wild state, it would be difficult for even a
skilful botanist to pronounce whether, as I believe to be the case,
hedgerow trees vary more than those growing in a primeval forest.
Trees when planted by man in woods or hedges do not grow where
they would naturally be able to hold their place against a host of
competitors, and are therefore exposed to conditions not strictly
natural: even this slight change would probably suffice to cause
seedlings raised from such trees to be variable. Whether or not our
half-wild English trees, as a general rule, are more variable than
trees growing in their native forests, there can hardly be a doubt
that they have yielded a greater number of strongly marked and
singular variations of structure.
In manner of growth, we have weeping or pendulous varieties of
the willow, ash, elm, oak, and yew, and other trees; and this weep-
ing habit is sometimes inherited, though in a singularly capricious
manner. In the Lombardy poplar, and in certain fastigate or pyra-
midal varieties of thorns, junipers, oaks, &c., we have an opposite
kind of growth. The Hessian oak,147 which is famous from its fasti-
gate habit and size, bears hardly any resemblance in general appear-
ance to a common oak; "its acorns are not sure to produce plants of
the same habit; some, however, turn out the same as the parent-tree.”
Another fastigate oak is said to have been found wild in the Pyrenees,
and this is a surprising circumstance; it generally comes so true by
seed, that De Candolle considered it as specifically distinct.148 The
fastigate Juniper (J. suecica) likewise transmits its character by
seed.149 Dr. Falconer informs me that in the Botanic Gardens at
Calcutta the great heat causes apple-trees to become fastigate; and
we thus see the same result following from the effects of climate and
from an innate spontaneous tendency.150
In foliage we have variegated leaves which are often inherited ;
dark purple or red leaves, as in the hazel, barberry, and beech, the
colour in these two latter trees being sometimes strongly and some-
times weakly inherited ; 151 deeply-cut leaves; and leaves covered
with prickles, as in the variety of the holly well called ferox, which
is said to reproduce itself by seed.152 In fact, nearly all the peculiar
varieties evince a tendency, more or less strongly marked, to repro-
147 Gardener's Chron.,' 1842, p. 36.
148 Loudon's Arboretum et Fruti-
cetum,' vol. iii. p. 1731.
149 Ibid., vol. iv. p. 2489.
150 Godron ('De l'Espèce,' tom. ii. p.
91) describes four varieties of Robinia
remarkable from their manner of growth.
151 Journal of a Horticultural Tour,
by Caledonian Hort. Soc.,' 1823, p. 107
Alph. De Candolle, 'Géograph. Bot.,' p.
1083. Verlot, Sur la Production des
Variétés,' 1865, p. 55, for the Barberrg.
152 Loudon's Arboretum et Frutice-
tum,' vol. ii. p. 508.

436
CHAP. X.
TREES.
" 155
duce themselves by seed.158 This is to a certain extent the case, ac-
cording to Bosc, 154 with three varieties of the elm, namely, the broad-
leafed, lime-leafed, and twisted elm, in which latter the fibres of the
wood are twisted. Even with the heterophyllous hornbeam (Car-
pinus betulus), which bears on each twig leaves of two shapes,
"several plants raised from seed all retained the same peculiarity.”
I will add only one other remarkable case of variation in foliage,
namely, the occurrence of two sub-varieties of the ash with simple
instead of pinnated leaves, and which generally transmit their cha-
racter by seed. 158 The occurrence, in trees belonging to widely dif-
ferent orders, of weeping and fastigate varieties, and of trees bearing
deeply cut, variegated, and purple leaves, shows that these deviations
of structure must result from some very general physiological laws.
Differences in general appearance and foliage, not more strongly
marked than those above indicated, have led good observers to rank
as distinct species certain forms which are now known to be mere
varieties. Thus a plane-tree long cultivated in England was con-
sidered by almost every one as a North American species; but is
now ascertained by old records, as I am informed by Dr. Hooker, to
be a variety. So again the Thuja pendula or filiformis was ranked
by such good observers as Lambert, Wallich, and others as a true
species; but it is now known that the original plants, five in number,
suddenly appeared in a bed of seedlings, raised at Mr. Loddige's
nursery, from T. orientalis; and Dr. Hooker has adduced excellent
evidence that at Turin seeds of T. pendula have reproduced the
parent-form, T. orientalis. 157
Every one must have noticed how certain individual trees regular-
ly put forth and shed their leaves earlier or later than others of the
same species. There is a famous horse-chesnut in the Tuileries
which is named from leafing so much earlier than the others. There
is also an oak near Edinburgh, which retains its leaves to a very late
period. These differences have been attributed by some authors to
the nature of the soil in which the trees grow; but Archbishop
Whately grafted an early thorn on a late one, and vice versa, and
both grafts kept to their proper periods, which differed by about a
fortnight, as if they still grew on their own stocks.168 There is a
Cornish variety of the elm which is almost an evergreen, and is so
153 Verlot, Des Variétés,' 1865, p. 92.
154 Loudon's Arboretum et Frutice-
tum,' vol. iii. p. 1376.
155 Gardener's Chronicle;' 1841, p.
687.
156 Godron, 'De l'Espèce,' tom. ii. p.
69. In Loudon's Gardener's Mag., vol.
xii. 1836, p. 371, a variegated bushy ash
is described and figured, as having sim-
ple leaves; it originated in Ireland.
157 Gardener's Chron.,' 1861, p. 575.
158 Quoted from Royal Irish Academy
in Gardener's Chron.,' 1841, p. 767.

CHAP. X.
437
TREES.
159
tender that the shoots are often killed by the frost ; and the varie-
ties of the Turkish oak (Q. cerris) may be arranged as deciduous,
sub-evergreen, and evergreen.
Scotch Fir (Pinus sylvestris).-I ailude to this tree as it bears on
the question of the greater variability of our hedgerow trees com-
pared with those under strictly natural conditions. A well-informed
writer 150 states that the Scotch fir presents few varieties in its na-
tive Scotch forests; but that it “varies much in figure and foliage,
“and in the size, shape, and colour of its cones, when several gen-
"erations have been produced away from its native locality." There
is little doubt that the highland and lowland varieties differ in the
value of their timber, and that they can be propagated truly by
seed; thus justifying Loudon's remark, that " a variety is often of
as much importance as a species, and sometimes far more so." 161 I
may mention one rather important point in which this tree occa-
sionally varies; in the classification of the Coniferæ, sections are
founded on whether two, three, or five leaves are included in the
same sheath; the Scotch fir has properly only two leaves thus en-
closed, but specimens have been observed with groups of three
leaves in a sheath. 162 Besides these differences in the semi-culti-
vated Scotch fir, there are in several parts of Europe natural or geo-
graphical races, which have been ranked by some authors as dis-
tinct species. 103 Loudon 164 considers P. pumilio, with its several
sub-varieties, as Mughus, nana, &c., which differ much when planted
in different soils and only come “tolerably true from seed," as alpine
varieties of the Scotch fir; if this were proved to be the case, it
would be an interesting fact as showing that dwarfing from long
exposure to a severe climate is to a certain extent inherited.
The Hawthorn (Crataegus oxycantha) has varied much. Besides
endless slighter variations in the form of the leaves, and in the size,
hardness, fleshiness, and shape of the berries, Loudon 165 enumerates
twenty-nine well-marked varieties. Besides those cultivated for
their pretty flowers, there are others with golden-yellow, black, and
whitish berries; others with woolly berries, and others with re-
curved thorns. Loudon truly remarks that the chief reason why
159 Loudon's Arboretum et Frutice- päischer Pinus-arten von Dr. Christ :
tum:' for Elm, see vol. iii. p. 1376; for Flora, 1864.' He shows that in the Ober-
Oak, p. 1846.
Engadin P. sylvestris and montana are
160 Gardener's Chronicle,' 1849, p. connected by intermediate links.
822.
164 · Arboretum et Fruticetum,' vol.
161 Arboretum et Fruticetum,' vol. iv. iv. pp. 1159 and 2189.
165 Ibid., vol. ii. p. 830; Loudon's
162 Gardener's Chron.,' 1852, p. 693. Gardener's Magazine,' vol. vi, 1830, p.
163 See Beiträge zur Kentniss Euro- 714.
p. 2150.

438
CHAP. X.
FLOWERS.
the hawthorn has yielded more varieties than most other trees, is
that curious nurserymen select any remarkable variety out of the
immense beds of seedlings which are annually raised for making
hedges. The flowers of the Hawthorn usually include from one to
three pistils; but in two varieties, named Monogyna and Sibirica,
there is only a single pistil; and d’Asso states that the common
thorn in Spain is constantly in this state.166 There is also a variety
which is apetalous, or has its petals reduced to mere rudiments.
The famous Glastonbury thorn flowers and leafs towards the end of
December, at which time it bears berries produced from an earlier
crop of flowers. 107 It is worth notice that several varieties of the
hawthorn, as well as of the lime and juniper, are very distinct in
their foliage and habit whilst young, but in the course of thirty or
forty years become extremely like each other ; 168 thus reminding us
of the well-known fact that the deodar, the cedar of Lebanon, and
that of the Atlas, are distinguished with the greatest ease whilst
young, but with difficulty when old.
FLOWERS
I SHALL not for several reasons treat the variability of plants which
are cultivated for their flowers alone at any great length. Many of
our favourite kinds in their present state are the descendants of two
or more species crossed and commingled together, and this circum-
stance alone would render it difficult to detect the differences due to
variation. For instance, our Roses, Petunias, Calceolarias, Fuchsias,
Verbenas, Gladioli, Pelargoniums, &c., certainly have had a multiple
origin. A botanist well acquainted with the parent-forms would
probably detect some curious structural differences in their crossed
and cultivated descendant; and he would certainly observe many
new and remarkable constitutional peculiarities. I will give a few
instances, all relating to the Pelargonium, and taken chiefly from
Mr. Beck, 169 a famous cultivator of this plant: some varieties re.
quire more water than others; some are "very impatient of the
knife if too greedily used in making cuttings;" some, when potted,
scarcely "show a root at the outside of the ball of the earth;" one
variety requires a certain amount of confinement in the pot to make
it throw up a flower-stem ; some varieties bloom well at the com-
mencement of the season, others at the close; one variety is
known,170 which will stand "even pine-apple top and bottom heat,
166 Loudon's Arboretum et Frutice-
tum,' vol. ii. p. 834.
187 Loudon's Gardener's Mag.,' vol.
ix. 1833, p. 123.
168 Ibid., vol. xi. 1835, p. 503.
169 Gardener's Chron.,'1845, p. 623.
170 D. Beaton, in 'Cottage Gardener,'
1860, p. 377. See also Mr. Beck, on the

CHAP. X.
439
FLOWERS.
without looking any more drawn than if it had stood in a common
greenhouse; and Blanche Fleur seems as if made on purpose for
growing in winter, like many bulbs, and to rest all summer."
These odd constitutional peculiarities would fit a plant when grow-
ing in a state of nature for widely different circumstances and
climates.
Flowers possess little interest under our present point of view,
because they have been almost exclusively attended to and selected
for their beautiful colours, size, perfect outline, and manner of
growth. In these particulars hardly one long-cultivated flower can
be named which has not varied greatly. What does a florist care
for the shape and structure of the organs of fructification, unless,
indeed, they add to the beauty of the flower? When this is the
case, flowers become modified in important points; stamens and
pistils may be converted into petals, and additional petals may be
developed, as in all double flowers. The process of gradual selection
by which fiowers have been rendered more and more double, each
step in the process of conversion being inherited, has been recorded
in several instances. In the so-called double flowers of the Compositæ,
the corollas of the central florets are greatly modified, and the modifi.
cations are likewise inherited. In the columbine (Aquilegia vulgaris)
some of the stamens are converted into petals having the shape of
nectaries, one neatly fitting into the other; but in one variety they
are converted into simple petals.171 In the hose and hose primulæ,
the calyx becomes brightly coloured and enlarged so as to resemble
a corolla ; and Mr. W. Wooler informs me that this peculiarity is
transmitted; for he crossed a common polyanthus with one having
a coloured calyx, 172 and some of the seedlings inherited the coloured
calyx during at least six generations. In the "hen-and-chicken”
daisy the main flower is surrounded by a brood of small flowers de-
veloped from buds in the axils of the scales of the involucre. A
wonderful poppy has been described, in which the stamens are con-
verted into pistils; and so strictly was this peculiarity inherited that,
out of 154 seedlings, one alone reverted to the ordinary and common
type.173 of the cock’s-comb (Celosia cristata), which is an annual,
there are several races in which the flower-stem is wonderfully
“fasciated” or compressed ; and one has been exhibited 174 actually
eighteen inches in breadth. Peloric races of Gloxinia speciosa and
p. 133.
habits of Queen Mab, in Gardener's
Chronicle,' 1845, p. 226.
173 Quoted by Alph. de Candolle,
171 Moquin-Tandon, 'Eléments de "Bibl. Univ.,' November, 1862, p. 58.
Tératologie,' 1841, p. 213.
174 Knight, Transact. Hort. Soc.,' vol.
172 See also Cottage Gardener, '1860, iv. p. 322.

440
CHAP. X.
FLOWERS.
Antirrhinum majus can be propagated by seed, and they differ in a
wonderful manner from the typical form both in structure and ap-
pearance.
A much more remarkable modification has been recorded by Sir
William and Dr. Hooker 175 in Begonia frigida. This plant properly
produces male and female flowers on the same fascicles; and in the
female flowers the perianth is superior ; but a plant at Kew produced,
besides the ordinary flowers, others which graduated towards a per-
fect hermaphrodite structure, and in these flowers the perianth was
inferior. To show the importance of this modification under a classi-
ficatory point of view, I may quote what Prof. Harvey says, namely,
that had it " occurred in a state of nature, and had a botanist col-
lected a plant with such flowers, he would not only have placed it
in a distinct genus from Begonia, but would probably have con-
sidered it as the type of a new natural order.” This modification
cannot in one sense be considered as a monstrosity, for analogous
structures naturally occur in other orders, as with Saxifragas and
Aristolochiaceæ. The interest of the case is largely added to by Mr.
C. W. Crocker's observation that seedlings from the normal flowers
produced plants which bore, in about the same proportion as the
parent-plant, hermaphrodite flowers having inferior perianths. The
hermaphrodite flowers fertilised with their own pollen were sterile.
If florists had attended to, selected, and propagated by seed other
modifications of structure besides those which are beautiful, a host
of curious varieties would certainly have been raised; and they
would probably have transmitted their characters so truly that the
cultivator would have felt aggrieved, as in the case of culinary
vegetables, if his whole bed had not presented a uniform appearance.
Florists have attended in some instances to the leaves of their plant,
and have thus produced the most elegant and symmetrical patterns
of white, red, and green, which, as in the case of the pelargonium, are
sometimes strictly inherited.176 Any one who will habitually examine
highly-cultivated flowers in gardens and greenhouses will observe
numerous deviations in structure; but most of these must be ranked
as mere monstrosities, and are only so far interesting as showing
how plastic the organisation becomes under high cultivation. From
175 Botanical Magazine,' tab. 5160,
fig. 4; Dr. Hooker, in *Gardener's
Chron.,' 1860, p. 190; Prof. Harvey, in
Gardener's Chron.,' 1860, p. 145; Mr.
Crocker, in 'Gardener's Chron.,' 1861,
Bot.,' p. 1083; 'Gard. Chronicle,' 1861,
p. 433. The inheritance of the white
and golden zones in Pelargonium largely
depends on the nature of the soil. See
D. Beaton, in Journal of Horticulture,
1861, p. 64.
p. 1092.
176 Alph. de Candolle, 'Géograph.

CHAP. X.
*441
FLOWERS.
this point of view such works as Professor Moquin-Tandon's Téra-
tologie' are highly instructive.
Roses. These flowers offer an instance of a number of forms
generally ranked as species, namely, R. centifolia, gallica, alba, da-
mascena, spinosissima, bracteata, Indica, semperflorens, moschata, &c.,
which have largely varied and been intercrossed. The genus Rosa
is a notoriously difficult one, and though some of the above forms
are admitted by all botanists to be distinct species, others are doubt-
ful; thus, with respect to the British forms, Babington makes seven-
teen, and Bentham only five species. The hybrids from some of the
most distinct forms—for instance, from R. Indica, fertilised by the
pollen of R. centifolia-produce an abundance of seed; I state this
on the authority of Mr. Rivers, 177 from whose work I have drawn
most of the following statements. As almost all the aboriginal
forms brought from different countries have been crossed and re-
crossed, it is no wonder that Targioni-Tozzetti, in speaking of the
common roses of the Italian gardens, remarks that "the native
country and precise form of the wild type of most of them are in-
volved in much uncertainty.” 178 Nevertheless Mr. Rivers in refer-
ring to R. Indica (p. 68) says that the descendants of each group
may generally be recognised by a close observer. The same author
often speaks of roses as having been a little hybridised ; but it is
evident that in very many cases the differences due to variation and
to hybridisation can now only be conjecturally distinguished.
The species have varied both by seed and by buds; such modified
buds being often called by gardeners sports. In the following chap-
ter I shall fully discuss this latter subject, and shall show that bud-
variations can be propagated not only by grafting and budding, but
often even by seed. Whenever a new rose appears with any pecu-
liar character, however produced, if it yields seed, Mr. Rivers (p. 4)
fully expects it to become the parent-type of a new family. The
tendency to vary is so strong in some kinds, as in the Village Maid
(Rivers, p. 16), that when grown in different soils it varies so much
in colour that it has been thought to form several distinct kinds.
Altogether the number of kinds is very great: thus M. Desportes,
in his Catalogue for 1829, enumerates 2562 as cultivated in France;
but no doubt a large proportion of these are merely nominal.
It would be useless to specify the many points of difference be-
tween the various kinds, but some constitutional peculiarities may
be mentioned. Several French roses (Rivers, p. 12) will not succeed
178 Journal Hort. Soc.,' vol. ix, 1855,
177 Rose Amateur's Guide,' T. Rivers,
1887, p. 21.
p. 182.
19*

442
CHAP. X.
FLOWERS.
in England; and an excellent horticulturist 179 remarks, that "Even
in the same garden you will find that a rose that will do nothing
under a south wall will do well under a north one. That is the case
with Paul Joseph here. It grows strongly and blooms beautifully
close to a north wall. For three years seven plants have done noth-
ing under a south wall.” Many roses can be forced,“ many are to-
tally unfit for forcing, among which is General Jacqueminot.” 180
From the effects of crossing and variation Mr. Rivers enthusiastically
anticipates (p. 87) that the day will come when all our roses, even
moss-roses, will have evergreen foliage, brilliant and fragrant flowers,
and the habit of blooming from June till November. "A distant
view this seems, but perseverance in gardening will yet achieve
wonders," as assuredly it has already achieved wonders.
It may be worth while briefly to give the well-known history of
one class of roses. In 1793 some wild Scotch roses (R. spinosissima)
were transplanted into a garden ; 181 and one of these bore flowers
slightly tinged with red, from which a plant was raised with semi-
monstrous flowers, also tinged with red; seedlings from this flower
were semi-double, and by continued selection, in about nine or ten
years, eight sub-varieties were raised. In the course of less than
twenty years these double Scotch roses had so much increased in
number and kind, that twenty-six well-marked varieties, classed in
eight sections, were described by Mr. Sabine. In 1841 182 it is said
that three hundred varieties could be procured in the nursery-gar-
dens near Glasgow; and these are described as blush, crimson, pur-
ple, red, marbled, two-coloured, white, and yellow, and as differing
much in the size and shape of the flower.
Pansy or Heartsease (Viola tricolor, &c.).—The history of this
flower seems to be pretty well known; it was grown in Evelyn's
garden in 1687; but the varieties were not attended to till 1810–
1812, when Lady Monke, together with Mr. Lee the well-known
nurseryman, energetically commenced their culture, and in the
course of a few years twenty varieties could be purchased.183 At
about the same period, namely in 1813 or 1814, Lord Gambier col-
lected some wild plants, and his gardener, Mr. Thomson, cultivated
them together with some common garden varieties, and soon effect-
ed a great improvement. The first great change was the conversion
179 The Rev. W.F. Radclyffe, in 'Jour-
nal of Horticulture,' March 14, 1865, p.
207.
180 Gardener's Chronicle,' 1861, p. 46.
181 Mr. Sabine, in Transact. Hort.
Soc.,' vol. iv. p. 285.
182 An Encyclop. of Plants,' by J. O.
Loudon, 1841, p. 443.
183 Loudon's Gardener's Magazine,
vol. xi. 1835, p. 427; also Journal of
Horticulture,' April 14, 1863, p. 275.

CHAP. X.
443
FLOWERS.
of the dark lines in the centre of the flower into a dark eye or centre,
which at that period had never been seen, but is now considered
one of the chief requisites of a first-rate flower. In 1835 a book
entirely devoted to this flower was published, and four hundred
named varieties were on sale. From these circumstances this plant
seemed to me worth studying, more especially from the great con-
trast between the small, dull, elongated, irregular flowers of the
wild pansy, and the beautiful, flat, symmetrical, circular, velvet-like
flowers, more than two inches in diameter, magnificently and va-
riously coloured, which are exhibited at our shows. But when I
came to inquire more closely, I found that, though the varieties were
so-modern, yet that much confusion and doubt prevailed about their
parentage. Florists believed that the varieties 184 are descended
from several wild stocks, namely, V. tricolor, lutea, grandiflora,
amana, and Altaica, more or less intercrossed. And when I looked to
botanical works to ascertain whether these forms ought to be rank-
ed as species, I found equal doubt and confusion. Viola Altaica
seems to be a distinct form, but what part it has played in the origin
of our varieties I know not; it is said to have been crossed with
V. lutea. Viola amana is now looked by all botanists as a
natural variety of V. grandiflora ; and this and V. sudetica have
been proved to be identical with V. lutea. The latter and V. tricolor
(including its admitted variety V. arvensis) are ranked as distinct
species by Babington ; and likewise by M. Gay,186 who has paid par-
ticular attention to the genus; but the specific distinction between
V. lutea and tricolor'is chiefly grounded on the one being strictly
and the other not strictly perennial, as well as on some other slight
and unimportant differences in the form of the stem and stipules.
Bentham unites these two forms; and a high authority on such mat-
ters, Mr. H. C. Watson, 187 says 'that, “while V. tricolor passes into
V. arvensis on the one side, it approximates so much towards V. lu-
tea and V. Curtisii on the other side, that a distinction becomes
scarcely more easy between them."
Hence, after having carefully compared numerous varieties, I
gave up the attempt as too difficult for any one except a professed
botanist. Most of the varieties present such inconstant characters,
that when grown in poor soil, or when flowering out of their
185
184 Loudon's Gardener's Magazine,
vol. viii. p. 575 ; vol. ix. p. 689.
185 Sir J. E. Smith, English Flora,'
vol. i. p. 306. H. C. Watson, Cybele
Britannica,' vol. i. 1847, p. 181.
186 Quoted from Annales des Sci-
ences,' in the Companion to the 'Bot.
Mag.,' vol. i. 1835, p. 159.
187 Cybele Britannica,' vol. i. p. 178.
See also Dr. Herbert on the changes of
colour in transplanted specimens, and
on the natural variations of V. grandi-
flora, in Transact. Hort. Soc.,' vol. iv.
p. 19.

444
CHAP. X.
FLOWERS.
proper season, they produce differently coloured and much smaller
flowers. Cultivators speak of this or that kind as being remarkably
constant or true; but by this they do not mean, as in other cases,
that the kind transmits its character by seed, but that the individual
plant does not change much under culture. The principle of in-
heritance, however, does hold good to a certain extent even with the
fleeting varieties of the Heartease, for to gain good sorts it is indis-
pensable to sow the seed of good sorts. Nevertheless in every large
seed-bed a few almost wild seedlings often reappear through rever
sion. On comparing the choicest varieties with the nearest allied
wild forms, besides the difference in the size, outline, and colour of
the flowers, the leaves are seen sometimes to differ in shape, as does
the calyx occasionally in the length and breadth of the sepals. The
differences in the form of the nectary more especially deserve notice;
because characters derived from this organ have been much used in
the discrimination of most of the species of Viola. In a large num-
ber of flowers compared in 1842 I found that in the greater number
the nectary was straight; in others the extremity was a little turned
upwards, or downwards, or inwards, so as to be completely hooked ;
in others, instead of being hooked, it was first turned rectangularly
downwards, and then backwards and upwards; in others the ex-
tremity was considerably enlarged ; and lastly, in some the basal
part was depressed, becoming, as usual, laterally compressed to-
wards the extremity. In a large number of flowers, on the other
hand, examined by me in 1856 from a nursery-garden in a different
part of England, the nectary hardly varied at all. Now M. Gay
says that in certain districts, especially in Auvergne, the nectary of
wild V.grandiflora varies in the manner just described. Must we
conclude from this that the cultivated varieties first mentioned were
all descended from V. grandiflora, and that the second lot, though
having the same general appearance, were descended from V. tri-
color, of which the nectary, according to M. Gay, is subject to little
variation? Or is it not more probable that both these wild forms
would be found under other conditions to vary in the same manner
and degree, thus showing that they ought not to be ranked as
specifically distinct ?
The Dahlia has been referred to by almost every author who has
written on the variation of plants, because it is believed that all the
varieties are descended from a single species, and because all have
arisen since 1802 in France, and since 1804 in England.188 Mr.
Sabine remarks that "it seems as if some period of cultivation had
188 Salisbury, in Transact. Hort. Soc.,' vol. i. 1912, pp. 84, 92. A semi-double
variety was produced in Madrid in 1790.

CHAP. X.
445
FLOWERS.
been required before the fixed qualities of the native plant gave way
and began to sport into those changes which now so delight us.” 189
The flowers have been greatly modified in shape from a flat to a
globular form. Anemone and ranunculus-like races, 190 which differ
in the form and arrangement of the florets, have arisen ; also
dwarfed races, one of which is only eighteen inches in height. The
seeds vary much in size. The petals are uniformly coloured or
tipped or striped, and present an almost infinite diversity of tints.
Seedlings of fourteen different colours 191 have been raised from the
same plant; yet, as Mr. Sabine has remarked, "many of the seed-
lings follow their parents in colour.” The period of flowering has
been considerably hastened, and this has probably been effected by
continued selection. Salisbury, writing 1808, says that they then
flowered from September to November; in 1828 some new dwarf
varieties began flowering in June ; 192 and Mr. Grieve informs me
that the dwarf purple Zelinda in his garden is in full bloom by the
middle of June and sometimes even earlier. Slight constitutional
differences have been observed between certain varieties: thus, some
kinds succeed much better in one part of England than in another,193
and it has been noticed that some varieties require much more mois-
ture than others.
Such flowers as the carnation, common tulip, and hyacinth, which
are believed to be descended, each from a single wild form, present
innumerable varieties, differing almost exclusively in the size, form,
and colour of the flowers. These and some other anciently cultivated
plants which have been long propagated by offsets, pipings, bulbs,
&c., become so excessively variable, that almost each new plant
raised from seed forms a new variety, "all of which to describe
particularly," as old Gerarde wrote in 1597,"were to roll Sisyphus's
stone, or to number the sands."
Hyacinth (Hyacinthus orientalis).-It may, however, be worth
while to give a short account of this plant, which was introduced
into England in 1596 from the Levant.195 The petals of the original
flower, says Mr. Paul, were narrow, wrinkled, pointed, and of a
flimsy texture; now they are broad, smooth, solid, and rounded.
194
193 Mr. Wildman, in Gardener's
Chron.,' 1843, p. 87.
194 Cottage Gardener,' April 8, 1856,
p. 33.
189 Transact. Hort. Soc.,' vol. iii.
1920, p. 225.
190 Loudon's Gardener's Mag.,' vol.
vi. 1830, p. 77.
191 Loudon's 'Encyclop. of Garden-
ing,' p. 1035.
192 Transact. Hort. Soc.,' vol. i. p. 91 ;
and Loudon's Gardener's Mag.,' vol.
ii. 1828, p. 179.
195 The best and fullest account of
this plant which I have met with is by
a famous horticulturist, Mr. Paul of
Waltham, in the Gardener's Chronicle,'
1864, p. 342.

446
CHAP. X.
FLOWERS.
66
The erectness, breadth, and length of the whole spike, and the size
of the flowers, have all increased. The colours have been intensified
and diversified. Gerarde, in 1597, enumerates four, and Parkinson,
in 1629, eight varieties. Now the varieties are very numerous, and
they were still more numerous a century ago. Mr. Paul remarks
that "it is interesting to compare the Hyacinths of 1629 with those
“of 1864, and to mark the improvement. Two hundred and thirty-
“ five years have elapsed since then, and this simple flower serves
“well to illustrate the great fact that the original forms of nature
“ do not remain fixed and stationary, at least when brought under
"cultivation. While looking at the extremes, we must not how-
ever forget that there are intermediate stages which are for the
“most part lost to us. Nature will sometimes indulge herself with
"a leap, but as a rule her march is slow and gradual.” He adds
that the cultivator should have "in his mind an ideal of beauty, for
“the realisation of which he works with head and hand.” We thus
see how clearly Mr. Paul, an eminently successful cultivator of this
flower, appreciates the action of methodical selection.
In a curious and apparently trustworthy treatise, published at
Amsterdam 196 in 1768, it is stated that nearly 2000 sorts were then
known; but in 1864 Mr. Paul found only 700 in the largest garden
at Haarlem. In this treatise it is said that not an instance is known
of any one variety reproducing itself truly by seed: the white kinds,
however, now 197 almost always 'yield white hyacinths, and the yel-
low kinds come nearly true. The hyacinth is remarkable from hav-
ing given rise to varieties with bright blue, pink, and distinctly
yellow flowers. These three primary colours do not occur in the
varieties of any other species; nor do they often all occur even in
the distinct species of the same genus. Although the several kinds
of hyacinths differ but slightly from each other except in colour, yet
each kind has its own individual character, which can be recognised
by a highly educated eye; thus the writer of the Amsterdam trea-
tise asserts (p. 43) that some experienced florists, such as the famous
G. Voorholm, seldom failed in a collection of above twelve hundred
sorts to recognise each variety by the bulb alone! This same writer
mentions some few singular variations: for instance, the hyacinth
commonly produces six leaves, but there is one kind (p. 35) which
scarcely ever has more than three leaves; another never more than
five; whilst others regularly produce either seven or eight leaves.
A variety, called la Coriphée, invariably produces (p. 116) two flower-
198 Des Jacinthes, de leur Anatomie,
Reproduction, et Culture,' Amsterdam,
1768
197 Alph. de Candolle, 'Géograph.
Bot.,' p. 1082.

CHAP. X.
447
FLOWERS.
stems, united together and covered by one skin. The flower-stem
in another kind (p. 128) comes out of the ground in a coloured
sheath, before the appearance of the leaves, and is consequently lia-
ble to suffer from frost. Another variety always pushes a second
flower-stem after the first has begun to develop itself. Lastly, white
hyacinths with red, purple, or violet centres (p. 129) are the most
liable to rot. Thus, the hyacinth, like so many previous plants,
when long cultivated and closely watched, is found to offer many
singular variations.
In the two last chapters I have given in some detail
the range of variation, and the history, as far as known,
of a considerable number of plants, which have been cul-
tivated for various purposes. But some of the most va-
riable plants, such as Kidney-beans, Capsicum, Millets,
Sorghum, &c., have been passed over; for botanists are
not agreed which kinds ought to rank as species and
which as varieties; and the wild parent-species are un-
known.198 Many plants long cultivated in tropical coun-
tries, such as the Banana, have produced numerous varie-
ties; but as these have never been described with even
moderate care, they also are here passed over. Never-
theless a sufficient, and perhaps more than sufficient, num-
ber of cases have been given, so that the reader may
be
enabled to judge for himself on the nature and extent of
the variation which cultivated plants have undergone.
198 Alph, de Candolle, Géograph. Bot.,' p. 983.

448
CHAP, XI.
BUD-VARIATION
CHAPTER XI.
ON BUD-VARIATION, AND ON CERTAIN ANOMALOUS
MODES OF REPRODUCTION AND VARIATION.
BUD-VARIATIONS IN THE PEACH, PLUM, CHIERRY, VINE, GOOSE-
BERRY, 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, TU-
LERS, AND BULBS — ON THE BREAKING OF TULIPS — BUD-VARIA-
TIONS GRADUATE INTO CHANGES CONSEQUENT ON CHANGED
CONDITIONS OF LIFE - CYTISUS ADAMI, ITS ORIGIN AND TRANS-
FORMATION - ON THE UNION OF TWO DIFFERENT EMBRYOS IN
ONE SEED --THE TRIFACIAL ORANGE-ON REVERSION BY BUDS
IN HYBRIDS AND MONGRELS - ON THE PRODUCTION OF MODIFIED
BUDS BY THE GRAFTING OF ONE VARIETY OR SPECIES ON
ANOTHER — ON THE DIRECT OR IMMEDIATE ACTION OF FOREIGN
POLLEN ON THE MOTHER-PLANT -- ON THE EFFECTS IN FEMALE
ANIMALS OF A FIRST IMPREGNATION ON THE SUBSEQUENT OFF-
SPRING - CONCLUSION AND SUMMARY.
Tuis chapter will be chiefly devoted to a subject in many
respects important, namely, bud-variation. By this term
I include all those sudden changes in structure or ap-
pearance which occasionally occur in full-grown plants
in their flower-buds or leaf-buds. Gardeners call such
changes “Sports;" but this, as previously remarked, is
an ill-defined expression, as it has often been applied to
strongly marked variations in seedling plants. The dif-
ference between seminal and bud reproduction is not
so great as it at first appears; for each bud is in one
sense a new and distinct individual; but such indi-
viduals are produced through the formation of various

CHAP. XI.
449
FRUIT.
kinds of buds without the aid of any special apparatus,
whilst fertile seeds are produced by the concourse of the
two sexual elements. The modifications which arise
through bud-variation can generally be propagated to
any extent by grafting, budding, cuttings, bulbs, &c.,
and occasionally even by seed. Some few of our most
beautiful and useful productions have arisen by bud-
variation.
Bud-variations have as yet been observed only in the
vegetable kingdom; but it is probable that if compound
animals, such as corals, &c., had been subjected to a long
course of domestication, they would have varied by buds;
for they resemble plants in many respects. Thus any
new or peculiar character presented by a compound
animal is propagated by budding, as occurs with differ-
ently coloured Hydras, and as Mr. Gosse has shown to
be the case with a singular variety of a true coral. Va-
rieties of the Hydra have also been grafted on other
varieties, and have retained their character.
I will in the first place give all the cases of bud-varia-
tions which I have been able to collect, and afterwards
show their importance. These cases prove that those
* authors who, like Pallas, attribute all variability to the
crossing either of distinct races, or of individuals belong-
ing to the same race but somewhat different from each
other, are in error; as are those authors who attribute
all variability to the mere act of sexual union. Nor can
we account in all cases for the appearance through bud-
variation of new characters by the principle of reversion
to long-lost characters. He who wishes to judge how
far the conditions of life directly cause each particular
variation ought to reflect well on the cases immediately
to be given. I will commence with bud-variations, as
exhibited in the fruit, and then pass on to flowers, and
finally to leaves.
Peach (Amygdalus Persica).-In the last chapter I gave two

450
СНАР. XI,
BUD-VARIATION.
cases of a peach-almond and double-flowered almond which sudden-
ly produced fruit closely resembling true peaches. I have also
recorded many cases of peach-trees producing buds, which, when
developed into branches, have yielded nectarines. We have seen
that no less than six named and several unnamed varieties of the
peach have thus produced several varieties of nectarine. I have
shown that it is highly improbable that all these peach-trees, some
of which are old varieties, and have been propagated by the mil-
lion, are hybrids from the peach and nectarine, and that it is
opposed to all analogy to attribute the occasional production of
nectarines on peach-trees to the direct action of pollen from some
neighbouring nectarine-tree. Several of the cases are highly re-
markable, because, firstly, the fruit thus produced has sometimes
been in part a nectarine and in part a peach ; secondly, because
nectarines thus suddenly produced have reproduced themselves by
seed ; and thirdly, because nectarines are produced from peach-trees
from seed as well as from buds. The seed of the nectarine, on the
other hand, occasionally produces peaches; and we have seen in
one instance that a nectarine-tree yielded peaches by bud-variation.
As the peach is certainly the oldest or primary variety, the pro-
duction of peaches from nectarines, either by seeds or buds, may
perhaps be considered as a case of reversion. Certain trees have
also been described as indifferently bearing peaches or nectarines,
and this may be considered as bud-variation carried to an extreme
degree.
The grosse mignonne peach at Montreuil produced “from a sport-
ing branch” the grosse mignonne tardive, “a most excellent varie-
ty," which ripens its fruit a fortnight later than the parent tree,
and is equally good. This same peach has likewise produced by
bud-variation the early grosse mignonne. Hunt's large tawny nec-
tarine originated from Hunt's small tawny nectarine, but not
through seminal reproduction."2
Plums.-Mr. Knight states that a tree of the yellow magnum
bonum plum, forty years old, which had always borne ordinary
fruit, produced a branch which yielded red magnum bonums.
Mr. Rivers, of Sawbridgeworth, informs me (Jan. 1863) that a sin-
gle tree out of 400 or 500 trees of the Early Prolific plum, which
is a purple kind, descended from an old French variety bearing
purple fruit, produced when about ten years old bright yellow
plums; these differed in no respect except colour from those on
1 Gardener's Chron.,' 1854, p. 821.
2 'Lindley's Guide to Orchard,' as
quoted in Gard. Chronicle,' 1852. p.
821. For the Early mignonne peach,
see Gardener's Chron.,'1864, p. 1251.
3. Transact. Hort. Soc.,' vol. ii. p. 160.

CHAP. XI.
451
FRUIT.
the other trees, but were unlike any other known kind of yellow
plum.
Cherry (Prunus cerasus).—Mr. Knight has recorded (idem) the
case of a branch of a May-Duke cherry, which, though certainly
never grafted, always produced fruit, ripening later, and more ob-
long, than the fruit on the other branches, Another account has
been given of two May-Duke cherry-trees in Scotland, with branches
bearing oblong, and very fine fruit, which invariably ripened, as in
Knight's case, a fortnight later than the other cherries.
Grapes (vitis vinifera).—The black or purple Frontignan in one
case produced during two successive years (and no doubt perma-
nently) spurs which bore white Frontignan grapes. In another
case, on the same foot-stalk, the lower berries" were well-coloured
black Frontignans; those next the stalk were white, with the ex-
ception of one black and one streaked berry;" and altogether there
were fifteen black and twelve white berries on the same stalk. In
another kind of grape black and amber-coloured berries were pro-
duced in the same cluster. Count Odart describes a variety which
often bears on the same stalk small round and large oblong ber-
ries; though the shape of the berry is generally a fixed character.
Here is another striking case given on the excellent authority of
M. Carrière :8 "a black Hamburgh grape (Frankenthal) was cut
down, and produced three suckers; one of these was layered, and
after a time produced much smaller berries, which always ripened
at least a fortnight earlier than the others. Of the remaining two
suckers, one produced every year fine grapes, whilst the other,
although it set an abundance of fruit, matured only a few, and
these of inferior quality.
Gooseberry (Ribes grossularia).-A remarkable case has been de-
scribed by Dr. Lindley of a bush which bore at the same time no
less than four kinds of berries, namely, hairy and red,-smooth,
small and red-green,-and yellow tinged with buff; the two lat-
ter kinds had a different flavour from the red berries, and their
seeds were coloured red. Three twigs on this bush grew close
together; the first bore three yellow berries and one red; the
second twig bore four yellow and one red ; and the third four red
and one yellow. Mr. Laxton also informs me that he has seen a
4 See alsoGardener's Chron.,' 1863,
p. 27.
5 Gard. Chron.,' 1552, p. 821.
6 Gardener's Chron.,' 1852, p. 629;
1856, p. 643 ; 18€4, p. 986. Other cases
are given by Braun, 'Rejuvenescence,'
in 'Ray Soc. Bot. Mem.,' 1853, p. 314.
7 'Ampelographie,' &c., 1849, p. 71.
8 Gardener's Chronicle,'1866, p. 970.
9 Gardener's Chronicle, 1855, pp.
597, 612.

452
CHAP. XT
BUD-VARIATION.
Red Warrington gooseberry bearing both red and yellow fruit on
the same branch.
Currant (Ribes rubrum).- A bush purchased as the Champagne,
which is a variety that bears blush-coloured fruit intermediate be-
tween red and white, produced during fourteen years, on separate
branches and mingled on the same branch, berries of the red,
white, and champagne kinds.20 The suspicion naturally arises
that this variety may have originated from a cross between a
red and white variety, and that the above transformation may
be accounted for by reversion to both parent-forms; but from the
foregoing complex case of the gooseberry this view is doubtful.
In France, a branch of a red-currant bush, about ten years old,
produced near the summit five white berries, and lower down,
amongst the red berries, one berry half red and half white.
Alexander Braun 12 also has often seen branches bearing red ber-
ries on white currants.
Pear (Pyrus communis).—Dureau de la Malle states that the
flowers on some trees of an ancient variety, the doyenné galeux,
were destroyed by frost: other flowers appeared in July, which
produced six pears; these exactly resembled in their skin and
taste the fruit of a distinct variety, the gros doyenné blanc, but
in shape were like the bon-chrétien : it was not ascertained whe-
ther this new variety could be propagated by budding or grafting.
The same author grafted a bon-chrétien on a quince, and it pro-
duced, besides its proper fruit, an apparently new variety, of a
peculiar form, with thick and rough skin.13
Apple (Pyrus malus).-In Canada, a tree of the variety called
Pound Sweet, produced, 14 between two of its proper fruit, an ap-
ple which was well russetted, small in size, different in shape, and
with a short peduncle. As no russet apple grew anywhere near,
this case apparently cannot be accounted for by the direct action of
foreign pollen. I shall hereafter give cases of apple-trees which
regularly produce fruit of two kinds, or half-and-half fruit; these
trees are generally supposed, and probably with truth, to be of
crossed parentage, and that the fruit reverts to both parent-forms.
Banana (Musa sapientium).—Sir R. Schomburgk states that he
10. Gardener's Chron.,' 1842, p. 873;
1855, p. 646. In the Chronicle,' 1866,
p. 876, Mr. P. Mackenzie states that the
bush still continues to bear the three
kinds of fruit, although they have
not been every year alike."
11 Revue Horticole,' quoted in Gard:
Chronicle,' 1844, p. 87.
12 'Rejuvenescence in Nature, Bot.
Memoirs Ray Soc.,' 1853, p. 314.
13 Comptes Rendus,' tom. xli., 1855,
p. 804. The second case is given on the
authority of Gaudichaud, idem, tom.
xxxiv., 1852, p. 748.
14 This case is given in the Gard,
Chronicle,' 1867, p. 403.

CH2P. XI.
453
FLOWERS.
saw in St. Domingo a raceme on the Fig Banana which bore
towards the base 125 fruits of the proper kind; and these were
succeeded, as is usual, higher up the raceme, by barren flowers, and
these by 420 fruits, having a widely different appearance, and ripen-
ing earlier than the proper fruit. The abnormal fruit closely re-
sembled, except in being smaller, that of the Musa Chinensis or
Cavendishi, which has generally been ranked as a distinct species. 15
FLOWERS.- Many cases have been recorded of a whole plant, or
single branch, or bud, suddenly producing flowers different from
the proper type in colour, form, size, doubleness, or other character.
Half the flower, or a smaller segment, sometimes changes colour.
Camellia. -The myrtle-leaved species (C. myrtifolia), and two or
three varieties of the common species, have been known to produce
hexagonal and imperfectly quadrangular flowers; and the branches
producing such flowers have been propagated by grafting 16 The
Pompone variety often bears" four distinguishable kinds of flowers,
“the pure white and the red-eyed, which appear promiscuously; the
“brindled pink and the rose-coloured, which may be kept separate
“with tolerable certainty by grafting from the branches that bear
"them.” A branch, also, on an old tree of the rose-coloured variety
has been seen to "revert to the pure white colour, an occurrence less
“common than the departure from it.” 17
Cratægus oxycantha.— A dark pink hawthorn has been known to
throw out a single tuft of pure white blossoms; 18 and Mr. A. Clap-
ham, nurseryman, of Bradford, informs me that his father had a
deep crimson thorn grafted on a white thorn, which, during several
years, always bore, high above the graft, bunches of white, pink,
and deep crimson flowers.
Azalea Indica is well known often to produce by buds new varie
ties. I have myself seen several cases. A plant of Azalea Indica
variegata has been exhibited bearing a truss of flowers of A. Ind.
Gledstanesii“ as true as could possibly be produced, thus evidencing
the origin of that fine variety." On another plant of A. Ind. varic-
gata a perfect flower of A. Ind. lateritia was produced; so that both
Gledstanesii and lateritia no doubt originally appeared as sporting
branches of A. Ind. variegata.
Cistus tricuspis.-A seedling of this plant, when some years old,
19
15 Journal of Proc. Linn. Soc.,' vol.
ii. Botany, p. 131.
16 Gard. Chronicle,' 1847, p. 207.
17 Herbert, Amaryllidaceæ,' 1838, p.
869.
18 Gardener's Chronicle,' 1843, p.
891.
19 Exhibited at Hort. Soc., London.
Repoxt in Gardener's Chron.,' 1814, p.
337.

454
CHAP. XI,
BUD-VARIATION.
produced, at Saharunpore,20 some branches "which bore leaves and
flowers widely different from the normal form.” “The abnormal
leaf is much less divided, and not acuminated. The petals are con-
siderably larger, and quite entire. There is also in the fresh state a
conspicuous, large, oblong gland, full of a viscid secretion, on the
back of each of the calycine segments.”
Althæa rosea.-A double yellow Hollyock suddenly turned one
year into a pure white single kind; subsequently a branch bearing
the original double yellow flowers reappeared in the midst of the
branches of the single white kind. 21
Pelargonium.—These highly cultivated plants seem eminently
liable to bud-variation. I will give only a few well-marked cases.
Gärtner has seen 22 a plant of P. zonale with a branch having white-
edged leaves, which remained constant for years, and bore flowers
of a deeper red than usual. Generally speaking, such branches
present little or no difference in their flowers: thus a writer 23
pinched off the leading shoot of a seedling P. zonale, and it threw
out three branches, which differed in the size and colour of their
leaves and stems; but on all three branches “the flowers were
identical,” except in being largest in the green-stemmed variety,
and smallest in that with variegated foliage: these three varieties
were subsequently propagated and distributed. Many branches,
and some whole plants, of a variety called compactum, which bears
orange-scarlet flowers, have been seen to produce pink flowers. 24
Hill's Hector, which is a pale red variety, produced a branch with
lilac flowers, and some trusses with both red and lilac flowers.
This apparently is a case of reversion, for Hill's Hector was a
seedling from a lilac variety 25 Of all Pelargoniums, Rollisson's
Unique seems to be the most sportive; its origin is not positively
known, but is believed to be from a cross. Mr. Salter, of Ham-
mersmith, states 26 that he has himself known this purple vari-
ety to produce the lilac, the rose-crimson or conspicuum, and the
red or coccineum varieties; the latter has also produced the rose
d'amour, so that altogether four varieties have originated by bud
variation from Rollisson's Unique. Mr. Salter remarks that these
four varieties "may now be considered as fixed, although they occa-
“sionally produce flowers of the original colour. This year coc-
2) Mr. W. Bell, Bot. Soc. of Edinburgh,
May, 1863
21 Revue Horticole,' quoted in 'Gard.
Chron.,'1845, p. 475.
22 Bastarderzeugung,' 1849, s. 76.
29 Jour. of Horticulture,' 1861, p. 336.
24 W. P. Ayres, in 'Gardener's Chron.,
1812, p. 791.
25 W. P. Ayres, idem.
28
Gardener's Chron.,' 1861, p. 968.

CHAP. XI.
455
FLOWERS.
"cineum has pushed flowers of three different colours, red, rose, and
"lilac, upon the same truss, and upon other trusses are flowers half
"red and half lilac.” Besides these four varieties, two other scarlet
Uniques are known to exist, both of which occasionally produce
lilac flowers identical with Rollisson's Unique ; 27 but one at least of
these did not arise through bud-variation, but is believed to be a
seedling from Rollisson's Unique.28 There are, also, in the trade 29
two other slightly different varieties, of unknown origin, of Rollis-
son's Unique: so that altogether we have a curiously complex case
of variation both by buds and seeds.30 An English wild plant, the
Geranium pratense, when cultivated in a garden, has been seen to
produce on the same plant both blue and white, and striped blue
and white flowers. 31
Chrysanthemum.- This plant frequently sports, both by its lateral
branches and occasionally by suckers. A seedling raised by Mr.
Salter has produced by bud-variation six distinct sorts, five different
in colour and one in foliage, all of which are now fixed. The
varieties which were first introduced from China were so exces-
sively variable, "that it was extremely difficult to tell which was
the original colour of the variety, and which was the sport.” The
same plant would produce one year only buff-coloured, and next
year only rose-coloured flowers; and then would change again, or
produce at the same time flowers of both colours. These fluctuat-
ing varieties are now all lost, and, when a branch sports into a new
variety, it can generally be propagated and kept true; but, as Mr.
Salter remarks, "every sport should be thoroughly tested in dif-
“ferent soils before it can be really considered as fixed, as many
“ have been known to run back when planted in rich compost ; but
“when sufficient care and time are expended in proving, there will
exist little danger of subsequent disappointment.” Mr. Salter in-
forms me that with all the varieties the commonest kind of bud-
variation is the production of yellow flowers, and, as this is the
primordial colour, these cases may be attributed to reversion. Mr.
Salter has given me a list of seven differently coloured chrysanthe-
mums, which have all produced branches with yellow flowers; but
66
27 Gardener's Chron.,' 1861, p. 945.
28 W. Paul, in 'Gardener's Chron.,'
1861, p. 968.
29 Idem, p. 945.
30 For other cases of bud-variation in
this same variety, see Gardener's
Chron.,' 1861, pp. 578, 600, 925. For
other distinct cases of bud-variation in
the genus Pelargonium, see Cottage
Gardener,' 1860, p. 194.
31 Rev. W. T. Bree, in Loudon's 'Gard.
Mag.,' vol. viii., 1882, p. 93.
32 The Chrysanthemum, its History
and Culture,' by J. Salter, 1865, p. 41,
&c.

456
CHAP. XI
BUD-VARIATION.
three of them have also sported into other colours. With any
change of colour in the flower, the foliage generally changes in a
corresponding manner in lightness or darkness.
Another Compositous plant, namely, Centauria cyanus, when cul-
tivated in a garden, not unfrequently produces on the same root
flowers of four different colours, viz., blue, white, dark-purple, and
particoloured.93 The flowers of Anthemis also vary on the same
plant.34
Roses.-Many varieties of the rose are known or are believed to
have originated by bud-variation.35 The common double moss-rose
was imported into England from Italy about the year 1735.36 Its
origin is unknown, but from analogy it probably arose from the
Provence rose (R. centifolia) by bud-variation ; for branches of the
common moss-rose have several times been known to produce Pro-
vence roses, wholly or partially destitute of moss: I have seen one
such instance, and several others have been recorded.37 Mr. Rivers
also informs me that he raised two or three roses of the Provence
class from seed of the old single moss-rose ; 38 and this latter kind
was produced in 1807 by bud-variation from the common moss-rose.
The white moss-rose was also produced in 1788 by an offset from
the common red moss-rose: it was at first pale blush-coloured, but
became white by continued budding. On cutting down the shoots
which had produced this white moss-rose, two weak shoots were
thrown up, and buds from these yielded the beautiful striped moss-
rose. The common moss-rose has yielded by bud-variation, besides
the old single red moss-rose, the old scarlet semi-double moss-rose,
and the sage-leaf moss-rose, which “has a delicate shell-like form,
and is of a beautiful blush-colour ; it is now (1852) nearly extinct." 39
A white moss-rose has been seen to bear a flower half white and
half pink.40 Although several moss-roses have thus certainly arisen
by bud-variation, the greater number probably owe their origin to
seed of moss-roses. For Mr. Rivers informs me that his seedlings
from the old single moss-rose almost always produced moss-roses ;
and the old single moss-rose was, as we have seen, the product by bud-
34 Bronn,
33 Bree, in Loudon's Gard. Mag.,'
vol. viii., 1832, p. 93.
Geschichte der Natur,' B.
ii, s. 123.
35 T. Rivers, 'Rose Amateur's Guide,'
1837, p. 4.
36 Mr. Shailer, quoted in Gardener's
Chron.,' 1848, p. 759.
37 Transact. Hort. Soc.' vol. iv.,
1822, p. 137; Gard. Chron.' 1842, p. 422.
38 See also Loudon's "Arboretum,'
vol. ii. p. 780.
39 All these statements on the origin
of the several varieties of the moss-
rose are given on the authority of Mr.
Shailer, who, together with his father,
was concerned in their original pro
pagation, in 'Gard. Chron.,' 1852, p.
759.
40 Gard. Chron.,' 1845, p. 564.

CHAP. XỈ.
457
FLOWERS.
variation of the double moss-rose originally imported from Italy.
That the original moss-rose was the product of bud-variation is pro-
bable, from the facts above given and from the moss-rose de Meaux
(also a var. of R. centifolia) 41 having appeared as a sporting branch
on the common rose de Meaux.
Prof. Caspary has carefully described the case of a six-year-old
white moss-rose, which sent up several suckers, one of which was
thorny, and produced red flowers, destitute of moss, exactly like
those of the Provence rose (R. centifolia): another shoot bore both
kinds of flowers and in addition longitudinally striped flowers. As
this white moss-rose had been grafted on the Provence rose, Prof.
Caspary attributes the above changes to the influence of the stock ;
but from the facts already given, and from others to be given, bud-
variation, with reversion, is probably a sufficient explanation.
Many other instances could be added of roses varying by buds.
The white Provence rose apparently thus originated. The double
and highly-coloured Belladonna rose has been known 44 to produce
by suckers both semi-double and almost single white roses; whilst
suckers from one of these semi-double white roses reverted to per-
fectly characterised Belladonnas. Varieties of the China rose pro-
pagated by cuttings in St. Domingo often revert after a year or two
into the old China rose. 15 Many cases have been recorded of roses
suddenly becoming striped or changing their character by seg-
ments: some plants of the Comtesse de Chabrillant, which is pro-
perly rose-coloured, were exhibited in 1862,46 with crimson flakes on
a rose ground. I have seen the Beauty of Billiard with a quarter
and with half the flower almost white. The Austrian bramble (R.
Tutea) not rarely 47 produces branches with pure yellow flowers; and
Prof. Henslow has seen exactly half the flower of a pure yellow,
and I have seen narrow yellow streaks on a single petal, of which
the rest was of the usual copper colour.
The following cases are highly remarkable. Mr. Rivers, as I am
informed by him, possessed a new French rose with delicate smooth
shoots, pale glaucous-green leaves, and semi-double pale flesh-coloured
flowers striped with dark red; and on branches thus characterised
there suddenly appeared, in more than one instance, the famous old
rose called the Baronne Prevost, with its stout thorny shoots, and
41 Transact. Hort. Soc.,' vol, ii. p.
242.
12 Schriften der Phys. Okon. Gesell.
zu Königsberg,' Feb. 3, 1865, s. 4. See
also Dr. Caspary's paper in Transac-
tions of the Hort. Congress of Amster-
dam,' 1865.
43 Gard. Chron.,'1852, p. 759.
44 Transact. Hort. Soc.,' vol. ii. p.
242.
45 Sir R. Schomburgk, Proc. Linn.
Soc. Bot.,' vol.ii. p. 132.
46 Gard. Chron.,' 1862, p. 619.
47 Hopkirk's 'Flora Anomala,' p. 167.
20

458
CHẤP. XI,
BUD-VARIATION.
immense, uniformly and richly coloured, double flowers; so that in
this case the shoots, leaves, and flowers, all at once changed their
character by bud-variation. According to M. Verlot 48 a variety call-
ed Rosa Cannabifolia, which has peculiarly shaped leaflets, and dif
fers from every member of the family in the leaves being opposite
instead of alternate, suddenly appeared on a plant of R. alba in the
gardens of the Luxembourg. Lastly, "a running shoot” was ob-
served by Mr. H. Curtis 40 on the old Aimée Vibert Noisette, and he
budded it on Celine; thus a climbing Aimée Vibert was first pro-
duced and afterwards propagated.
Dianthus.—It is quite common with the Sweet William (D. bar-
batus) to see differently coloured flowers on the same root; and I
have observed on the same truss four differently coloured and shaded
flowers. Carnations and pinks (D.caryophyllus, &c.) occasionally vary
by layers; and some kinds are so little certain in character that they
are called by floriculturists "catch-flowers." 50 Mr. Dickson has ably
discussed the "running" of particoloured or striped carnations, and
says it cannot be accounted for by the compost in which they are
grown: "layers from the same clean flower would come part of them
“clean and part foul, even when subjected to precisely the same
"treatment; and frequently one flower alone appears influenced by
"the taint, the remainder coming perfectly clean." 51 This running
of the parti-coloured flowers apparently is a case of reversion by buds
to the original uniform tint of the species.
I will briefly mention some other cases of bud-variation to show
how many plants belonging to many orders have varied in their
flowers; numerous cases might be added. I have seen on a snap-
dragon (Antirrhinum majus) white, pink, and striped flowers on the
same plant, and branches with striped flowers on a red-coloured va-
riety. On a double stock (Matthiola incana) I have seen a branch
bearing single flowers; and on a dingy-purple, double variety of the
wall-flower (Cheiranthus cheiri) a branch which had reverted to the
ordinary copper colour. On other branches of the same plant, some
flowers were exactly divided across the middle, one half being pur-
ple and the other coppery; but some of the smaller petals towards
the centre of these same flowers were purple longitudinally streaked
with coppery colour, or coppery streaked with purple. A Cyclamen 52
has been observed to bear white and pink flowers of two forms, the
one resembling the Persicum strain, and the other the Coum strain.
48 Sur la Production et la Fixation
des Variétés,' 1865, p. 4.
49 Journal of Horticulture, March,
1865, p. 233.
60 Gard. Chron.,' 1843, p. 135.
51 Ibid., 1842, p. 55.
62 Gard. Chron.,' 1867, p. 235.

CHAP. XI.
459
LEAVES AND SHOOTS.
Oenothera biennis has been seen 53 bearing flowers of three different
colours. The hybrid Gladiolus colvillii occasionally bears uniformly
coloured flowers, and one case is recorded 54 of all the flowers on a
plant thus changing colour. A Fuchsia has been seen 55 bearing
two kinds of flowers. Mirabilis jalapa is eminently sportive, some-
times bearing on the same root pure red, yellow, and white flowers,
and others striped with various combinations of these three colours. 56
The plants of the Mirabilis which bear such extraordinarily variable
flowers, in most, probably in all cases, owe their origin, as shown
by Prof. Lecoq, to crosses between differently-coloured varieties.
Leaves and Shoots.-Changes, through bud variation, in fruits and
flowers have hitherto been treated of, but incidentally some remark-
able modifications in the leaves and shoots of the rose and Cistus,
and in a lesser degree in the foliage of the Pelargonium and Chry-
santhemum, have been noticed. I will now add a few more cases on
variation in leaf-buds. Verlot 57 states that on Aralia trifoliata,
which properly has leaves with three leaflets, branches bearing sim-
ple leaves of various forms frequently appear; these can be propa-
gated by buds or grafting, and have given rise, as he states, to se-
veral nominal species.
With respect to trees, the history of but few of the many varieties
with curious or ornamental foliage is known; but several probably
have originated by bud-variation. Here is one case ;- An old ash-
tree (Fraxinus excelsior) in the grounds of Necton, as Mr. Mason
states," for many years has had one bough of a totally different
character to the rest of the tree, or of any other ash-tree which I
have seen; being short-jointed and densely covered with foliage."
It was ascertained that this variety could be propagated by grafts.68
The varieties of some trees with cut leaves, as the oak-leaved labur-
num, the parsley-leaved vine, and especially the fern-leaved beech,
are apt to revert by buds to the common form.59 The fern-like leaves
of the beech sometimes revert only partially, and the branches dis-
play here and there sprouts bearing common leaves, fern-like, an
variously shaped leaves. Such cases differ but little from the so-
called heterophyllous varieties, in which the tree habitually bears
Fécondation,' 1862, p. 303.
57 Des Variétés,'1865, p. 5.
63 W. Mason, in 'Gard. Chron.,'1843,
53 Gärtner, 'Bastarderzeugung,' s. 305.
54 Mr. D. Beaton, in Cottage Garden-
er,' 1860, p. 250.
55 Gard. Chron.,' 1850, p. 536.
56 Braun, 'Ray Soc. Bot. Mem.,' 1853,
p. 315; Hopkirk's 'Flora Anomala,' p.
164; Lecoq, 'Géograph. Bot. de l'Eu-
rope,' tom. iii., 1854, p. 405; and 'De la
p. 878.
59 Alex. Braun, 'Ray Soc. Bot. Mem.,'
1953, p. 315; Gard. Chron.,' 1841, p.
329.

460
CHAP. XL
BUD-VARIATION
leaves of various forms; but it is probable that most heterophyllous
trees have originated as seedlings. There is a sub-variety of the
weeping willow with leaves rolled up into a spiral coil; and Mr.
Masters states that a tree of this kind kept true in his garden for
twenty-five years, and then threw out a single upright shoot bearing
flat leaves.60
I have often noticed single twigs and branches on beech and
other trees with their leaves fully expanded before those on the
other branches had opened ; and as there was nothing in their ex-
posure or character to account for this difference, I presume that
they had appeared as bud-variations, like the early and late fruit-
maturing varieties of the peach and nectarine.
Cryptogamic plants are liable to bud-variation, for fronds on the
same fern are often seen to display remarkable deviations of struc-
ture. Spores, which are of the nature of buds, taken from such ab-
normal fronds, reproduce, with remarkable fidelity, the same variety,
after passing through the sexual stage.62
With respect to colour, leaves often become by bud-variation zoned,
blotched, or spotted with white, yellow, and red; and this occa-
sionally occurs even with plants in a state of nature. Variegation,
however, appears still more frequently in plants produced from seed;
even the cotyledons or seed-leaves being thus affected.62 There
have been endless disputes whether variegation should be consider-
ed as a disease. In a future chapter we shall see that it is much in-
fluenced, both in the case of seedlings and of mature plants, by the
nature of the soil. Plants which have become variegated as seed-
lings, generally transmit their character by seed to a large propor-
tion of their progeny ; and Mr. Salter has given me a list of eight
genera in which this occurred.63 Sir F. Pollock has given me more
precise information: he sowed seed from a variegated plant of Bal-
lota nigra, which was found growing wild, and thirty per cent. of
the seedlings were variegated ; seed from these latter being sown,
sixty per cent. came up variegated. When branches become varie-
gated by bud-variation, and the variety is attempted to be propa-
gated by seed, the seedlings are rarely variegated; Mr. Salter found
this to be the case with plants belonging to eleven genera, in which
the greater number of the seedlings proved to be green-leaved ; yet
a few were slightly variegated, or were quite white, but none were
60 Dr. M. T. Masters, 'Royal Institu-
tion Lecture,' March 16, 1860.
61 See Mr. W. K. Bridgman's curious
paper in 'Annals and Mag. of Nat. Hist.,'
December, 1861; also Mr. J. Scott, Bot.
Soc. Edinburgh,' June 12, 1862.
62 Journal of Horticulture,' 1861, p..
336; Verlot, 'Des Variétés,' p. 76.
63 See also Verlot, Des Variétés,' p.

CHAP, XI.
461
BY SUCKERS, TUBERS, AND BULBS.
1964
worth keeping Variegated plants, whether originally produced
from seeds or buds, can generally be propagated by budding, graft-
ing, &c. ; but all are apt to revert by bud-variation to their ordinary
foliage. This tendency, however, differs much in the varieties of
even the same species; for instance, the golden-striped variety of
Euonymus Japonicus" is very liable to run back to the green-leaved,
while the silver-striped variety hardly ever changes. I have seen
a variety of the holly, with its leaves having a central yellow patch,
which had everywhere partially reverted to the ordinary foliage, so
that on the same small branch there were many twigs of both kinds.
In the pelargonium, and in some other plants, variegation is general-
ly accompanied by some degree of dwarfing, as is well exemplified
in the “Dandy" pelargonium. When such dwarf varieties sport
back by buds or suckers to the ordinary foliage, the dwarfed sta-
ture sometimes still remains. It is remarkable that plants propa-
gated from branches which have reverted from variegated to plain
leaves 66 do not always (or never, as one observer asserts) perfectly
resemble the original plain-leaved plant from which the variegated
branch arose : it seems that a plant, in passing by bud-variation
from plain leaves to variegated, and back again from variegated to
plain, is generally in some degree affected so as to assume a slightly
different aspect.
Bud-variation by Suckers, Tubers, and Bulbs.-All the cases hither-
to given of bud-variation in fruits, flowers, leaves, and shoots, have
been confined to buds on the stems or branches, with the exception
of a few cases incidentally noticed of varying suckers in the rose,
pelargonium, and chrysanthemum. I will now give a few instances
of variation in subterranean buds, that is, by suckers, tubers, and
bulbs; not that there is any essential difference between buds above
and beneath the ground. Mr. Salter informs me that two varie-
gated varieties of Phlox originated as suckers; but I should not
have thought these worth mentioning, had not Mr. Salter found,
after repeated trials, that he could not propagate them by "root-
joints," whereas, the variegated Tussilago farfara can thus be safe-
ly propagated ; 67 but this latter plant may have originated as a varie-
gated seedling, which would account for its greater fixedness of
64 Gard. Chron.,' 1814, p. 86.
65 Ibid., 1861, p. 968.
68 Ibid., 1861, p. 433. Cottage Gar-
dener,' 1560, p. 2.
67 M. Lemoine (quoted in Gard.
Chron.,'1867, p. 74) has lately observed
that the Symphitum with variegated
leaves cannot be propagated by division
of the roots. He also found that out of
500 plants of a Phlox with striped flow-
ers, which had been propagated by root-
division, only seven or eight produced
striped flowers. See also, on striped Pe-
largoniums, 'Gard. Chron.' 1867, p. 1000.

462
CHAP, XI.
BUD-VARIATION
character. The Barberry (Berberis vulgaris) offers an analogous
case; there is a well-known variety with seedless fruit, which can be
propagated by cuttings or layers; but suckers always revert to the
common form, which produces fruit containing seeds.68 My father
repeatedly tried this experiment, and always with the same result.
Turning now to tubers: in the common Potato (Solanum tubero-
sum) a single bud or eye sometimes varies and produces a new va-
riety; or, occasionally, and this is a much more remarkable circum-
stance, all the eyes in a tuber vary in the same manner and at the
same time, so that the whole tuber assumes a new character. For
instance, a single eye in a tuber of the old Forty-fold potato, which
is a purple variety, was oberved 9 to become white; this eye was
cut out and planted separately, and the kind has since been largely
propagated. Kemp's Potato is properly white, but a plant in Lan-
cashire produced two tubers which were red, and two which were
white; the red kind was propagated in the usual manner by eyes,
and kept true to its new colour, and, being found a more productive
variety, soon became widely known under the name of Taylor's
Forty-fold.' The Old Forty-fold potato, as already stated, is a purple
variety; but a plant long cultivated on the same ground produced,
not as in the case above given a single white eye, but a whole white
tuber, which has since been propagated and keeps true. Several
cases have been recorded of large portions of whole rows of potatoes
slightly changing their character.72
Dahlias propagated by tubers under the hot climate of St. Do-
mingo vary much; Sir R. Schomburgk gives the case of the “But-
terfly variety," which the second year produced on the same plant
“double and single flowers; here white petals edged with maroon;
“there of a uniform deep maroon.
Mr. Bree also mentions a
plant “which bore two different kinds of self-coloured flowers, as
“well as a third kind which partook of both colours beautifully in-
"termixed.” 74 Another case is described of a dahlia with purple
flowers which bore a white flower streaked with purple."
Considering how long and extensively many Bulbous plants have
been cultivated, and how numerous are the varieties produced from
seed, these plants have not varied so much by offsets,—that is, by
the production of new bulbs,--as might have been expected. With
73
75
68 Anderson's 'Recreations in Agricul-
ture,' vol. v. p. 152.
69 Gard. Chron.,' 1857, p. 662.
70 Ibid., 1841, p. 814.
71 Ibid., 1857, p. 613.
72 Ibid., 1857, p. 679. See also Phil-
lips, 'Hist. of Vegetables,' vol. ii. p. 91,
for other and similar accounts.
73 Journal of Proc. Linn. Soc.,' vol.
ii. Botany, p. 132.
74 Loudon's Gard. Mag.,' vol. viii.,
1832, p. 94.
75 Gard. Chron.,' 1950, p. 536; and
1842, p. 729.

CHAP. XI. BY SUCKERS, TUBERS, AND BULBS.
463
the Hyacinth a case has been recorded of a blue variety which for
three successive years gave offsets which produced white flowers
with a red centre.76 Another hyacinth has been described" as
bearing on the same truss a perfectly pink and a perfectly blue
flower.
Mr. John Scott informs me that in 1862 Imatophyllum miniatum,
in the Botanic Gardens of Edinburgh, threw up a sucker which dif-
fered from the normal form, in the leaves being two-ranked instead
of four-ranked. The leaves were also smaller, with the upper sur-
face raised instead of being channelled.
In the propagation of Tulips, seedlings are raised, called selfs or
breeders, which consist of one plain colour on a white or yellow
“bottom. These, being cultivated on a dry and rather poor soil,
" become broken or variegated and produce new varieties. The
" time that elapses before they break varies from one to twenty
"years or more, and sometimes this change never takes place.” 78
The various broken or variegated colours which give value to all
tulips are due to bud-variation; for although the Bybloemens and
some other kinds have been raised from several distinct breeders, yet
all the Baguets are said to have come from a single breeder or seed-
ling. This bud-variation, in accordance with the views of MM. Vil-
morin and Verlot,79 is probably an attempt to revert to that uni-
form colour which is natural to the species. A tulip, however,
which has already become broken, when treated with too strong
manure, is liable to flush or lose by a second act of reversion its
variegated colours. Some kinds, as Imperatrix Florum, are much
more liable than others to flushing; and Mr. Dickson maintains 50
that this can no more be accounted for than the variation of any
other plant. He believes that English growers, from care in choos-
ing seed from broken flowers instead of from plain flowers, have to
a certain extent diminished the tendency in flowers already broken
to flushing or secondary reversion.
During two consecutive years all the early flowers in a bed of
Tigridia conchiflora el resembled those of the old T. pavonia ; but
the later flowers assumed their proper colour of fine yellow spotted
with crimson. An apparently authentic account has been published 62
of two forms of Hemerocallis, which have been universally con-
p. 63.
80 Gard. Chron.,'1811, p. 752; 1842,
76 Des Jacinthes,' &c., Amsterdam,
1768, p. 122.
77 Gard. Chron.,' 1945, p. 212.
78 Loudon's Encyclop. of Gardening,
p. 55.
p. 1024.
81 Gard. Chron., '1849, p. 565.
82 Transact. Linn. Soc.,' vol. ii. p.
354.
79 Production des Variétés,' 1865,

464
CHAP. X1.
BUD-VARIATION.
sidered as distinct species, changing into each other; for the roots
of the large-flowered tawny H. fulva, being divided and planted in
a different soil and place, produced the small-flowered yellow H.
flava, as well as some intermediate forms. It is doubtful whether
such cases as these latter, as well as the "flushing” of broken tulips
and the “running” of particoloured carnations,—that is, their more
or less complete return to a uniform tint, ought to be classed under
bud-variation, or ought to be retained for the chapter in which I
treat of the direct action of the conditions of life on organic beings.
These cases, however, have this much in common with bud-varia-
tion, that the change is effected through buds and not through
seminal reproduction. But, on the other hand, there is this differ-
ence—that in ordinary cases of bud-variation, one bud alone changes,
whilst in the foregoing cases all the buds on the same plant were
modified together; yet we have an intermediate case, for with the
potato all the eyes in one tuber alone simultaneously changed their
character.
I will conclude with a few allied cases, which may be ranked
either under bud-variation, or under the direct action of the condi-
tions of life. When the common Hepatica is transplanted from its
native woods, the flowers change colour, even during the first year.83
It is notorious that the improved varieties of the Heartsease (Viola
tricolor) when transplanted often produce flowers widely different in
size, form, and colour: for instance, I transplanted a large uniformly-
coloured dark purple variety, whilst in full flower, and it then pro-
duced much smaller, more elongated flowers, with the lower petals
yellow; these were succeeded by flowers marked with large purple
spots, and ultimately, towards the end of the same summer, by the
original large dark purple flowers. The slight changes which some
fruit-trees undergo from being grafted and regrafted on various
stocks,84 were considered by Andrew Knight 85 as closely allied to
“sporting branches," or bud-variations. Again, we have the case of
young fruit-trees changing their character as they grow old ; seed-
ling pears, for instance, lose with age their spines and improve in
the flavour of their fruit. Weeping birch-trees, when grafted on the
common variety, do not acquire a perfect pendulous habit until they
83 Godron, De l'Espèce,' tom. ii. p.
84.
84 M. Carrière has lately described,
in the 'Révue Horticole' (Dec. 1, 1866,
p. 457), an extraordinary case. He twice
inserted grafts of the Aria vestita on
thorn-trees (épines) growing in pots;
and the grafts, as they grew, produced
shoots with bark, buds, leaves, petioles,
petals, and flower-stalks all widely dif-
ferent from those of the Aria. The
grafted shoots were also much hardier,
and flowered earlier, than those on the
ungrafted Aria.
85 Transact. Hort. Soc.,' vol. ii. p.
160.

CHAP. XI.
465
ANOMALOUS REPRODUCTION, ETC.
grow old : on the other hand, I shall hereafter give the case of some
weeping ashes which slowly and gradually assumed an upright
habit of growth. All such changes, dependent on age, may be
compared with the changes, alluded to in the last chapter, which
many trees naturally undergo; as in the case of the Deodar and
Cedar of Lebanon, which are unlike in youth and closely resemble
each other in old age; and as with certain oaks, and with some
varieties of the lime and hawthorn.86
Before giving a summary on Bud-variation I will dis-
cuss some singular and anomalous cases, which are more
or less closely related to this same subject. I will begin
with the famous case of Adam's laburnum or Cytisus
Adami, a form or hybrid intermediate between two very
distinct species, namely, C. laburnum and purpureus,
the common and purple laburnum; but as this tree has
often been described, I will be as brief as I can.
Throughout Europe, in different soils and under different climates,
branches on this tree have repeatedly and suddenly reverted to both
parent-species in their flowers and leaves. To behold mingled on
the same tree tufts of dingy-red, bright yellow, and purple flowers,
borne on branches having widely different leaves and manner of
growth, is a surprising sight. The same raceme sometimes bears
two kinds of flowers; and I have seen a single flower exactly di-
vided into halves, one side being bright yellow and the other pur-
ple; so that one half of the standard-petal was yellow and of larger
size, and the other half purple and smaller. In another flower the
whole corolla was bright yellow, but exactly half the calyx was
purple. In another, one of the dingy-red wing-petals had a bright
yellow narrow stripe on it, and lastly, in another flower, one of the
stamens, which had become slightly foliaceous, was half yellow
and half purple; so that the tendency to segregation of character
or reversion affects even single parts and organs.87 The most re-
markable fact about this tree is that in its intermediate state, even
when growing near both parent-species, it is quite sterile; but when
the flowers become pure yellow or pure purple they yield seed. I
believe that the pods from the yellow flowers yield a full comple-
88 For the cases of oaks see Alph. De
Candolle in Bibl. Univers.,' Geneva,
Nov. 1862; for limes, &c., Loudon's
"Gard, Mag.,' vol. xi, 1835, p. 503,
87 For analogous facts, see Braun,
"Rejuvenescence,' in "Ray Soc. Bot.
Mem.,' 1853, p. 320; and 'Gard. Chron.,'
1842, p. 397.
20*

466
CEAP. XI.
ANOMALOUS MODES
ment of seed; they certainly yield a large number. Two seedlings
raised by Mr. Herbert from such seed exhibited a purple tinge on.
the stalks of their flowers; but several seedlings raised by myself
resembled in every character the common laburnum, with the ex-
ception that some of them had remarkably long racemes: these
seedlings were perfectly fertile. That such purity of character and
fertility should be suddenly reacquired from so hybridized and ste-
rile a form is an astonishing phenomenon. The branches with pur-
ple flowers appear at first sight exactly to resemble those of 0.
purpureus; but on careful comparison I found that they differed
from the pure species in the shoots being thicker, the leaves a little
broader, and the flowers slightly shorter, with the corolla and calyx
less brightly purple: the basal part of the standard-petal also plain-
ly showed a trace of the yellow stain. So that the flowers, at least
in this instance, had not perfectly recovered their true character;
and in accordance with this, they were not perfectly fertile, for
many of the pods contained no seed, some produced one, and very
few contained as many as two seeds; whilst numerous pods on a
tree of the pure C. purpureus in my garden contained three, four,
and five fine seeds. The pollen, moreover, was very imperfect, a
multitude of grains being small and shrivelled ; and this is a singu-
lar fact; for, as we shall immediately see, the pollen-grains in the
dingy-red and sterile flowers on the parent-tree, were, in external
appearance, in a much better state, and included very few shrivelled
grain. Although the pollen of the reverted purple flowers was in
so poor a condition, the ovules were well-formed, and, when ma-
ture, germinated freely with me. Mr. Herbert also raised plants
from seeds of the reverted purple flowers, and they differed very
little from the usual state of C. purpureus ; but this expression
shows that they had not perfectly recovered their proper character.
Prof. Caspary has examined the ovules of the dingy-red and
sterile flowers in several plants of C. adami on the Continent,' and
finds them generally monstrous. In three plants examined by me
in England, the ovules were likewise monstrous, the nucleus vary-
ing auch in shape, and projecting irregularly beyond the proper
coats. The pollen-grains, on the other hand, judging from their
external appearance, were remarkably good, and readily protruded
their tubes. By repeatedly counting, under the microscope, the
proportional number of bad grains, Prof. Caspary ascertained that
only 2-5 per cent. were bad, which is a less proportion than in the
88 Journal of Hort. Soc.,' vol. ii.,
1947, p. 100.
89 See Transact. of Ilort. Congress
of Amsterdam,' 1865; but I owe most
of the following information to Prof. Cas-
pary's letters.

CHAP. XI. OF REPRODUCTION AND VARIATION,
467
pollen of three pure species of Cytisus in their cultivated state, viz.
.C. purpureus, laburnum, and alpinus. Although the pollen of C.
adami is thus in appearance good, it does not follow, according
to M. Naudin's observations " on Mirabilis, that it would be func-
tionally effective. The fact of the ovules of C. adami being mon-
strous, and the pollen apparently sound, is all the more remarka-
ble, because it is opposed to what usually occurs not only with
most hybrids, 91 but with two hybrids in the same genus, namely in
C. purpureo-elongatus, and C. alpino-laburnum. In both these by-
brids, the ovules, as observed by Prof. Caspary and myself, were
well-formed, whilst many of the pollen-grains were ill-formed ; in
the latter hybrid 20-3 per cent., and in the former no less than 84:8
per cent. of the grains were ascertained by Prof. Caspary to be bad.
This unusual condition of the male and female reproductive ele-
ments in C. adami has been used by Prof. Caspary as an argument
against this plant being considered as an ordinary hybrid produced
from seed; but we should remember that with hybrids the ovules
have not been examined nearly so frequently as the pollen, and
they may be much oftener imperfect than is generally supposed.
Dr. E. Bornet, of Antibes, informs me (through Mr. J. Traherne Mog-
gridge) that with hybrid Cisti the ovarium is frequently deformed,
the ovules being in some cases quite absent, and in other cases in
capable of fertilisation.
Several theories have been propounded to account for the origin
of C. adami, and for the transformations which it undergoes. These
transformations have been attributed by some authors to simple bud-
variation ; but considering the wide difference between C. laburnum
and purpureus, both of which are natural species, and considering
the sterility of the intermediate form, this view may be summarily
rejected. We shall presently see that, with hybrid plants, two
different embryos may be developed within the same seed and co-
here, and it has been supposed that C. adami might have thus
originated. It is known that when a plant with variegated leaves
is budded on a plain stock, the latter is sometimes affected, and it is
believed by some that the laburnum has been thus affected. Thus
Mr. Purser states 92 that a common laburnum-tree in his garden,
into which three grafts of the Cytisus purpureus had been inserted,
gradually assumed the character of C. adami; but more evidence
90 Nouvelles Archives du Muséum,'
tom. i. p. 143.
91 See on this head, Naudin, idem, p.
141
92 The statement is believed by Dr.
Lindley in 'Gard. Chron.,' 1857, pp. 332,
400.

468
CRAP. XI.
ANOMALOUS MODES
and copious details would be requisite to make so extraordinary a
statement credible.
Many authors maintain that C. adami is a hybrid produced in
the common way by seed, and that it has reverted by buds to its
two parent-forms. Negative results are of little value ; but Reis-
seck, Caspary, and I myself, tried in vain to cross C. laburnum and
purpureus; when I fertilised the former with pollen of the latter,
I had the nearest approach to success, for pods were formed, but in
sixteen days after the withering of the flowers they fell off. Never-
theless, the belief that C. adami is a spontaneously produced hybrid
between these two species is strongly supported by the fact that hy-
brids between these species and two others have spontaneously arisen.
In a bed of seedlings from C. elongatus, which grew near to C. pur-
pureus, and was probably fertilised by it, through the agency of in-
sects (for these, as I know by experiment, play an important part in
the fertilisation of the laburnum), the sterile hybrid C. purpureo-
elongatus appeared.93 Thus, also, Waterer's laburnum, the C. alpino-
laburnum, 94 spontaneously appeared, as I am informed by Mr. Water-
er, in a bed of seedlings.
On the other hand, we have a clear and distinct account given by
M. Adam, who raised the plant, to Poiteau," showing that C. adami
is not an ordinary hybrid. M. Adam inserted in the usual manner
a shield of the bark of C. purpureus into a stock of C. laburnum; and
the bud lay dormant, as often happens, for a year; the shield then
produced many buds and shoots, one of which grew more upright
and vigorous with larger leaves than the shoots of C. purpureus, and
was consequently propagated. Now it deserves especial notice that
these plants were sold by M. Adam, as a variety of C. purpureus,
before they had flowered ; and the account was published by Poi-
teau after the plants had flowered, but before they had exhibited
their remarkable tendency to revert into the two parent-species.
08 Braun, in . Bot. Mem. Ray Soc.,'
1953, p. xxiii.
94 This hybrid has never been de-
scribed. It is exactly intermediate in
foliage, time of flowering, dark striæ at
the base of the standard petal, hairiness
of the ovarium, and in almost every
other character, between C. laburnum
and alpinus; but it approaches the
former species more nearly in colour,
and exceeds it in the length of the
racemes. We have before seen that 20:3
per cent. of its pollen-grains are ill-form-
ed and worthless. My plant, though
growing not above thirty or forty yards
from both parent-species, during some
seasons yielded no good seeds; but in
1866 it was unusually fertile, and its
long racemes produced from one to occa-
sionally even four pods. Many of the
pods contained no good seeds, but gene-
rally they contained a single apparently
good seed, sometimes two, and in one
case three seeds. Some of the seeds ger-
minated.
95 Annales de la Soc. de Hort. de
Paris,' tom. vii., 1830, p. 93.

CHAP. XI. OF REPRODUCTION AND VARIATION.
469
So that there was no conceivable motive for falsification, and it is
difficult to see how there could have been any error. If we admit
as true M. Adam's account, we must admit the extraordinary fact
that two distinct species can unite by their cellular tissue, and sub-
sequently produce a plant bearing leaves and sterile flowers inter-
mediate in character between the scion and stock, and producing
buds liable to reversion; in short, resembling in every important
respect a hybrid formed in the ordinary way by seminal reproduc-
tion. Such plants, if really thus formed, might be called graft-hy-
brids.
I will now give all the facts which I have been able to collect
illustrative of the above theories, not for the sake of merely throw-
ing light on the origin of C. adami, but to show in how many extra-
ordinary and complex methods one kind of plant may affect another,
generally in connection with bud-variation. The supposition that
either C. laburnum or purpureus produced by ordinary bud-variation
the intermediate and the other form, may, as already remarked, be
absolutely excluded, from the want of any evidence, from the great
amount of change thus implied, and from the sterility of the inter-
mediate form. Nevertheless such cases as nectarines suddenly ap-
pearing on peach-trees, occasionally with the fruit half-and-half in
nature,-moss-roses appearing on other roses, with the flowers divid-
ed into halves, or striped with different colours --and other such
cases, are closely analogous in the result produced, though not in
origin, with the case of O. adami.
A distinguished botanist, Mr. G. H. Thwaites 96 has recorded a re-
markable case of a seed from Fuchsia coccinea fertilised by F. ful-
gens, which contained two embryos, and was “a true vegetable
twin.” The two plants produced from the two embryos were
tremely different in appearance and character," though both resem-
bled other hybrids of the same parentage produced at the same time.
These twin plants “ were closely coherent, below the two pairs of
“cotyledon-leaves, into a single cylindrical stem, so that they had
“subsequently the appearance of being branches on one trunk.”
Had the two united stems grown up to their full height, instead of
dying, a curiously mixed hybrid would have been produced ; but
even if some of the buds had subsequently reverted to both parent-
forms, the case, although more complex, would not have been strict-
ly analogous with that of C. adami. On the other hand, a mongrel
melon described by Sageret ºr perhaps did thus originate ; for the
ex-
96 Annals and Mag. of Nat. Hist.,'
March, 1848.
97 Pomologie Physiolog.,' 1830, p.
126.

470
CHAP. XI.
ANOMALOUS MODES
two main branches, which arose from two cotyledon-buds, produced
very different fruit,-on the one branch like that of the pater-
nal variety, and on the other branch to a certain extent like that of
the maternal variety, the melon of China.
The famous bizzarria Orange offers a strictly parallel case to that
of Cytisus adami. The gardener who in 1644 in Florence raised this
tree, declared that it was a seedling which had been grafted ; and
after the graft had perished, the stock sprouted and produced the
bizzarria. Gallesio, who carefully examined several living specimens
and compared them with the description given by the original de-
scriber P. Nato, 98 states that the tree produces at the same time
leaves, flowers, and fruit, identical with the bitter orange and with
the citron of Florence, and likewise compound fruit with the two
kinds either blended together, both externally and internally, or
segregated in various ways. This tree can be propagated by cut-
tings, and retains its diversified character. The so-called trifacial
orange of Alexandria and Smyrna º' resembles in its general nature
the bizzarria, but differs from it in the sweet orange and citron being
blended together in the same fruit, and separately produced on the
same tree: nothing is known of its origin. In regard to the bizzar-
ria, many authors believe that it is a graft-hybrid ; Gallesio on the
other hand thinks that it is an ordinary hybrid, with the habit of
partially reverting by buds to the two parent-forms; and we have
seen in the last chapter that the species in this genus often cross
spontaneously.
Here is another analogous, but doubtful case. A writer in the
* Gardener's Chronicle 100 states that an Æsculus rubicunda in his
garden yearly produced on one of its branches "spikes of pale yellow
flowers, smaller in size and somewhat similar in colour to those of
Æ. flava." If as the editor believes Æsculus rubicunda is a hybrid
descended on one side from Æ. flava, we have a case of partial re-
version to one of the parent-forms. If, as some botanists maintain,
Æ. rubicunda is not a hybrid, but a natural species, the case is one
of simple bud-variation.
The following facts show that hybrids produced from seed in the
ordinary way, certainly sometimes revert by buds to their parent-
forms. Hybrids between Tropæolum minus and majus 101 at first
98 Gallesio, Gli Agrumi dei Giard.
Bot. Agrar, di Firenze,' 1839, p. 11. In
his Traité du Citrus,' 1811, p. 146, he
speaks as if the compound fruit con-
sisted in part of lemons, but this appa-
rently was a mistake.
99 Gard. Chron.,' 1855, p. 628. See
also Prof. Caspary, in Transact. Hort.
Congress of Amsterdam,' 1865.
100 Gard. Chron.,' 1851, p. 406.
101 Gärtner, Bastarderzeugung,' s.
549. It is, however, doubtful whether
these plants should be ranked as species
or varieties.

CHAP. XI.
471
OF REPRODUCTION AND VARIATION.
produced flowers intermediate in size, colour, and structure between
their two parents; but later in the season some of these plants pro-
duced flowers in all respects like those of the mother-form, mingled
with flowers still retaining the usual intermediate condition. А
hybrid Cereus between C. speciosissimus and phyllanthus, 102 plants
which are widely different in appearance, produced for the first three
years angular, five-sided stems, and then some flat stems like those
of C. phyllanthus. Kölreuter also gives cases of hybrid Lobelias and
Verbascums, which at first produced flowers of one colour, and later
in the season flowers of a different colour.103 Naudin 104 raised forty
hybrids from Datura lævis fertilised by D. stramonium ; and three
of these hybrids produced many capsules, of which a half, or quarter,
or lesser segment was smooth and of small size like the capsule of the
pure D. lævis, the remaining part being spinose and of larger size like
the capsule of the pure D. stramonium : from one of these composite
capsules, plants were raised which perfectly resembled both parent-
forms.
Turning now to varieties. A seedling apple, conjectured to be
of crossed parentage, has been described in France, 105 which bears
fruit, with one half larger than the other, of a red colour, acid taste,
and peculiar odour; the other side being greenish-yellow and very
sweet: it is said scarcely ever to include perfectly developed seed.
I suppose that this is not the same tree with that which Gaudi-
chaud 106 exhibited before the French Institute, bearing on the same
branch two distinct kinds of apples, one a reinette rouge, and the
other like a reinette canada jaunâtre : this double-bearing variety
can be propagated by grafts, and continues to produce both kinds;
its origin is unknown. The Rev. J. D. La Touche sent me a coloured
drawing of an apple which he brought from Canada, of which half,
surrounding and including the whole of the calyx and the insertion
of the footstalk, is green, the other half being brown and of the na-
ture of the pomme gris apple, with the line of separation between
the two halves exactly defined. The tree was a grafted one, and
Mr. La Touche thinks that the branch which bore this curious apple
sprung from the point of junction of the graft and stock: had this
fact been ascertained, the case would probably have come into the
small class of graft-hybrids presently to be given. But the branch
may have sprung from the stock, which no doubt was a seedling.
102 Gärtner, 'Bastarderz.,', s. 550. 105 L'Hermès, Jan. 14, 1837, quoted
103 Journal de Physique,' tom. xxiii., in Loudon's Gard. Mag.,' vol. xiii. p.
1783, p. 100. Act. Acad. St. Peters- 230.
burgh,' 1781, part. i. p. 249.
106 Comptes Rendus,' tom. xxxiv.,
104 Nouvelles Archives du Muséum, 1852, p. 746.
tom. i. p. 49.

472
CHAP. XI.
ANOMALOUS MODES
Prof. H. Lecoq, who has made a great number of crosses between
the differently coloured varieties of Mirabilis japala,107 finds that in
the seedlings the colours rarely combine, but form distinct stripes;
or half the flower is of one colour and half of a different colour. Some
varieties regularly bear flowers striped with yellow, white, and
red; but plants of such varieties occasionally produce on the same
root branches with uniformly coloured flowers of all three tints, and
other branches with half-and-half coloured flowers and others with
marbled flowers. "Gallesio 108 crossed reciprocally white and red car-
nations, and the seedlings were striped ; but some of the striped
plants also bore entirely white and entirely red flowers. Some of
these plants produced one year red flowers alone, and in the follow-
ing year striped flowers; or conversely, some plants, after having
borne for two or three years striped flowers, would revert and bear
exclusively red flowers. It may be worth mentioning that I fertil-
ised the Purple Sweet-pea (Lathyrus odoratus) with pollen from the
light-coloured Painted Lady: seedlings raised from one and the
same pod were not intermediate in character, but perfectly resem-
bled both parents. Later in the summer, the plants which had at
first borne flowers identical with those of the Painted Lady, pro-
duced flowers streaked and blotched with purple; showing in these
darker marks a tendency to reversion to the mother-variety. Andrew
Knight 109 fertilised two white grapes with pollen of the Aleppo
grape, which is darkly variegated both in its leaves and fruit. The
result was that the young seedlings were not at first variegated,
but all became variegated during the succeeding summer; besides
this, many produced on the same plant bunches of grapes which
were all black, or all white, or lead-coloured striped with white, or
white dotted with minute black stripes; and grapes of all these
shades could frequently be found on the same footstalk.
In most of these cases of crossed varieties, and in some
of the cases of crossed species, the colours proper to both
parents appeared in the seedlings, as soon as they first
flowered, in the form of stripes or larger segments, or as
whole flowers or fruit of two kinds borne on the same
plant; and in this case the appearance of the two colours
cannot strictly be said to be due to reversion, but to some
incapacity of fusion, leading to their segregation. When,
107 Géograph. Bot. de l'Europe,'
tom. iii. 1854, p. 405; and 'De la Fécon-
dation,' 1862, p. 302.
108 Traité du Citrus,' 1811, p. 45.
109 Transact. Linn. Soc.,' vol. ix. p.
268.

CHAP. XI.
473
OF REPRODUCTION AND VARIATION.
however, the later flowers or fruit, produced during the
same season or during a succeeding year or generation,
become striped or half-in-half, &c., the segregation of the
two colours is strictly a case of reversion by bud-varia-
tion. In a future chapter I shall show that, with animals
of crossed parentage, the same individual has been known
to change its character during growth, and to revert to
one of its parents which it did not at first resemble.
From the various facts now given there can be no doubt
that the same individual plant, whether a hybrid or a
mongrel, sometimes returns in its leaves, flowers, and
fruit, either wholly or by segments, to both parent-forms
in the same manner as the Cytisus adami, and the .Biz-
zarria Orange.
We will now consider the few facts which have been
recorded in support of the belief that a variety when
grafted or budded on another variety sometimes affects
the whole stock, or at the point of junction gives rise to.
a bud, or graft-hybrid, which partakes of the characters
of both stock and scion.
110
It is notorious that when the variegated Jessamine is budded on
the common kind, the stock sometimes produces buds bearing vari-
egated leaves: Mr. Rivers, as he informs me, has seen instances of
this. The same thing occurs with the Oleander. 11 Mr. Rivers, on
the authority of a trustworthy friend, states that some buds of a
golden-variegated ash, which were inserted into common ashes, all
died except one, but the ash-stocks were affected, 111 and produced,
both above and below the points of insertion of the plates of bark
bearing the dead buds, shoots which bore variegated leaves. Mr. J.
Anderson Henry has communicated to me a nearly similar case:
Mr. Brown, of Perth, observed many years ago, in a Highland glen,
an ash-tree with yellow leaves; and buds taken from this tree were
inserted into common ashes, which in consequence were affected,
and produced the Blotched Breadalbane Ash. This variety has been
110 Gärtner ("Bastarderzeugung,' s.
611) gives many references on this sub-
ject.
111 A nearly similar account was given
by Bradley, in 1724, in his 'Treatise on
Husbandry,' vol. i. p. 199.

474
CHAP, XI.
ANOMALOUS MODES
propagated, and has preserved its character during the last fifty
years. Weeping ashes, also, were budded on the affected stocks,
and became similarly variegated. Many authors consider variega-
tion as the result of disease; and on this view, which however is
doubtful, for some variegated plants are perfectly healthy and vi-
gorous, the foregoing cases may be looked at as the direct result of
the inoculation of a disease. Variegation is much influenced, as we
shall hereafter see, by the nature of the soil in which the plants are
grown; and it does not seem improbable that whatever change in
the sap or tissues certain soils induce, whether or not called a dis-
ease, might spread from the inserted piece of bark to the stock.
But a change of this kind cannot be considered to be of the nature
of a graft-hybrid.
There is a variety of the hazel with dark-purple leaves, like those
of the copper-beech: no one has attributed this colour to disease,
and it apparently is only an exaggeration of a tint which may often
be seen on the leaves of the common hazel. When this variety is
grafted on the common hazel,112 it sometimes colours, as has been
asserted, the leaves below the graft; but I should add that Mr.
Rivers, who has possessed hundreds of such grafted trees, has never
seen an instance.
Gärtner 113 quotes two separate accounts of branches of dark and
white-fruited vines which had been united in various ways, such as
being split longitudinally, and then joined, &c.; and these branches
produced distinct bunches of grapes of the two colours, and other
bunches with grapes either striped or of an intermediate and new
tint. Even the leaves in one case were variegated. These facts are
the more remarkable because Andrew Knight never succeeded in
raising variegated grapes by fertilising white kinds by pollen of
dark kinds; though, as we have seen, he obtained seedlings with
variegated fruit and leaves, by fertilising a white variety by the
variegated dark Aleppo grape. Gärtner attributes the above-quoted
cases merely to bud-variation ; but it is a strange coincidence that
the branches which had been grafted in a peculiar manner should
alone have thus varied; and H. Adorne de Tscharner positively as-
serts that he produced the described result more than once, and
could do so at will, by splitting and uniting the branches in the
manner described by him.
I should not have quoted the following case had not the author
of Des Jacinthes' 114 impressed me with the belief not only of his
extensive knowledge, but of his truthfulness: he says that bulbs
112 Loudon's Arboretum,' vol. iv. P.
2595.
113 Bastarderzeugung,' s. 619.
114 Amsterdam, 1768, p. 124.

Cuap. XI. OF REPRODUCTION AND VARIATION. 475
of blue and red hyacinths may be cut in two, and that they will
grow together and throw up a united stem (and this I have myself
seen), with flowers of the two colours on the opposite sides. But the
remarkable point is, that flowers are sometimes produced with the
two colours blended together, which makes the case closely analo-
gous with that of the blended colours of the grapes on the united
vine-branches.
Mr. R. Trail stated in 1867, before the Botanical Society of Edin-
burgh (and has since given me fuller information), that several years
ago he cut about sixty blue and white potatoes into halves through
the eyes or buds, and then carefully joined them, destroying at the
same time the other eyes. Some of these united tubers produced
white, and others blue tubers, and it is probable that in these cases
the one half alone of the bud grew. Some, however, produced
tubers partly white and partly blue; and the tubers from about four
or five were regularly mottled with the two colours. In these latter
cases we may conclude that a stem had been formed by the union
of the bisected buds; and as tubers are produced by the enlarge-
ment of subterranean branches arising from the main stem, their
mottled colour apparently affords clear evidence of the intimate com-
mingling of the two varieties. I have repeated these experiments
on the potato and on the hyacinth on a large scale, but with no
success.
The most reliable instance known to me of the formation of a
graft-hybrid is one, recorded by Mr. Poynter, 115 who assures me, in
a letter, of the entire accuracy of the statement. Rosa Devoniensis
had been budded some years previously on a white Banksian rose;
and from the much enlarged point of junction, whence the Devoni-
ensis and Banksian still continued to grow, a third branch issued,
which was neither pure Banksian nor pure Devoniensis, but partook
of the character of both; the flowers resembled, but were superior
in character to those of the variety called Lamarque (one of the
Noisettes), while the shoots were similar in their manner of growth
to those of the Banksian rose, with the exception that the longer
and more robust shoots were furnished with prickles. This rose was
exhibited before the Floral Committee of the Horticultural Society
of London. Dr. Lindley examined it, and concluded that it had cer-
tainly been produced by the mingling of R. Banksic with some
rose like R. Devoniensis, " for while it was very greatly increased
in vigour and in the size of all the parts, the leaves were half-way
between a Banksian and Tea-scented rose." It appears that rose-
115 Gard. Chron.,' 1960, p. 672, with a woodcut.

476
CHAP. XI.
DIRECT ACTION OF THE MALE
growers were aware that the Banksian rose sometimes affects other
roses. Had it not been for this latter statement, it might have been
suspected that this new variety was simply due to bud-variation, and
that it had occurred by a mere accident at the point of junction be-
tween the two old kinds.
To sum up the foregoing facts: the statement that
Cytisus adami originated as a graft-hybrid is so precise
that it can hardly be rejected, and, as we have just seen,
some analogous facts render the statement to a certain
extent probable. The peculiar, monstrous condition of
the ovules, and the apparently sound condition of the pol-
len, favour the belief that it is not an ordinary or seminal
hybrid. On the other hand, the fact that the same two
species, viz. C. laburnum and purpureus, have spontane-
ously produced hybrids by seed, is a strong argument in
support of the belief that C. adami originated in a simi-
lar manner. With respect to the extraordinary tendency
which this tree exhibits to complete or partial reversion,
we have seen that undoubted seminal hybrids and mon-
grels are similarly liable. On the whole, I am inclined
to put trust in M. Adam's statement; and if it should
ever be proved true, the same view would probably have
to be extended to the Bizzarria and Trifacial oranges and
to the apples above described ; but more evidence is
requisite before the possibility of the production of graft-
hybrids can be fully admitted. Although it is at present
impossible to arrive at any certain conclusion with respect
to the origin of these remarkable trees, the various facts
above given appear to me to deserve attention under seve-
ral points of view, more especially as showing that the
power of reversion is inherent in Buds.
On the direct or immediate action of the Male Element
on the Mother Form.- Another remarkable class of facts
must be here considered, because they have been sup-
posed to account for some cases of bud-variation: I refer
to the direct action of the male element, not in the ordi-

CHAP XI,
ELEMENT ON THE MOTHER FORM. 477
nary way on the ovules, but on certain parts of the female
plant, or in the case of animals on the subsequent progeny
of the female by a second male.
I may premise that
with plants the ovarium and the coats of the ovules
are obviously parts of the female, and it could not have
been anticipated that they would be affected by the pollen
of a foreign variety or species, although the develop-
ment of the embryo, within the embryonic sack, within
the ovule, within the ovarium, of course depends on the
male element.
Even as long ago as 1729 it was observed 116 that white and blue
varieties of the Pea, when planted near each other, mutually cross-
ed, no doubt through the agency of bees, and in the autumn blue
and white peas were found within the same pods. Wiegmann
made an exactly similar observation in the present century. The
same result has followed several times when a variety with peas of
one colour has been artificially crossed by a differently-coloured
variety. 117 These statements led Gärtner, who was highly sceptical
on the subject, carefully to try a long series of experiments: he se-
lected the most constant varieties, and the result conclusively showed
that the colour of the skin of the pea is modified when pollen of a
differently coloured variety is used. This conclusion has since been
confirmed by experiments made by the Rev. J. M. Berkeley. 118
Mr. Laxton of Stamford, whilst making experiments on peas for
the express purpose of ascertaining the influence of foreign pollen
on the mother-plant, has recently 119 observed an important additional
fact. He fertilised the Tall Sugar pea, which bears very thin green
pods, becoming brownish-white when dry, with pollen of the Purple.
podded pea, which, as its name expresses, has dark-purple pods with
very thick skin, becoming pale reddish-purple when dry. Mr. Lax-
ton has cultivated the tall sugar-pea during twenty years, and has
never seen or heard of it producing a purple pod ; nevertheless, a
flower fertilised by pollen from the purple-pod yielded a pod clouded
with purplish-red, which Mr. Laxton kindly gave to me.
of about two inches in length towards the extremity of the pod, and
a smaller space near the stalk, were thus coloured. On comparing
A space
116 Philosophical Transact.,' vol.
xliii., 1744-45, p. 525.
117 Mr. Swayne, in 'Transact. Hort.
Soc.,' vol. v. p. 234; and Gärtner, 'Bas-
tarderzeugung,' 1849, s. 51 and 499.
118 Gard. Chron.,' 1851, p. 404.
119 Ibid., 1866, p. 900.

478
CHAP. XI.
DIRECT ACTION OF THE MALE
the colour with that of the purple-pod, both pods having been first
dried and then soaked in water, it was found to be identically the
same; and in both the colour was confined to the cells lying imme-
diately beneath the outer skin of the pod. The valves of the crossed
pod were also decidedly thicker and stronger than those of the pods
of the mother-plant, but this may have been an accidental circum-
stance, for I know not how far their thickness in the Tall Sugar-pea
is a variable character.
The peas of the Tall Sugar-pea, when dry, are pale greenish-brown,
thickly covered with dots of dark purple so minute as to be visible
only through a lens, and Mr. Laxton has never seen or heard of this
variety producing a purple pea ; but in the crossed pod one of the
peas was of a uniform beautiful violet-purple tint, and a second was
irregularly clouded with pale purple. The colour lies in the outer
of the two coats which surround the pea. As the peas of the purple-
podded variety when dry are of a pale greenish-buff, it would at
first appear that this remarkable change of colour in the peas in the
crossed pod could not have been caused by the direct action of the
pollen of the purple-pod : but when we bear in mind that this latter
variety has purple flowers, purple marks on its stipules, and purple
pods; and that the Tall sugar-pea likewise has purple flowers and sti-
pules, and microscopically minute purple dots on the peas, we can
hardly doubt that the tendency to the production of purple in both
parents has in combination modified the colour of the peas in the
crossed pod. After having examined these specimens, I crossed the
same two varieties, and the peas in one pod, but not the pods them-
selves, were clouded and tinted with purplish-red in a much more
conspicuous manner than the peas in the uncrossed pods produced at
the same time by the same plants. I may notice as a caution that
Mr. Laxton sent me various other crossed peas slightly, or even
greatly, modified in colour; but the change in these cases was due,
as had been suspected by Mr. Laxton, to the altered colour of the
cotyledons, seen through the transparent coats of the peas; and as
the cotyledons are parts of the embryo, these cases are not in any
way remarkable.
Turning now to the genus Matthiola. The pollen of one kind of
stock sometimes affects the colour of the seeds of another kind,
used as the mother-plant. I give the following case the more read-
ily, as Gärtner doubted similar statements with respect to the stock
previously made by other observers. A well-known horticulturist,
Major Trevor Clarke, informs me 120 that the seeds of the large red-
120 See also a paper by this observer, read before the International Hort. and Bot.
Congress of London, 1866.

CHAP. XI.
479
ELEMENT ON THE MOTHER FORM.
flowered biennial stock (Matthiola annua ; Cocardeau of the French)
are light brown, and those of the purple branching Queen stock (M.
incana) are violet-black; and he found that, when flowers of the red
stock were fertilised by pollen from the purple stock, they yielded
about fifty per cent. of black seeds. He sent me four pods from a red-
flowered plant, two of which had been fertilised by their own pollen,
and they included pale brown seed: and two which had been crossed
by pollen from the purple kind, and they included seeds all deeply
tinged with black. These latter seeds yielded purple-flowered plants
like their father, whilst the pale brown seeds yielded normal red-
flowered plants; and Major Clarke, by sowing similar seeds, has ob-
served on a greater scale the same result. The evidence in this case
of the direct action of the pollen of one species on the colour of the
seeds of another species appears to me conclusive.
In the foregoing cases, with the exception of that of
the purple-podded pea, the coats of the seeds alone have
been affected in colour. We shall now see that the ova-
rium itself, whether forming a large fleshy fruit or a mere
thin envelope, may be modified by foreign pollen, in
colour, flavour, texture, size, and shape.
The most remarkable instance, because carefully recorded by
highly competent authorities, is one of which I have seen an account
in a letter written, in 1867, by M. Naudin to Dr. Hooker. M. Naudin
states that he has seen fruit growing on Chamærops humilis, which
had been fertilised by M. Denis with pollen from the Phoenix or
date-palm. The fruit or drupe thus produced was twice as large as,
and more elongated than, that proper to the Chamærops ; so that it
was intermediate in these respects, as well as in texture, between
the fruit of the two parents. These hybridised seeds germinated,
and produced young plants likewise intermediate in character. This
case is the more remarkable as the Chamærops and Phoenix belong
not only to distinct genera, but in the estimation of some botanists
to distinct sections of the family.
Gallesio 121 fertilised the flowers of an orange with pollen from
the lemon; and one fruit thus produced bore a longitudinal stripe
of peel having the colour, flavour, and other characters of the lemon.
Mr. Anderson 122 fertilised a green-fleshed melon with pollen from a
scarlet-fleshed kind; in two of the fruits "a sensible change was
121 Traité du Citrus,' p. 40.
122 Transact. Hort. Soc.,' vol. iv. p. 318. See also vol. v. p. 65.

480
CHAP. XI.
DIRECT ACTION OF THE MALE
perceptible; and four other fruits were somewhat altered both inter-
naliy and externally." The seeds of the two first-mentioned fruits
produced plants partaking of the good properties of both parents.
In the United States, where Cucurbitacea are largely cultivated, it
is the popular belief 123 that the fruit is thus directly affected by
foreign pollen; and I have received a similar statement with respect
to the cucumber in England. It is known that grapes have been
thus affected in colour, size, and shape: in France a pale-coloured
grape had its juice tinted by the pollen of the dark-coloured Tein-
turier; in Germany a variety bore berries which were affected by
the pollen of two adjoining kinds; some of the berries being only
partially affected or mottled. 124 As long ago as 1751 125 it was ob-
served that, when differently coloured varieties of maize grow near
each other, they mutually affect each other's seeds, and this is now
a popular belief in the United States. Dr. Savi 126 tried the experi-
ment with care: he sowed yellow and black-seeded maize together,
and on the same ear some of the seeds were yellow, some black, and
some mottled, 127 the differently coloured seeds being arranged in
rows or irregularly. Mr. Sabine states 128 that he has seen the form
of the nearly globular seed-capsule of Amaryllis vittata altered by
the application of the pollen of another species, of which the capsule
has gibbous angles. Mr. J. Anderson Henry
Mr. J. Anderson Henry 129 crossed Rhododen-
dron Dalhousic with the pollen of R. Nuttallii, which is one of the
largest-flowered and noblest species of the genus. The largest pod
produced by the former species, when fertilised with its own pollen,
123 Prof. Asa Gray, ' Proc. Acad. Sc.,' the two parents do not readily unite, as
Boston, vol. iv., 1860, p. 21.
in the cases of Mirabilis and Dianthus
124 For the French case, see 'Proc. given a few pages back. Thirdly, in
Hort. Soc.,' vol. i. new series, 1866, p. 50. crossed plants of a subsequent genera-
For Germany, see M. Jack, quoted in tion, by reversion, through either bud or
Henfrey's Botanical Gazette,' vol. i. p. seminal generation. Fourthly, by rever-
277. A case in England has recently sion to a character not originally gained
been alluded to by the Rev. J. M. Berke- by a cross, but which had long been lost,
ley before the Hort. Soc. of London. as with white-flowered varieties, which
125 Philosophical Transactions,' vol. we shall hereafter see often become
xlvii., 1751-52, p. 206.
striped with some other colour. Lastly,
126 Gallesio, Teoria della Riprodu- there are cases, as when peaches are pro-
zione,' 1816, p. 95.
duced with a half or quarter of the fruit
127 It may be worth while to call at- like a nectarine, in which the change is
tention to the several means by which apparently due to mere variation through
flowers and fruit become striped or mot- either bud or seminal generation.
tled. Firstly, by the direct action of the 128 Transact. Hort. Soc.,' vol. v. p.
pollen of another variety or species, as 69.
with the above-given cases of oranges 129 Journal of Horticulture,' Jan. 20,
and maize. Secondly, in crosses of the 1863, p. 46.
first generation, when the colours of

CHAP. XI. ELEMENT ON THE MOTHER FORM.
481
measured 13 in length and 11 in girth ; whilst three of the pods
which had been fertilised by pollen of R. Nuttallii measured 15 inch
in length and no less than 2 inches in girth. Here we see the
effect of foreign pollen apparently confined to increasing the size of
the ovarium ; but we must be cautious in assuming, as the follow-
ing case shows, that in this instance size has been directly trans-
ferred from the male parent to the capsule of the female plant. Mr.
Henry fertilised Arabis blepharophylla with pollen of A. Soyeri, and
the pods thus produced, of which he was so kind as to send me de-
tailed measurements and sketches, were much larger in all their di-
mensions than those naturally produced by either the male or female
parent-species. In a future chapter we shall see that the organs of
vegetation in hybrid plants, independently of the character of either
parent, are sometimes developed to a monstrous size; and the in-
creased size of the pods in the foregoing cases may be an analogous
fact.
No case of the direct action of the pollen of one variety on another
is better authenticated or more remarkable than that of the common
apple. The fruit here consists of the lower part of the calyx and of
the upper part of the flower-peduncle 130 in a metamorphosed condi-
tion, so that the effect of the foreign pollen has extended even be-
yond the limits of the ovarium. Cases of apples thus affected were
recorded by Bradley in the early part of the last century; and other
cases are given in old volumes of the Philosophical Transactions ;132
in one of these a Russeting apple and an adjoining kind mutually
affected each other's fruit; and in another case a smooth apple af-
fected a rough-coated kind. Another instance has been given 132 of
two very different apple-trees growing close to each other, which
bore fruit resembling each other, but only on the adjoining branches.
It is, however, almost superfluous to adduce these or other cases,
after that of the St. Valery apple, which, from the abortion of the
stamens, does not produce pollen, but, being annually fertilised by
the girls of the neighbourhood with pollen of many kinds, bears
fruit, “ differing from each other in size, flavour, and colour, but re-
sembling in character the hermaphrodite kinds by which they have
been fertilised.'' 133
130 See on this head the high autho-
rity of Prof. Decaisne, in a paper trans-
lated in Proc. Hort. Soc.,' vol. i. new
series, 1866, p. 48.
131 Vol. xliii., 1744-45, p. 525; vol.
xlv., 1747-49, p. 602.
132 Transact. Hort. Soc.,' vol. v. pp.
69 and 63. Puvis also has collected ('De
21
la Dégénération,' 1937, p. 36) several
other instances; but it is not in all cases
possible to distinguish between the di-
rect action of foreign pollen and bud-
variations.
133 T. de Clermont-Tonnerre, in 'Mém.
de la Soc. Linn. de Paris,' tom. iii., 1825,
p. 164

482
CHAP. XI.
DIRECT ACTION OF THE MALE
I have now shown, on the authority of several excel-
lent observers, in the case of plants belonging to widely
different orders, that the pollen of one species or variety,
when applied to a distinct form, occasionally causes the
coats of the seeds and the ovarium or fruit, including
even in one instance the calyx and upper part of the pe-
duncle of the mother-plant, to become modified. Some-
times the whole of the ovarium or all the seeds are thus
affected; sometimes only a certain number of the seeds,
as in the case of the pea, or only a part of the ovarium,
as with the striped orange, mottled grapes and maize,
are thus affected. It must not be supposed that any di-
rect or immediate effect invariably follows the use of
foreign pollen: this is far from being the case; nor is it
known on what conditions the result depends. Mr.
Knight 134 expressly states that he has never seen the
fruit thus affected, though he has crossed thousands of
apple and other fruit-trees. There is not the least reason to
believe that a branch which has borne seed or fruit direct-
ly modified by foreign pollen is itself affected, so as sub-
sequently to produce modified buds: such an occurrence,
from the temporary connection of the flower with the
stem, would be hardly possible. Hence but very few, if
any, of the cases of sudden modifications in the fruit of
trees, given in the early part of this chapter, can be ac-
counted for by the action of foreign pollen ; for such modi-
fied fruits have commonly been afterwards propagated by
budding or grafting. It is also obvious that changes of
colour in the flower which necessarily supervene long be-
fore it is ready for fertilisation, and changes in the shape
or colour of the leaves, can have no relation to the action
of foreign pollen: all such cases must be attributed to
simple bud-variation.
The proofs of the action of foreign pollen on the mother-
plant have been given in considerable detail, because this
154 Transact of Hort. Soc.,' vol, v, p. 68.

CHAP. XI.
483
ELEMENT ON THE MOTIIER FORM.
.
action, as we shall see in a future chapter, is of the highest
theoretical importance, and because it is in itself a re-
markable and apparently anomalous circumstance. That
it is remarkable under a physiological point of view is
clear, for the male element not only affects, in accordance
with its proper function, the germ, but the surrounding
tissues of the mother-plant. That the action is anomalous
in appearance is true, but hardly so in reality, for appa-
rently it plays the same part in the ordinary fertilisation
of many flowers. Gärtner has shown, as by gradually in-
creasing the number of pollen-grains until he succeeded
in fertilising a Malva, that many grains are expended in
the development, or, as he expresses it, in the satiation,
of the pistil and ovarium. Again, when one plant is fer-
tilised by a widely distinct species, it often happens that
the ovarium is fully and quickly developed without any
seeds being formed, or the coats of the seeds are devel-
oped without an embryo being formed within. Dr. Hil-
debrand also has lately shown in a valuable paper that,
with several Orchideæ, the action of the plant's own pol-
len is necessary for the development of the ovarium, and
that this development takes place not only long before
the pollen-tubes have reached the ovules, but even before
the placenta and ovules have been formed; so that with
these orchids the pollen apparently acts directly on the
ovarium. On the other hand, we must not overrate the
efficacy of pollen in this respect ; for in the case of hy-
bridised plants it might be argued that an embryo had
been formed and had affected the surrounding tissues of
the mother-plant before it perished at a very early age.
Again, it is well known that with many plants the ovarium
may be fully developed, though pollen be wholly excluded.
And lastly, Mr. Smith, the late Curator at Kew (as I hear
136
135 Beitrage zur Kenntniss der Be-
fruchtung,' 1814, s. 347-351.
136 Die Fruchtbildung der Orchideen,
ein Beweis für doppelte Wirkung des
Pollen,' Botanische Zeitung, No. 44 et
seq., Oct. 30, 1963; and 1565, s. 249.

484
CHAP, XI.
DIRECT ACTION OF THE MALE
through Dr. Hooker), observed the singular fact with
an orchid, the Bonatea speciosa, the development of the
ovarium could be effected by mechanical irritation of the
stigma. Nevertheless, from the number of the pollen-
grains expended “in the satiation of the ovarium and
pistil,”—from the generality of the formation of the
ovarium and seed-coats in sterile hybridised plants,—and
from Dr. Hildebrand's observations on orchids, we may
admit that in most cases the swelling of the ovarium,
and the formation of the seed-coats, are at least aided, if
not wholly caused, by the direct action of the pollen, in-
dependently of the intervention of the fertilised germ.
Therefore, in the previously-given cases we have only to
add to our belief in the power of the plant's own pollen
on the development of the ovarium and seed-coats, its
further power, when applied to a distinct species or
variety, of influencing the shape, size, colour, texture,
&c., of these same parts.
Turning now to the animal kingdom. If we could
imagine the same flower to yield seeds during successive
years, then it would not be very surprising that a flower
of which the ovarium had been modified by foreign pol-
len should next year produce, when self-fertilised, off-
spring modified by the previous male influence. Closely
analogous cases have actually occurred with animals.
In the case often quoted from Lord Morton,187 a nearly
purely-bred, Arabian, chesnut mare bore a hybrid to a
quagga; she was subsequently sent to Sir Gore Ouseley,
and produced two colts by a black Arabian horse. These
colts were partially dun-coloured, and were striped on the
legs more plainly than the real hybrid, or even than the
quagga.
One of the two colts had its neck and some
other parts of its body plainly marked with stripes.
Stripes on the body, not to mention those on the legs,
137 Philos. Transact.,' 1821, p. 20.

CHAP. XI. ELEMENT ON THE MOTHER FORM.
485
and the dun colour, are extremely rare, -I speak after
having long attended to the subject,-with horses of
all kinds in Europe, and are unknown in the case of
Arabians. But what makes the case still more striking
is that the hair of the mane in these colts resembled that
of the quagga, being short, stiff, and upright. Hence
there can be no doubt that the quagga affected the cha-
racter of the offspring subsequently begot by the black
Arabian horse. With respect to the varieties of our do-
mesticated animals, many similar and well-authenticated
facts have been published,138 and others have been com-
municated to me, plainly showing the influence of the
first male on the progeny subsequently borne by the
mother to other males. It will suffice to give a single in-
stance, recorded in the Philosophical Transactions,' in a
paper following that by Lord Morton: Mr. Giles put a
sow of Lord Western's black and white Essex breed to a
wild boar of a deep chesnut colour; and the “pigs pro-
duced partook in appearance of both boar and sow, but in
some the chesnut colour of the boar strongly prevailed.”
After the boar had long been dead, the sow was put to
a boar of her own black and white breed,-a kind which
is well known to breed very true and never to show any
chesnut colour, -yet from this union the sow produced
some young pigs which were plainly marked with the
same chesnut tint as in the first litter. Similar cases
have so frequently occurred, that careful breeders avoid
putting a choice female to an inferior male on account of
138 Dr. Alex. Harvey on 'A remark-
able Effect of Cross-breeding,' 1851. On
the Physiology of Breeding,' by Mr.
Reginald Orton, 1855. "Intermarriage,'
by Alex. Walker, 1837. L'Hérédité
Naturelle,' by Dr. Prosper Lucas, tom.
ii. p. 58. Mr. W. Sedgwick in 'British
and Foreign Medico-Chirurgical Review,
1863, July, p. 183. Bronn, in his 'Ge-
schichte der Natur,' 1843, B. ii. s. 127,
has collected several cases with respect
to mares, sows, and dogs. Mr. W. C.
L. Martin ("History of the Dog,' 1845,
p. 104) says he can personally vouch for
the influence of the male parent of the
first litter on the subsequent litters by
other fathers. A French poet, Jacques
Savary, who wrote in 1665 on dogs, was
aware of this singular fact.

486
CHAP, XI.
CONCLUSION AND SUMMARY
the injury to her subsequent progeny which may be ex-
pected to follow.
Some physiologists have attempted to account for these
remarkable results from a first impregnation by the close
attachment and freely intercommunicating blood vessels
between the modified embryo and the mother. But it is
a most improbable hypothesis that the mere blood of one
individual should affect the reproductive organs of an-
other individual in such a manner as to modify the sub-
sequent offspring. The analogy from the direct action
of foreign pollen on the ovarium and seed-coats of the
mother-plant strongly supports the belief that the male
element acts directly on the reproductive organs of the
female, wonderful as is this action, and not through the
intervention of the crossed embryo. With birds there is
no such close connection between the embryo and mother
as in the case of mammals : yet a careful observer, Dr.
Chapuis, states 139 that with pigeons the influence of a
first male sometimes makes itself perceived in the suc-
ceeding broods; but this statement, before it can be fully
trusted, requires confirmation.
Conclusion and Summary of the Chapter. The facts
given in the latter half of this chapter are well worthy of
consideration, as they show us in how many extraordina-
ry modes one organic form may lead to the modification
of another, and often without the intervention of seminal
reproduction. There is ample evidence, as we have just
seen, that the male element may either directly affect the
structure of the female, or in the case of animals lead to
the modification of her offspring. There is a considerable
but insufficient body of evidence showing that the tissues
of two plants may unite and form a bud having a blend-
ed character; or again, that buds inserted into a stock
may affect all the buds subsequently produced by this
139 "Le Pigeon Voyageur Belge,' 1865, p. 59.

CHAP. XI.
487
OF THE CHAPTER.
stock. Two embryos, differing from each other and con-
tained in the same seed, may cohere and form a single
plant. Offspring from a cross between two species or
varieties may in the first or in a succeeding generation
revert in various degrees by bud-variation to their parent-
forms; and this reversion or segregation of character may
affect the whole flower, fruit, or leaf-bud, or only the half
or smaller segment, or a single organ. In some cases this
segregation of character apparently depends on some in-
capacity of union rather than on reversion, for the flow-
ers or fruit which are first produced display by segments
the characters of both parents. In the Cytisus adami
and the Bizzarria orange, whatever their origin may have
been, the two parent species occur blended together un-
der the form of a sterile hybrid, or reappear with their
characters perfect and their reproductive organs effec-
tive; and these trees, retaining the same sportive charac-
ter, can be propagated by buds. These various facts
ought to be well considered by any one who wishes to
embrace under a single point of view the various modes
of reproduction by gemmation, division, and sexual union,
the reparation of lost parts, variation, inheritance, rever-
sion, and other such phenomena. In a chapter towards
the close of the following volume I shall attempt to con-
nect these facts together by a provisional hypothesis.
In the early half of this chapter I have given a long
list of plants in which through bud-variation, that is, in-
dependently of reproduction by seed, the fruit has sud-
denly become modified in size, colour, flavour, hairiness,
shape, and time of maturity ; flowers have similarly
changed in shape, colour, and doubleness, and greatly in
the character of the calyx; young branches or shoots
have changed in colour, in bearing spines, and in habit
of growth, as in climbing and weeping; leaves have
changed in colour, variegation, shape, period of unfolding,
and in their arrangement on the axis. Buds of all kinds,
whether produced on ordinary branches or on subterranean

488
CTAP. XI,
CONCLUSION AND SUMMARY
stems, whether simple or, as in tubers and bulbs, much
modified and supplied with a stock of nutriment, are all
liable to sudden variations of the same general nature.
In the list, many of the cases are certainly due to re-
version to characters not acquired from a cross, but which
were formerly present, and have been lost for a longer or
shorter period of time ;-as when a bud on a variegated
plant produces plain leaves, or when variously-coloured
flowers on the Chysanthemum revert to the aboriginal
yellow tint. Many other cases included in the list are
probably due to the plants being of crossed parentage,
and to the buds reverting to one of the two parent-forms.
In illustration of the origin of Cytisus adami, several cases
were given of partial or complete reversion, both with
hybrid and mongrel plants; hence we may suspect that
the strong tendency in the Chrysanthemum, for instance,
to produce by bud-variation differently-coloured flowers,
results from the varieties formerly having been intention-
ally or accidentally crossed ; and that their descendants
at the present day still occasionally revert by buds to the
colours of the more persistent parent-varieties. This is
almost certainly the case with Rollison's Unique Pelargo-
nium; and so it may be to a large extent with the bud-
varieties of the Dahlia and with the 6 broken colours"
of Tulips.
Many cases of bud-variation, however, cannot be attri-
buted to reversion, but to spontaneous variability, such
as so commonly occurs with cultivated plants when raised
from seed. As a single variety of the Chrysanthemum
has produced by buds six other varieties, and as one va-
riety of the gooseberry has borne at the same time four
distinct varieties of fruit, it is scarcely possible to believe
that all these variations are reversions to former parents.
We can hardly believe, as remarked in a previous chapter,
that all the many peaches which have yielded nectarine.
buds are of crossed parentage. Lastly, in such cases as
that of the moss-rose with its peculiar calyx, and of the

CAAP. XI.
489
OF THE CHAPTER. .
rose which bears opposite leaves, in that of the Imato-
phyllum, &c., there is no known natural species or seed-
ling variety, from which the characters in question could
have been derived by crossing. We must attribute all
such cases to actual variability in the buds. The varie-
ties which have thus arisen cannot be distinguished by
any external character from seedlings; this is notoriously
the case with the varieties of the Rose, Azalea, and many
other plants. It deserves notice that all the plants which
have yielded bud-variations have likewise varied greatly
by seed.
These plants belong to so many orders that we may
infer that almost every plant would be liable to bud-
variation if placed under the proper exciting conditions.
These conditions, as far as we can judge, mainly depend
on long-continued and high cultivation ; for almost all
the plants in the foregoing lists are perennials, and have
been largely propagated in many soils and under different
climates, by cuttings, offsets, bulbs, tubers, and especially
by budding or grafting. The instances of annuals vary-
ing by buds, or producing on the same plant differently
coloured flowers are comparatively rare: Hopkirk140 has
seen this with Convolvulus tricolor; and it is not rare
with the balsam and annual Delphinium. According to
Sir R. Schomburgk, plants from the warmer temperate
regions, when cultivated under the hot climate of St. Do-
mingo, are eminently liable to bud-variation ; but change
of climate is by no means a necessary contingent, as we
see with the gooseberry, currant, and some others. Plants
living under their natural conditions are very rarely
subject to bud-variation : variegated and coloured leaves
have, however, been occasionally observed ; and I have
given an instance of the variation of buds on an ash-tree;
but it is doubtful whether any tree planted in ornamental
grounds can be considered as living under strictly natural
140 Flora Anomala.' p. 164.

-490
CHAP. XI,
CONCLUSION AND SUMMARY
conditions. Gärtner has seen white and dark-red flowers
produced from the same root of the wild Achillea mille-
folium, and Prof. Caspary has seen Viola lutea, in a com-
pletely wild condition, bearing flowers of different colours
and sizes.141
As wild plants are so rarely liable to bud-variation,
whilst highly cultivated plants long propagated by arti-
ficial means have yielded by this form of reproduction
many varieties, we are led through a series such as the
following, -namely, all the eyes in the same tuber of the
potato varying in the same manner,-all the fruit on a pur-
ple plum-tree suddenly becoming yellow,-all the fruit on
a doubled-flowered almond suddenly becoming peachlike,
-all the buds on grafted trees being in some very slight
degree affected by the stock on which they have been
worked,—all the flowers on a transplanted heartsease
changing for a time in color, size, and shape,—we are led
through such facts to look at every case of bud-variation
as the direct result of the particular conditions of life to
which the plant has been exposed. But if we turn to the
other end of the series, namely, to such cases as that of a
peach-tree which, after having been cultivated by tens of
thousands during many years in many countries, and after
having annually produced thousands of buds, all of which
have apparently been exposed to precisely the same con-
ditions, yet at last suddenly produces a single bud with
its whole character greatly transformed, we are driven to
an opposite conclusion. In such cases as the latter it
would
appear
that the transformation stands in no direct
relation to the conditions of life.
We have seen that varieties produced from seeds and
from buds resemble each other so closely in general ap-
pearance, that they cannot possibly be distinguished.
Just as certain species and groups of species, when propa-
gated by seed, are more variable than other species or
141 Schriften der Phys-Ökon. Gesell. zu Königsberg,' Band vi., Feb. 3, 1865, s. 4.

CHAP XI,
491
OF THE CHAPTER.
genera, so it is in the case of certain bud-varieties. Thus
the Queen of England Chrysanthemum has produced by
this latter process no less than six, and Rollison's Unique
Pelargonium four distinct varieties; moss-roses have also
produced several other moss-roses. The Rosaceæ have
varied by buds more than any other group of plants;
but this may be in large part due to so many members
having been long cultivated; but within this one group,
the peach has often varied by buds, whilst the apple and
pear, both grafted trees extensively cultivated, have af-
forded, as far as I can ascertain, extremely few instances
of bud-variation.
The law of analogous variation holds good with vari-
eties produced by buds, as with those produced from
seed: more than one kind of rose has sported into a
moss-rose; more than one kind of camellia has assumed
an hexagonal form; and at least seven or eight varieties
of the peach have produced nectarines.
The laws of inheritance seem to be nearly the same
with seminal and bud-varieties. We know how com-
monly reversion comes into play with both, and it may
affect the whole, or only segments, of a leaf, flower, or
fruit. When the tendency to reversion affects many
buds
on the same tree, it becomes covered with different kinds
of leaves, flowers, or fruit; but there is reason to believe
that such fluctuating varieties have generally arisen from
seed. It is well known that, out of a number of seed-
ling varieties, some transmit their character much more
truly by seed than others; so with bud-varieties some re-
tain their character by successive buds more truly than
others; of which instances have been given with two
kinds of variegated Euonymus and with certain kinds of
tulips. Notwithstanding the sudden production of bud-
varieties, the characters thus acquired are sometimes ca-
pable of transmission by seminal reproduction : Mr. Ri-
vers has found that moss-roses generally reproduce them-
selves by seed; and the mossy character has been trans-

492
CHAP. XI,
CONCLUSION AND SUMMARY
ferred by crossing, from one species of rose to another.
The Boston nectarine, which appeared as a bud-variation,
produced by seed a closely allied nectarine. We have
however seen, on the authority of Mr. Salter, that seed
taken from a branch with leaves variegated through bud-
variation, transmits this character very feebly; whilst
many plants, which became variegated as seedlings, trans-
mit variegation to a large proportion of their progeny.
Although I have been able to collect a good many
cases of bud-variation, as shown in the previous lists,
and might probably, by searching foreign horticultural
works, have collected more cases, yet their total number
is as nothing in comparison with that of seminal varie-
ties. With seedlings raised from the more variable culti-
vated plants, the variations are almost infinitely nume-
rous, but their differences are generally slight: only at
long intervals of time a strongly marked modification
appears. On the other hand, it is a singular and inexpli-
cable fact that, when plants vary by buds, the variations,
though they occur with comparative rarity, are often, or
even generally, strongly pronounced. It struck me that
this might perhaps be a delusion, and that slight changes
often occurred in buds, but from being of no value were
overlooked or not recorded. Accordingly I applied to
two great authorities on this subject, namely, to Mr.
Rivers with respect to fruit-trees, and to Mr. Salter with
respect to flowers. Mr. Rivers is doubtful, but does not
remember having noticed very slight variations in fruit-
buds. Mr. Salter informs me that with flowers such do
occur, but, if propagated, they generally lose their new
character in the following year; yet he concurs with me
that bud-variations usually at once assume a decided and
permanent character. We can hardly doubt that this is
the rule, when we reflect on such cases as that of the
peach, which has been so carefully observed and of which
such trifling seminal varieties have been propagated, yet
this tree has repeatedly produced by bud-variation nec-

CHAP. XI.
493
OF THE CHAPTER.
tarines, and only twice (as far as I can learn) any other
variety, namely, the Early and Late Grosse Mignonne
peaches; and these differ from the parent-tree in hardly
any character except the period of maturity.
To my surprise I hear from Mr. Salter that he brings
the great principle of selection to bear on variegated
plants propagated by buds, and has thus greatly improved
and fixed several varieties. He informs me that at first
a branch often produces variegated leaves on one side
alone, and that the leaves are marked only with an ir-
regular edging or with a few lines of white and yellow.
To improve and fix such varieties, he finds it necessary to
encourage the buds at the bases of the most distinctly
marked leaves, and to propagate from them alone. By
following with perseverance this plan during three or
four successive seasons, a distinct and fixed variety can
generally be secured.
Finally, the facts given in this chapter prove in how
close and remarkable a manner the germ of a fertilised
seed and the small cellular mass forming a bud resemble
each other in function, in their powers of inheritance
with occasional reversion,—and in their capacity for vari-
ation of the same general nature, in obedience to the
same laws. This resemblance, or rather identity, is ren-
dered far more striking if the facts can be trusted which
apparently render it probable that the cellular tissue of
one species or variety, when budded or grafted on another,
may give rise to a bud having an intermediate character.
In this chapter we clearly see that variability is not ne-
cessarily contingent on sexual generation, though much
more frequently its concomitant than on bud-reproduc-
tion. We see that bud-variability is not solely depend-
ent on reversion or atavism to long-lost characters, or to
those formerly acquired from a cross, but that it is often
spontaneous. But when we ask ourselves what is the
cause of any particular bud-variation, we are lost in doubt,
being driven in some cases to look to the direct action of

494
CHAP, XI,
CONCLUSION AND SUMMARY.
the external conditions of life as sufficient, and in other
cases to feel a profound conviction that these have played
a quite subordinate part, of not more importance than
the nature of the spark which ignites a mass of combusti-
ble matter.
END OF VOL. I.

GARDENING FOR PROFIT,
In the Market and Family Garden.
BY PETER HENDERSON.
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CONTENTS.
Men fitted for the Business of Gardening.
The Amount of Capital Required, and
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Profits of Market Gardening.
Location, Situation, and Laying Out.
Soils, Drainage, and Preparation.
Manures, Implements.
Uses and Management of Cold Frames.
Formation and Management of Hot-beds.
Forcing Pits or Green-houses.
Seeds and Seed Raising.
How, When, and Where to Sow Seeds.
Transplanting, Insects.
Packing of Vegetables for Shipping.
Preservation of Vegetables in Winter.
Vegetables, their Varieties and Cultivation.
In the last chapter, the most valuable kinds are described, and
the culture proper to each is given in detail.
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Τ Η Ε
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CHAP. I. BARBERRY.
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CHAP. V. DWARF CHERRY.
CHAP. VI. CURRANT.
CHAP. VII. GOOSEBERRY.
CHAP. VIII. CORNELIAN CHERRY.
CHAP. IX. CRANBERRY.
CHAP X. HUCKLEBERRY.
CHAP. XI. SHEPERDIA.
CHAP. XII. PREPARATION FOR
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AMERICAN POMOLOGY.
APPLES.
By Doct. JOHN A. WARDER,
PRESIDENT OHIO POMOLOGICAL SOCIETY ; VICE-PRESIDENT AMERICAN POMOLOGICAL
SOCIETY.
293 ILLUSTRATIONS.
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CONTENTS
Chapter 1.--INTRODUCTORY.
Chapter II.-HISTORY OF THE APPLE.
Chapter III.-PROPAGATION.
Buds and Cuttings-Grafting-Budding-The Nursery.
Chapter IV.-DWARFING.
Chapter V.-DISEASES.
Chapter VI. THE SITE FOR AN ORCHARD.
Chapter VII.-PREPARATION OF SOIL FOR AN ORCHARD.
Chapter VIII.-SELECTION AND PLANTING.
Chapter IX.-CULTURE, Etc.
Chapter X.-PHILOSOPHY OF PRUNING.
Chapter XI.-THINNING.
Chapter XII.-RIPENING AND PRESERVING FRUITS.
Chapter XIII and XIV.-INSECTS.
Chapter XV.-CHARACTERS OF FRUITS AND THEIR
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Chapter XVI.-CLASSIFICATION.
Necessity for-Basis of-Characters-Shape-Its Regn-
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GARDENER, the FRUIT CULTURIST, and the AMATEUR.
It is adapted not only to those owning large and Elegant Estates, but con-
tains directions for the best arrangement of the smallest Plots, down to the
City Yard, the Roof or Window Garden, or the simple Flower Stand. It
also gives an abstract of the Principles, Construction, and Management of
AQUARIUMS. Among numerous other matters it treats of
Draining, Giving best methods, estimates of cost, trenches, tiles, etc.,
thus enabling almost any one properly to perform this important work.
Cattle are carefully noticed with reference to the special merits of dif-
ferent breeds for dairying or fattening.
Sheep Management, including Breeding, Feeding, Prices, Profits,
etc., receives attention, and a very full treatise on the Merinos is given.
Grape Culture occupies a large space, embracing the opinions of
men in all parts of the country, as to best sorts, planting, training, diseases,
and general management for home use or marketing.
Full Lists of Ornamental Trees and Shrubs, Fruits, Flowers, Green
and Hot-house Plants, etc., are given, with directions for management each
month in the year.
The Kitchen Garden receives particular attention, with iefer-
ence to the best way to grow and preserve each kind of Vegetable.
In short, as its name indicates, the book treats of almost every subject that
needs consideration by those living in the country, or having any thing to do
with the cultivation of the soil.
Sent Post-Paid.
Price, $5.00.
NEW-YORK:
Orange Judd & Co., 245 Broadway.

MY VINEYARD AT LAKEVIEW ;
OR,
SUCCESSFUL GRAPE CULTURE,
BY A WESTERN GRAPE GROWER.
ILLUSTRATED.
To any one who wishes to grow grapes, whether a single vine or & vine
Fard, th
book is full of valuable teachings. The author gives not only his
success, but, what is of quite as much importance, his failure. It tells just
what the beginner in grape culture wishes to know, with the charm that
always attends the relation of personal experience.
It is especially valuable as giving an account of the processes actually
followed in
CELEBRATED GRAPE REGIONS
in Western New-York and on the shores and islands of Lake Erie.
This book is noticed by a writer in the Horticulturist for August last as
follows: "Two works very different in character and value have just been
published, and seem to demand a passing notice. The better and less pre-
tentious of the two is 'MY VINEYARD AT LAKEVIEW,' a charming little book
that professes to give the actual experience of a western grape grower, de-
tailing not only his successes, but his blunders and failures. It is written
in a pleasant style, without any attempt at display, and contains much ad.
vice that will prove useful to a beginner--the more useful because derived
from the experience of a man who had no leisure for fanciful experiments,
but has been obliged to make his vineyard support himself and his family."
Written in a simple and attractive style, and relating the experience of one who felt
his way along into the successful cultivation of a vineyard in Ohio.-Mass. Ploughman.
It is the experience of a practical grape grower, and not the theory of an experi-
menter. -Bath Daily Sentinel and Times.
It has no superior as an attractive narrative of country life.- Hartford Daily Post.
Many books have been written on the grape, but this is the only work that gives an
account of grape growing as actually practiced at the successful vineyards in the grape
region of the West, and will be welcomed by a large class of readers.-New-Bedford
Standard.
This little volume contains, in an attractive form, and in clear and concise language,
just the information needed to enable any one to become thoroughly posted up in this
delightful and profitable branch of horticulture.- Vermont Farmer.
Just the manual for a beginner, by one who says "he is well rewarded in the success
Butained." Adding, "It might have been reached in half the time, had I possessed the
knowledge imparted to the reader of this book."-Boston Cultivator.
Sent Post-paid.
Price, $1.25.
ORANGE JUDD & CO., 245 Broadway, New-York.

THE AMERICAN
Horticultural Annual
FOR 1868.
A Year-Book
FOR EVERY HOME,
The second number of this serial is now ready. It contains a
popular record of horticultural progress during the past year,
besides valuable articles from
EMINENT HORTICULTURISTS.
Among those who contributed to its pages are
HON. MARSHALL P. WILDER, DOCTOR JOHN A. WARDER,
PETER HENDERSON,
S. B. PARSONS,
THOMAS MEEHAN,
JAS. J. H. GREGORY,
JOSIAH HOOPES,
GEORGE SUCH,
WM. S. CARPENTER,
ANDREW S. FULLER,
CEORGE W. CAMPBELL,
JOHN SAUL,
DOCTOR VAN KEUREN,
JAMES VICK,
and other well-known pomological and floricultural writers.
The engravings, which have been prepared expressly for the
work, are numerous, and make it the
MOST BEAUTIFULLY ILLUSTRATED
work of its kind ever published in this or any other country. It
contains Tables, Lists of Nurserymen, Seedsmen, and Florists, and
other useful matters of reference. Sent post-paid. Price, fancy
paper covers, 50 cts; cloth, 75 cts.
ORANGE JUDD & CO.,
245 Broadway, New-York

THE AMERICAN
Agricultural Annual
FOR 1868.
A Year-Book
Wanted by Everybody.
This volume is now ready, and contains much of interest to
every agriculturist. Besides the general record of agricultural
progress, it has a valuable article on
Factory Dairy Practice,
By GARDNER B. WEEKS, Esq., Secretary of the American Dairy-
men's Association, in which he discusses the reasons for the best
practice and the most approved apparatus, buildings, etc., fully il-
lustrated, and is equally interesting to the practical dairyman and
to the novice.
Sewers and Earth-Closets
In their relations to Agriculture, by Col. GEO. E. WARING, Jr.
Winter Wheat,
Describing, with engravings, new and valuable varieties by JOSEPH
HARRIS and JOHN JOHNSTON ; an article upon
Scythes and Cradles,
By JOHN W. DOUGLAS, (fully illustrated ;) also articles on Horse-
Breaking and on Bitting Colts, by SAM’L F. HEADLY, Esq., (il-
lustrated ;) on Recent Progress in Agricultural Science, by Prof
S. W. JOHNSON ; on Commercial Fertilizers, Veterinary Medicine
and Jurisprudence, Progress of Invention Affecting Agriculture,
Valuable Tables for Farmers and others, etc.
It is intended that the work shall be practical, excellent in the
beauty of its illustrations, and in its adaptation to the wants of
American Farmers, superior to anything of the kind heretofore
published
In its general features it is like the Agricultural Annual for 1867,
containing an Almanac and Calendar, and there will be added a
list of dealers in Agricultural Implements, Seeds, etc. Sent post-
paid. Price, fancy paper covers, 50 cts.; cloth, 75 cts.
ORANGE JUDD & CO.,
245 Broadway, New-York.

THE
GRAPE CULTURIST,
BY
ANDREW S. FULLER.
NEW AND ENLARGED EDITION.
THE STANDARD WORK
ON THE CULTIVATION OF THE HARDY GRAPE,
AS IT NOT ONLY DISCUSSES PRINCIPLES,
BUT
ILLUSTRATES PRACTICE.
Every thing is made perfectly plain, and its teach-
ings may be followed upon
ONE VINE ORA VINEYARD
The following are some of the topics that are treated :
GROWING NEW VARIETIES FROM SEED.
PROPAGATION BY SINGLE BUDS OR EYES.
PROPAGATING HOUSES AND THEIR MANAGEMENT FULLY DESCRIBED.
HOW TO Grow.
CUTTINGS IN OPEN AIR, AND HOW TO MAKE LAYERS.
GRAFTING THE GRAPE-A SIMPLE AND SUCCESSFUL METHOD.
HYBRIDIZING AND CROSSING_MODE OF OPERATION.
SOIL AND SITUATION-PLANTING AND CULTIVATION.
PRUNING, TRAINING, AND TRELLISES - ALL THE SYSTEMS EXPLAINED,
GARDEN CULTURE—How To Grow VINES IN A Door-YARD.
INSECTS, Mildew, SUN-SCALD, AND OTHER TROUBLES.
DESCRIPTION OF THE VALUABLE AND THE DISCARDED VARIETIES.
Sent post-paid. Price $1.50.
Orange Judd & Co., 245 Broadway.

[Established in 1842.]
A Good, Cheap, and very Valuable Paper for
Every Man, Woman and Child,
IN CITY, VILLAGE and COUNTRY,
THE
AMERICAN AGRICULTURIST,
FOR THE
FARM, GARDEN AND HOUSEHOLD,
Including a Special Department of Interesting and
Instructive Reading for CHILDREN and YOUTH.
The Agriculturist is a large periodical of Thirty-two pages, quarto, not octavo,
beautifully printed, and filled with plain, practical, reliable, original matter, includ-
ing hundreds of beautiful and instructive Engravings in every annual volume.
It contains each month a Calendar of Operations to be performed on the Farm,
in the Orchard and Garden, in and around the Dwelling, etc.
The thousands of hints and suggestions given in every volume are prepared by prac-
tical, intelligent working men, who know what they talk and write about. The
articles are thoroughly edited, and every way reliable,
The Household Department is valuable to every Housekeeper, affording
very many useful hints and directions calculated to lighten and facilitate in-door work.
The Department for Children and Youth, is prepared with special care
not only to amuse, but also to inculcate knowledge and sound moral principles.
Terms.The circulation of the American Agriculturist, (about 150,000) is so
large that it can be furnished at the low price of $1.50 a year; four copies, one year, for
$5; ten copies, one year, for $12; twenty or more copies, one year. $1 each; single
copies, 15 certs each. An extra copy to the one furnishing a club of ten or twenty.
TRY IT A YEAR.
ORANGE JUDD & CO.,
Publishers & Proprietors,
No. 245 Broadway, New-York City.

LIST OF
RURAL BOOKS
PUBLISHED AND FOR SALE BY
ORANGE JUDD & CO.,
NO. 245 BROADWAY, NEW YORK.
OF Any Book on this list will be forwarded, post-paid, to
any address in the United States, (except those Territories reached
by the Overland California Mail only, on receipt of the price.
66
66
Allen's (L. F.) Rural Architecture..$1 50
Allen's (R. L.) American Farm Book 1 50
Allen's (R. L.) Diseases of Domestic
Animals.......
..... 100
American Agricultural Annual,paper 50
cloth 75
American Horticultural Annual,paper 50
cloth 75
American Bird Fancier.
30
American Pomology.
3 00
American Rose Culturist..
30
American Weeds and Useful Plants. 1 75
Bement's Rabbit Fancier............ 30
Bommer's Method of Making Manures 25
Boussingault's Rural Economy...... 1 60
Breck's New Book of Flowers...... 1 75
Buist's Flower Garden Directory.... 1 50
Buist's Family Kitchen Gardener... 1 00
Chorlton's Grape Grower's Guide... 75
Cobbett's American Gardener...... 75
Cole's (S. W.) American Fruit Book 775
Cole's Veterinarian.....
75
Copeland's Country Life, 8vo, cloth. 5 00
Cotton Culture, (Lyman)...... 1 50
Cotton-Planter's Manual, (Turner).. 1 50
Dadd's (G. H.) Modern Horse Doctor 1 50
Dada's American Cattle Doctor..... 1 50
Dana's Muck Manual.......
1 25
Dog and Gun, (Hooper's), paper.... 30
cloth.... 60
Downing's Landscape Gardening.... 6 50
Draining for Profit and Health.. 1 50
Eastwood on Cranberry.....
Guenon on Milch Cows............. 75
Henderson's Gardening for Profit... 1 50
Herbert's Hints to Horse-Keepers... 1 75
Hop Culture.....
40
Hoopes on Evergreens..
Johnston's Agricultural Chemistry.. 1 75
Johnston's Elements of Agricultural
Chemistry........ ....... 1 50
Leuchar's How to Build Hot-Houses 1 50
Miles on the lIorse's Foot..........
Mohr on the Grape Vine....... 1 00
My Vineyard at Lakeview...
1 25
Norton's Scientific Agriculture......
Onion Culture.....
20
Our Farm of Four Acres, paper..... 30
cloth...... 60
Pardee on Strawberry Culture..
Peat and its Uses.....
1 25
Pedder's Land Measurer.
60
Quinby's Mysteries of Bee-Keeping. 1 50
Rural Annual, (Harris), 8 Nos. bound,
2 vols
each.. 1 50
Randall's Sheep Husbandry.......... 1 50
Randall's Fine Wool Sheep Husbandry1 00
Richardson on the Dog, paper...
30
cloth...... 60
Rivers' Miniature Fruit Garden. 1 00
Saunders' Domestic Poultry, paper..
cloth.. 75
Schenck's Gardener's Text Book....
Skillful Housewife...
75
Elliott's Western Fruit Grower's Guide1 50
Flax Culture......
50
French's Farm Drainage.
1 50
Field's (Thos. W.) Pear Culture..... 1 25
Fuller's Grape Culturist.....
1 50
Fuller's Small Fruit Culturist. 1 50
Fuller's Strawberry Culturist.
20
Gregory on Squash Culture..... 30
75
Stewart's (John) Stable Book.... 1 50
Thompson's Food of Animals... 1 00
Timothy Banker Papers.......
Tobacco Culture......
25
Warder's Hedges ani Evergreens... 1 50
Youatt and Spooner on the Horse... 1 50
Youatt and Martin on Cattle...... 1 50
Youаtt on the Hog...
1 00
Youаtt on Sheep......
1 00
60
66
65
66
66





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