The Variation of Animals and Plants under Domestication Volume 1
#14 in our series by Charles Darwin
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Title: The Variation of Animals and Plants under Domestication
Volume I
Author: Charles Darwin
Release Date: October, 2001 [Etext #2871]
Edition: 10
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Plants under Domestication Volume I by Charles Darwin
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THE VARIATION OF
ANIMALS AND PLANTS
UNDER DOMESTICATION
BY
CHARLES DARWIN, M.A., F.R.S., ETC.
IN TWO VOLUMES
VOLUME I.
PREFACE TO THE SECOND EDITION.
During the seven years which have elapsed since the publication in 1868 of
the first edition of this Work, I have continued to attend to the same
subjects, as far as lay in my power; and I have thus accumulated a large
body of additional facts, chiefly through the kindness of many
correspondents. Of these facts I have been able here to use only those
which seemed to me the more important. I have omitted some statements, and
corrected some errors, the discovery of which I owe to my reviewers. Many
additional references have been given. The eleventh chapter, and that on
Pangenesis, are those which have been most altered, parts having been re-
modelled; but I will give a list of the more important alterations for the
sake of those who may possess the first edition of this book.
CONTENTS.
INTRODUCTION.
CHAPTER 1.I.
DOMESTIC DOGS AND CATS.
ANCIENT VARIETIES OF THE DOG--RESEMBLANCE OF DOMESTIC DOGS IN VARIOUS
COUNTRIES TO NATIVE CANINE SPECIES--ANIMALS NOT ACQUAINTED WITH MAN AT
FIRST FEARLESS--DOGS RESEMBLING WOLVES AND JACKALS--HABIT OF BARKING
ACQUIRED AND LOST--FERAL DOGS--TAN-COLOURED EYE-SPOTS--PERIOD OF GESTATION-
-OFFENSIVE ODOUR--FERTILITY OF THE RACES WHEN CROSSED--DIFFERENCES IN THE
SEVERAL RACES IN PART DUE TO DESCENT FROM DISTINCT SPECIES--DIFFERENCES IN
THE SKULL AND TEETH--DIFFERENCES IN THE BODY, IN CONSTITUTION--FEW
IMPORTANT DIFFERENCES HAVE BEEN FIXED BY SELECTION--DIRECT ACTION OF
CLIMATE--WATER-DOGS WITH PALMATED FEET--HISTORY OF THE CHANGES WHICH
CERTAIN ENGLISH RACES OF THE DOG HAVE GRADUALLY UNDERGONE THROUGH
SELECTION--EXTINCTION OF THE LESS IMPROVED SUB-BREEDS.
CATS, CROSSED WITH SEVERAL SPECIES--DIFFERENT BREEDS FOUND ONLY IN
SEPARATED COUNTRIES--DIRECT EFFECTS OF THE CONDITIONS OF LIFE--FERAL CATS--
INDIVIDUAL VARIABILITY.
CHAPTER 1.II.
HORSES AND ASSES.
HORSE--DIFFERENCES IN THE BREEDS--INDIVIDUAL VARIABILITY OF--DIRECT EFFECTS
OF THE CONDITIONS OF LIFE--CAN WITHSTAND MUCH COLD--BREEDS MUCH MODIFIED BY
SELECTION--COLOURS OF THE HORSE--DAPPLING--DARK STRIPES ON THE SPINE, LEGS,
SHOULDERS, AND FOREHEAD--DUN-COLOURED HORSES MOST FREQUENTLY STRIPED--
STRIPES PROBABLY DUE TO REVERSION TO THE PRIMITIVE STATE OF THE HORSE.
ASSES--BREEDS OF--COLOUR OF--LEG- AND SHOULDER-STRIPES--SHOULDER-STRIPES
SOMETIMES ABSENT, SOMETIMES FORKED.
CHAPTER 1.III.
PIGS--CATTLE--SHEEP--GOATS.
PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICUS--TORFSCHWEIN--
JAPAN PIGS--FERTILITY OF CROSSED PIGS--CHANGES IN THE SKULL OF THE HIGHLY
CULTIVATED RACES--CONVERGENCE OF CHARACTER--GESTATION--SOLID-HOOFED SWINE--
CURIOUS APPENDAGES TO THE JAWS--DECREASE IN SIZE OF THE TUSKS--YOUNG PIGS
LONGITUDINALLY STRIPED--FERAL PIGS--CROSSED BREEDS.
CATTLE--ZEBU A DISTINCT SPECIES--EUROPEAN CATTLE PROBABLY DESCENDED FROM
THREE WILD FORMS--ALL THE RACES NOW FERTILE TOGETHER--BRITISH PARK CATTLE--
ON THE COLOUR OF THE ABORIGINAL SPECIES--CONSTITUTIONAL DIFFERENCES--SOUTH
AFRICAN RACES--SOUTH AMERICAN RACES--NIATA CATTLE--ORIGIN OF THE VARIOUS
RACES OF CATTLE.
SHEEP--REMARKABLE RACES OF--VARIATIONS ATTACHED TO THE MALE SEX--
ADAPTATIONS TO VARIOUS CONDITIONS--GESTATION OF--CHANGES IN THE WOOL--SEMI-
MONSTROUS BREEDS.
GOATS--REMARKABLE VARIATIONS OF.
CHAPTER 1.IV.
DOMESTIC RABBITS.
DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RABBIT--ANCIENT
DOMESTICATION--ANCIENT SELECTION--LARGE LOP-EARED RABBITS--VARIOUS BREEDS--
FLUCTUATING CHARACTERS--ORIGIN OF THE HIMALAYAN BREED--CURIOUS CASE OF
INHERITANCE--FERAL RABBITS IN JAMAICA AND THE FALKLAND ISLANDS--PORTO SANTO
FERAL RABBITS--OSTEOLOGICAL CHARACTERS--SKULL--SKULL OF HALF-LOP RABBITS--
VARIATIONS IN THE SKULL ANALOGOUS TO DIFFERENCES IN DIFFERENT SPECIES OF
HARES--VERTEBRAE--STERNUM--SCAPULA--EFFECTS OF USE AND DISUSE ON THE
PROPORTIONS OF THE LIMBS AND BODY--CAPACITY OF THE SKULL AND REDUCED SIZE
OF THE BRAIN--SUMMARY ON THE MODIFICATIONS OF DOMESTICATED RABBITS.
CHAPTER 1.V.
DOMESTIC PIGEONS.
ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS--INDIVIDUAL VARIABILITY--
VARIATIONS OF A REMARKABLE NATURE--OSTEOLOGICAL CHARACTERS: SKULL, LOWER
JAW, NUMBER OF VERTEBRAE--CORRELATION OF GROWTH: TONGUE WITH BEAK; EYELIDS
AND NOSTRILS WITH WATTLED SKIN--NUMBER OF WING-FEATHERS AND LENGTH OF WING-
-COLOUR AND DOWN--WEBBED AND FEATHERED FEET--ON THE EFFECTS OF DISUSE--
LENGTH OF FEET IN CORRELATION WITH LENGTH OF BEAK--LENGTH OF STERNUM,
SCAPULA, AND FURCULUM--LENGTH OF WINGS--SUMMARY ON THE POINTS OF DIFFERENCE
IN THE SEVERAL BREEDS.
CHAPTER 1.VI.
PIGEONS--continued.
ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES--HABITS OF
LIFE--WILD RACES OF THE ROCK-PIGEON--DOVECOTE-PIGEONS--PROOFS OF THE
DESCENT OF THE SEVERAL RACES FROM COLUMBA LIVIA--FERTILITY OF THE RACES
WHEN CROSSED--REVERSION TO THE PLUMAGE OF THE WILD ROCK-PIGEON--
CIRCUMSTANCES FAVOURABLE TO THE FORMATION OF THE RACES--ANTIQUITY AND
HISTORY OF THE PRINCIPAL RACES--MANNER OF THEIR FORMATION--SELECTION--
UNCONSCIOUS SELECTION--CARE TAKEN BY FANCIERS IN SELECTING THEIR BIRDS--
SLIGHTLY DIFFERENT STRAINS GRADUALLY CHANGE INTO WELL-MARKED BREEDS--
EXTINCTION OF INTERMEDIATE FORMS--CERTAIN BREEDS REMAIN PERMANENT, WHILST
OTHERS CHANGE--SUMMARY.
CHAPTER 1.VII.
FOWLS.
BRIEF DESCRIPTIONS OF THE CHIEF BREEDS--ARGUMENTS IN FAVOUR OF THEIR
DESCENT FROM SEVERAL SPECIES--ARGUMENTS IN FAVOUR OF ALL THE BREEDS HAVING
DESCENDED FROM GALLUS BANKIVA--REVERSION TO THE PARENT-STOCK IN COLOUR--
ANALOGOUS VARIATIONS--ANCIENT HISTORY OF THE FOWL--EXTERNAL DIFFERENCES
BETWEEN THE SEVERAL BREEDS--EGGS--CHICKENS--SECONDARY SEXUAL CHARACTERS--
WING-AND TAIL-FEATHERS, VOICE, DISPOSITION, ETC.--OSTEOLOGICAL DIFFERENCES
IN THE SKULL, VERTEBRAE, ETC.--EFFECTS OF USE AND DISUSE ON CERTAIN PARTS--
CORRELATION OF GROWTH.
CHAPTER 1.VIII.
DUCK--GOOSE--PEACOCK--TURKEY--GUINEA-FOWL--CANARY-BIRD--GOLD-FISH--HIVE-
BEES--SILK-MOTHS.
DUCKS, SEVERAL BREEDS OF--PROGRESS OF DOMESTICATION--ORIGIN OF FROM THE
COMMON WILD-DUCK--DIFFERENCES IN THE DIFFERENT BREEDS--OSTEOLOGICAL
DIFFERENCES--EFFECTS OF USE AND DISUSE ON THE LIMB-BONES.
GOOSE, ANCIENTLY DOMESTICATED--LITTLE VARIATION OF--SEBASTOPOL BREED.
PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.
TURKEY, BREEDS OF--CROSSED WITH THE UNITED STATES SPECIES--EFFECTS OF
CLIMATE ON.
GUINEA-FOWL, CANARY-BIRD, GOLD-FISH, HIVE-BEE.
SILK-MOTHS, SPECIES AND BREEDS OF--ANCIENTLY DOMESTICATED--CARE IN THEIR
SELECTION--DIFFERENCES IN THE DIFFERENT RACES--IN THE EGG, CATERPILLAR, AND
COCOON STATES--INHERITANCE OF CHARACTERS--IMPERFECT WINGS--LOST INSTINCTS--
CORRELATED CHARACTERS.
CHAPTER 1.IX.
CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.
PRELIMINARY REMARKS ON THE NUMBER AND PARENTAGE OF CULTIVATED PLANTS--FIRST
STEPS IN CULTIVATION--GEOGRAPHICAL DISTRIBUTION OF CULTIVATED PLANTS.
CEREALIA--DOUBTS ON THE NUMBER OF SPECIES--WHEAT: VARIETIES OF--INDIVIDUAL
VARIABILITY--CHANGED HABITS--SELECTION--ANCIENT HISTORY OF THE VARIETIES--
MAIZE: GREAT VARIATION OF--DIRECT ACTION OF CLIMATE ON.
CULINARY PLANTS--CABBAGES: VARIETIES OF, IN FOLIAGE AND STEMS, BUT NOT IN
OTHER PARTS--PARENTAGE OF--OTHER SPECIES OF BRASSICA--PEAS: AMOUNT OF
DIFFERENCE IN THE SEVERAL KINDS, CHIEFLY IN THE PODS AND SEED--SOME
VARIETIES CONSTANT, SOME HIGHLY VARIABLE--DO NOT INTERCROSS--BEANS--
POTATOES: NUMEROUS VARIETIES OF--DIFFER LITTLE EXCEPT IN THE TUBERS--
CHARACTERS INHERITED.
CHAPTER 1.X.
PLANTS continued--FRUITS--ORNAMENTAL TREES--FLOWERS.
FRUITS--GRAPES--VARY IN ODD AND TRIFLING PARTICULARS--MULBERRY--THE ORANGE
GROUP--SINGULAR RESULTS FROM CROSSING--PEACH AND NECTARINE--BUD-VARIATION--
ANALOGOUS VARIATION--RELATION TO THE ALMOND--APRICOT--PLUMS--VARIATION IN
THEIR STONES--CHERRIES--SINGULAR VARIETIES OF--APPLE--PEAR--STRAWBERRY--
INTERBLENDING OF THE ORIGINAL FORMS--GOOSEBERRY--STEADY INCREASE IN SIZE OF
THE FRUIT--VARIETIES OF--WALNUT--NUT--CUCURBITACEOUS PLANTS--WONDERFUL
VARIATION OF.
ORNAMENTAL TREES--THEIR VARIATION IN DEGREE AND KIND--ASH-TREE--SCOTCH-FIR-
-HAWTHORN.
FLOWERS--MULTIPLE ORIGIN OF MANY KINDS--VARIATION IN CONSTITUTIONAL
PECULIARITIES--KIND OF VARIATION--ROSES--SEVERAL SPECIES CULTIVATED--PANSY-
-DAHLIA--HYACINTH--HISTORY AND VARIATION OF.
CHAPTER 1.XI.
ON BUD-VARIATION, AND ON CERTAIN ANOMALOUS MODES OF REPRODUCTION AND
VARIATION.
BUD-VARIATION IN THE PEACH, PLUM, CHERRY, VINE, GOOSEBERRY, CURRANT, AND
BANANA, AS SHOWN BY THE MODIFIED FRUIT--IN FLOWERS: CAMELLIAS, AZALEAS,
CHRYSANTHEMUMS, ROSES, ETC.--ON THE RUNNING OF THE COLOUR IN CARNATIONS
--BUD-VARIATIONS IN LEAVES--VARIATIONS BY SUCKERS, TUBERS, AND BULBS--ON
THE BREAKING OF TULIPS--BUD-VARIATIONS GRADUATE INTO CHANGES CONSEQUENT ON
CHANGED CONDITIONS OF LIFE--GRAFT-HYBRIDS--ON THE SEGREGATION OF THE
PARENTAL CHARACTERS IN SEMINAL HYBRIDS BY BUD-VARIATION--ON THE DIRECT OR
IMMEDIATE ACTION OF FOREIGN POLLEN ON THE MOTHER-PLANT--ON THE EFFECTS OF A
PREVIOUS IMPREGNATION ON THE SUBSEQUENT OFFSPRING OF FEMALE ANIMALS--
CONCLUSION AND SUMMARY.
CHAPTER 1.XII.
INHERITANCE.
WONDERFUL NATURE OF INHERITANCE--PEDIGREES OF OUR DOMESTICATED ANIMALS--
INHERITANCE NOT DUE TO CHANCE--TRIFLING CHARACTERS INHERITED--DISEASES
INHERITED--PECULIARITIES IN THE EYE INHERITED--DISEASES IN THE HORSE--
LONGEVITY AND VIGOUR--ASYMMETRICAL DEVIATIONS OF STRUCTURE--POLYDACTYLISM
AND REGROWTH OF SUPERNUMERARY DIGITS AFTER AMPUTATION--CASES OF SEVERAL
CHILDREN SIMILARLY AFFECTED FROM NON-AFFECTED PARENTS--WEAK AND FLUCTUATING
INHERITANCE: IN WEEPING TREES, IN DWARFNESS, COLOUR OF FRUIT AND FLOWERS--
COLOUR OF HORSES--NON-INHERITANCE IN CERTAIN CASES--INHERITANCE OF
STRUCTURE AND HABITS OVERBORNE BY HOSTILE CONDITIONS OF LIFE, BY
INCESSANTLY RECURRING VARIABILITY, AND BY REVERSION--CONCLUSION.
LIST OF ILLUSTRATIONS.
FIGURE.
1. DUN DEVONSHIRE PONY, WITH SHOULDER, SPINAL, AND LEG STRIPES.
2. HEAD OF JAPAN OR MASKED PIG.
3. HEAD OF WILD BOAR, AND OF "GOLDEN DAYS," A PIG OF THE YORKSHIRE LARGE
BREED.
4. OLD IRISH PIG WITH JAW-APPENDAGES.
5. HALF-LOP RABBIT.
6. SKULL OF WILD RABBIT.
7. SKULL OF LARGE LOP-EARED RABBIT.
8. PART OF ZYGOMATIC ARCH, SHOWING THE PROJECTING END OF THE MALAR BONE OF
THE AUDITORY MEATUS, OF RABBITS.
9. POSTERIOR END OF SKULL, SHOWING THE INTER-PARIETAL BONE, OF RABBITS.
10. OCCIPITAL FORAMEN OF RABBITS.
11. SKULL OF HALF-LOP RABBIT.
12. ATLAS VERTEBRAE OF RABBITS.
13. THIRD CERVICAL VERTEBRAE OF RABBITS.
14. DORSAL VERTEBRAE, FROM SIXTH TO TENTH INCLUSIVE, OF RABBITS.
15. TERMINAL BONE OF STERNUM OF RABBITS.
16. ACROMION OF SCAPULA OF RABBITS.
17. THE ROCK-PIGEON, OR COLUMBA LIVIA.
18. ENGLISH POUTER.
19. ENGLISH CARRIER.
20. ENGLISH BARB.
21. ENGLISH FANTAIL.
22. AFRICAN OWL.
23. SHORT-FACED ENGLISH TUMBLER.
24. SKULLS OF PIGEONS, VIEWED LATERALLY.
25. LOWER JAWS OF PIGEONS, SEEN FROM ABOVE.
26. SKULL OF RUNT, SEEN FROM ABOVE.
27. LATERAL VIEW OF JAWS OF PIGEONS.
28. SCAPULAE OF PIGEONS.
29. FURCULA OF PIGEONS.
30. SPANISH FOWL.
31. HAMBURGH FOWL.
32. POLISH FOWL.
33. OCCIPITAL FORAMEN OF THE SKULLS OF FOWLS.
34. SKULLS OF FOWLS, VIEWED FROM ABOVE, A LITTLE OBLIQUELY.
35. LONGITUDINAL SECTIONS OF SKULLS OF FOWLS, VIEWED LATERALLY.
36. SKULL OF HORNED FOWL, VIEWED FROM ABOVE, A LITTLE OBLIQUELY.
37. SIXTH CERVICAL VERTEBRAE OF FOWLS, VIEWED LATERALLY.
38. EXTREMITY OF THE FURCULA OF FOWLS, VIEWED LATERALLY.
39. SKULLS OF DUCKS, VIEWED LATERALLY, REDUCED TO TWO-THIRDS OF THE
NATURAL SIZE.
40. CERVICAL VERTEBRAE OF DUCKS, OF NATURAL SIZE.
41. PODS OF THE COMMON PEA.
42. PEACH AND ALMOND STONES, OF NATURAL SIZE, VIEWED EDGEWAYS.
43. PLUM STONES, OF NATURAL SIZE, VIEWED LATERALLY.
TABLE 1: PRINCIPAL ADDITIONS AND CORRECTIONS IN THIS (SECOND) EDITION.
First Edition, Volume I., Page 34.
Second Edition Volume I., Page 35.
Dr. Burt Wilder's observations on the brains of different breeds of the
Dog.
First Edition, Volume I., Page 38.
Second Edition Volume I., Page 40.
Degeneracy of Dogs imported into Guinea.
First Edition, Volume I., Page 51.
Second Edition Volume I., Page 54.
Difference in the number of the lumbar vertebrae in the races or species of
the Horse.
First Edition, Volume I., Page 102.
Second Edition Volume I., Page 106.
Hairy appendages to the throats of Goats.
First Edition, Volume I., Page 162.
Second Edition Volume I., Page 170.
Sexual differences in colour in the domestic Pigeon.
First Edition, Volume I., Page 217.
Second Edition Volume I., Page 228.
Movements like those of the Tumbler-pigeon, caused by injury to the brain.
First Edition, Volume I., Page 290.
Second Edition Volume I., Page 306.
Additional facts with respect to the Black-shouldered Peacock.
First Edition, Volume I., Page 296.
Second Edition Volume I., Page 312.
Ancient selection of Gold-fish in China.
First Edition, Volume I., Page 314.
Second Edition Volume I., Page 332.
Major Hallett's 'Pedigree Wheat.'
First Edition, Volume I., Page 326.
Second Edition Volume I., Page 345.
The common radish descended from Raphanus raphanistrum.
First Edition, Volume I., Page 374.
Second Edition Volume I., Page 398.
Several additional cases of bud-variation given.
First Edition, Volume I., Page 396.
Second Edition Volume I., Page 420.
An abstract of all the cases recently published of graft-hybrids in the
potato, together with a general summary on graft-hybridisation.
First Edition, Volume I., Page 399.
Second Edition Volume I., Page 429.
An erroneous statement with respect to the pollen of the date-palm
affecting the fruit of the Chamaerops omitted.
First Edition, Volume I., Page 400.
Second Edition Volume I., Page 430.
New cases of the direct action of pollen on the mother-plant.
First Edition, Volume I., Page 404.
Second Edition Volume I., Page 435.
Additional and remarkable instances of the action of the male parent on the
future progeny of the female.
First Edition, Volume II., Page 14.
Second Edition Volume I., Page 459.
An erroneous statement corrected, with respect to the regrowth of
supernumerary digits after amputation.
First Edition, Volume II., Page 23.
Second Edition Volume I., Page 467.
Additional facts with respect to the inherited effects of circumcision.
First Edition, Volume II., Page 23.
Second Edition Volume I., Page 467.
Dr. Brown-Sequard on the inherited effects of operations on the Guinea-pig.
First Edition, Volume II., Page 24.
Second Edition Volume I., Page 469.
Other cases of inherited mutilations.
First Edition, Volume II., Page 43.
Second Edition Volume II., Page 17.
An additional case of reversion due to a cross.
First Edition, Volume II., Page 72.
Second Edition Volume II., Page 48.
Inheritance as limited by sex.
First Edition, Volume II., Page 105.
Second Edition Volume II., Page 83.
Two varieties of maize which cannot be crossed.
First Edition, Volume II., Page 120.
Second Edition Volume II., Page 99.
Some additional facts on the advantages of cross-breeding in animals.
First Edition, Volume II., Page 123.
Second Edition Volume II., Page 103.
Discussion on the effects of close interbreeding in the case of man.
First Edition, Volume II., Page 135 to 141.
Second Edition Volume II., Page 117 to 122.
Additional cases of plants sterile with pollen from the same plant.
First Edition, Volume II., Page 149.
Second Edition Volume II., Page 131.
Mr. Sclater on the infertility of animals under confinement.
First Edition, Volume II., Page 152.
Second Edition Volume II., Page 134.
The Aperea a distinct species from the Guinea-pig.
First Edition, Volume II., Page 230.
Second Edition Volume II., Page 215.
Professor Jager on hawks killing light-coloured pigeons.
First Edition, Volume II., Page 273.
Second Edition Volume II., Page 262.
Professor Weismann on the effects of isolation in the development of
species.
First Edition, Volume II., Page 281.
Second Edition Volume II., Page 271.
The direct action of the conditions of life in causing variation.
First Edition, Volume II., Page 317.
Second Edition Volume II., Page 309.
Mr. Romanes on rudimentary parts.
First Edition, Volume II., Page 324 to 328.
Second Edition Volume II., Page 316 to 327.
Some additional cases of correlated variability.
First Edition, Volume II., Page 339.
Second Edition Volume II., Page 333.
On Geoffroy St.-Hilaire's law of "soi pour soi."
First Edition, Volume II., Page 357 to 404.
Second Edition Volume II., Page 349 to 399.
The chapter on Pangenesis has been largely altered and re-modelled; but the
essential principles remain the same.
THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION.
INTRODUCTION.
The object of this work is not to describe all the many races of animals
which have been domesticated by man, and of the plants which have been
cultivated by him; even if I possessed the requisite knowledge, so gigantic
an undertaking would be here superfluous. It is my intention to give under
the head of each species only such facts as I have been able to collect or
observe, showing the amount and nature of the changes which animals and
plants have undergone whilst under man's dominion, or which bear on the
general principles of variation. In one case alone, namely in that of the
domestic pigeon, I will describe fully all the chief races, their history,
the amount and nature of their differences, and the probable steps by which
they have been formed. I have selected this case, because, as we shall
hereafter see, the materials are better than in any other; and one case
fully described will in fact illustrate all others. But I shall also
describe domesticated rabbits, fowls, and ducks, with considerable fulness.
The subjects discussed in this volume are so connected that it is not a
little difficult to decide how they can be best arranged. I have determined
in the first part to give, under the heads of the various animals and
plants, a large body of facts, some of which may at first appear but little
related to our subject, and to devote the latter part to general
discussions. Whenever I have found it necessary to give numerous details,
in support of any proposition or conclusion, small type has been used.
(Here shown with [].) The reader will, I think, find this plan a
convenience, for, if he does not doubt the conclusion or care about the
details, he can easily pass them over; yet I may be permitted to say that
some of the discussions thus printed deserve attention, at least from the
professed naturalist.
It may be useful to those who have read nothing about Natural Selection, if
I here give a brief sketch of the whole subject and of its bearing on the
origin of species. (Introduction/1. To any one who has attentively read my
'Origin of Species' this Introduction will be superfluous. As I stated in
that work that I should soon publish the facts on which the conclusions
given in it were founded, I here beg permission to remark that the great
delay in publishing this first work has been caused by continued ill-
health.) This is the more desirable, as it is impossible in the present
work to avoid many allusions to questions which will be fully discussed in
future volumes.
From a remote period, in all parts of the world, man has subjected many
animals and plants to domestication or culture. Man has no power of
altering the absolute conditions of life; he cannot change the climate of
any country; he adds no new element to the soil; but he can remove an
animal or plant from one climate or soil to another, and give it food on
which it did not subsist in its natural state. It is an error to speak of
man "tampering with nature" and causing variability. If a man drops a piece
of iron into sulphuric acid, it cannot be said strictly that he makes the
sulphate of iron, he only allows their elective affinities to come into
play. If organic beings had not possessed an inherent tendency to vary, man
could have done nothing. (Introduction/2. M. Pouchet has recently
('Plurality of Races' English Translation 1864 page 83 etc.) insisted that
variation under domestication throws no light on the natural modification
of species. I cannot perceive the force of his arguments, or, to speak more
accurately, of his assertions to this effect.) He unintentionally exposes
his animals and plants to various conditions of life, and variability
supervenes, which he cannot even prevent or check. Consider the simple case
of a plant which has been cultivated during a long time in its native
country, and which consequently has not been subjected to any change of
climate. It has been protected to a certain extent from the competing roots
of plants of other kinds; it has generally been grown in manured soil; but
probably not richer than that of many an alluvial flat; and lastly, it has
been exposed to changes in its conditions, being grown sometimes in one
district and sometimes in another, in different soils. Under such
circumstances, scarcely a plant can be named, though cultivated in the
rudest manner, which has not given birth to several varieties. It can
hardly be maintained that during the many changes which this earth has
undergone, and during the natural migrations of plants from one land or
island to another, tenanted by different species, that such plants will not
often have been subjected to changes in their conditions analogous to those
which almost inevitably cause cultivated plants to vary. No doubt man
selects varying individuals, sows their seeds, and again selects their
varying offspring. But the initial variation on which man works, and
without which he can do nothing, is caused by slight changes in the
conditions of life, which must often have occurred under nature. Man,
therefore, may be said to have been trying an experiment on a gigantic
scale; and it is an experiment which nature during the long lapse of time
has incessantly tried. Hence it follows that the principles of
domestication are important for us. The main result is that organic beings
thus treated have varied largely, and the variations have been inherited.
This has apparently been one chief cause of the belief long held by some
few naturalists that species in a state of nature undergo change.
I shall in this volume treat, as fully as my materials permit, the whole
subject of variation under domestication. We may thus hope to obtain some
light, little though it be, on the causes of variability,--on the laws
which govern it, such as the direct action of climate and food, the effects
of use and disuse, and of correlation of growth,--and on the amount of
change to which domesticated organisms are liable. We shall learn something
of the laws of inheritance, of the effects of crossing different breeds,
and on that sterility which often supervenes when organic beings are
removed from their natural conditions of life, and likewise when they are
too closely interbred. During this investigation we shall see that the
principle of Selection is highly important. Although man does not cause
variability and cannot even prevent it, he can select, preserve, and
accumulate the variations given to him by the hand of nature almost in any
way which he chooses; and thus he can certainly produce a great result.
Selection may be followed either methodically and intentionally, or
unconsciously and unintentionally. Man may select and preserve each
successive variation, with the distinct intention of improving and altering
a breed, in accordance with a preconceived idea; and by thus adding up
variations, often so slight as to be imperceptible by an uneducated eye, he
has effected wonderful changes and improvements. It can, also, be clearly
shown that man, without any intention or thought of improving the breed, by
preserving in each successive generation the individuals which he prizes
most, and by destroying the worthless individuals, slowly, though surely,
induces great changes. As the will of man thus comes into play, we can
understand how it is that domesticated breeds show adaptation to his wants
and pleasures. We can further understand how it is that domestic races of
animals and cultivated races of plants often exhibit an abnormal character,
as compared with natural species; for they have been modified not for their
own benefit, but for that of man.
In another work I shall discuss, if time and health permit, the variability
of organic beings in a state of nature; namely, the individual differences
presented by animals and plants, and those slightly greater and generally
inherited differences which are ranked by naturalists as varieties or
geographical races. We shall see how difficult, or rather how impossible it
often is, to distinguish between races and sub-species, as the less well-
marked forms have sometimes been denominated; and again between sub-species
and true species. I shall further attempt to show that it is the common and
widely ranging, or, as they may be called, the dominant species, which most
frequently vary; and that it is the large and flourishing genera which
include the greatest number of varying species. Varieties, as we shall see,
may justly be called incipient species.
But it may be urged, granting that organic beings in a state of nature
present some varieties,--that their organisation is in some slight degree
plastic; granting that many animals and plants have varied greatly under
domestication, and that man by his power of selection has gone on
accumulating such variations until he has made strongly marked and firmly
inherited races; granting all this, how, it may be asked, have species
arisen in a state of nature? The differences between natural varieties are
slight; whereas the differences are considerable between the species of the
same genus, and great between the species of distinct genera. How do these
lesser differences become augmented into the greater difference? How do
varieties, or as I have called them incipient species, become converted
into true and well-defined species? How has each new species been adapted
to the surrounding physical conditions, and to the other forms of life on
which it in any way depends? We see on every side of us innumerable
adaptations and contrivances, which have justly excited the highest
admiration of every observer. There is, for instance, a fly (Cecidomyia
(Introduction/3. Leon Dufour in 'Annales des Science. Nat.' (3rd series,
Zoolog.) tome 5 page 6.)) which deposits its eggs within the stamens of a
Scrophularia, and secretes a poison which produces a gall, on which the
larva feeds; but there is another insect (Misocampus) which deposits its
eggs within the body of the larva within the gall, and is thus nourished by
its living prey; so that here a hymenopterous insect depends on a dipterous
insect, and this depends on its power of producing a monstrous growth in a
particular organ of a particular plant. So it is, in a more or less plainly
marked manner, in thousands and tens of thousands of cases, with the lowest
as well as with the highest productions of nature.
This problem of the conversion of varieties into species,--that is, the
augmentation of the slight differences characteristic of varieties into the
greater differences characteristic of species and genera, including the
admirable adaptations of each being to its complex organic and inorganic
conditions of life,--has been briefly treated in my 'Origin of Species.' It
was there shown that all organic beings, without exception, tend to
increase at so high a ratio, that no district, no station, not even the
whole surface of the land or the whole ocean, would hold the progeny of a
single pair after a certain number of generations. The inevitable result is
an ever-recurrent Struggle for Existence. It has truly been said that all
nature is at war; the strongest ultimately prevail, the weakest fail; and
we well know that myriads of forms have disappeared from the face of the
earth. If then organic beings in a state of nature vary even in a slight
degree, owing to changes in the surrounding conditions, of which we have
abundant geological evidence, or from any other cause; if, in the long
course of ages, inheritable variations ever arise in any way advantageous
to any being under its excessively complex and changing relations of life;
and it would be a strange fact if beneficial variations did never arise,
seeing how many have arisen which man has taken advantage of for his own
profit or pleasure; if then these contingencies ever occur, and I do not
see how the probability of their occurrence can be doubted, then the severe
and often-recurrent struggle for existence will determine that those
variations, however slight, which are favourable shall be preserved or
selected, and those which are unfavourable shall be destroyed.
This preservation, during the battle for life, of varieties which possess
any advantage in structure, constitution, or instinct, I have called
Natural Selection; and Mr. Herbert Spencer has well expressed the same idea
by the Survival of the Fittest. The term "natural selection" is in some
respects a bad one, as it seems to imply conscious choice; but this will be
disregarded after a little familiarity. No one objects to chemists speaking
of "elective affinity;" and certainly an acid has no more choice in
combining with a base, than the conditions of life have in determining
whether or not a new form be selected or preserved. The term is so far a
good one as it brings into connection the production of domestic races by
man's power of selection, and the natural preservation of varieties and
species in a state of nature. For brevity sake I sometimes speak of natural
selection as an intelligent power;--in the same way as astronomers speak of
the attraction of gravity as ruling the movements of the planets, or as
agriculturists speak of man making domestic races by his power of
selection. In the one case, as in the other, selection does nothing without
variability, and this depends in some manner on the action of the
surrounding circumstances on the organism. I have, also, often personified
the word Nature; for I have found it difficult to avoid this ambiguity; but
I mean by nature only the aggregate action and product of many natural
laws,--and by laws only the ascertained sequence of events.
It has been shown from many facts that the largest amount of life can be
supported on each area, by great diversification or divergence in the
structure and constitution of its inhabitants. We have, also, seen that the
continued production of new forms through natural selection, which implies
that each new variety has some advantage over others, inevitably leads to
the extermination of the older and less improved forms. These latter are
almost necessarily intermediate in structure, as well as in descent,
between the last-produced forms and their original parent-species. Now, if
we suppose a species to produce two or more varieties, and these in the
course of time to produce other varieties, the principal of good being
derived from diversification of structure will generally lead to the
preservation of the most divergent varieties; thus the lesser differences
characteristic of varieties come to be augmented into the greater
differences characteristic of species, and, by the extermination of the
older intermediate forms, new species end by being distinctly defined
objects. Thus, also, we shall see how it is that organic beings can be
classed by what is called a natural method in distinct groups--species
under genera, and genera under families.
As all the inhabitants of each country may be said, owing to their high
rate of reproduction, to be striving to increase in numbers; as each form
comes into competition with many other forms in the struggle for life,--for
destroy any one and its place will be seized by others; as every part of
the organisation occasionally varies in some slight degree, and as natural
selection acts exclusively by the preservation of variations which are
advantageous under the excessively complex conditions to which each being
is exposed, no limit exists to the number, singularity, and perfection of
the contrivances and co-adaptations which may thus be produced. An animal
or a plant may thus slowly become related in its structure and habits in
the most intricate manner to many other animals and plants, and to the
physical conditions of its home. Variations in the organisation will in
some cases be aided by habit, or by the use and disuse of parts, and they
will be governed by the direct action of the surrounding physical
conditions and by correlation of growth.
On the principles here briefly sketched out, there is no innate or
necessary tendency in each being to its own advancement in the scale of
organisation. We are almost compelled to look at the specialisation or
differentiation of parts or organs for different functions as the best or
even sole standard of advancement; for by such division of labour each
function of body and mind is better performed. And as natural selection
acts exclusively through the preservation of profitable modifications of
structure, and as the conditions of life in each area generally become more
and more complex from the increasing number of different forms which
inhabit it and from most of these forms acquiring a more and more perfect
structure, we may confidently believe, that, on the whole, organisation
advances. Nevertheless a very simple form fitted for very simple conditions
of life might remain for indefinite ages unaltered or unimproved; for what
would it profit an infusorial animalcule, for instance, or an intestinal
worm, to become highly organised? Members of a high group might even
become, and this apparently has often occurred, fitted for simpler
conditions of life; and in this case natural selection would tend to
simplify or degrade the organisation, for complicated mechanism for simple
actions would be useless or even disadvantageous.
The arguments opposed to the theory of Natural Selection, have been
discussed in my 'Origin of Species,' as far as the size of that work
permitted, under the following heads: the difficulty in understanding how
very simple organs have been converted by small and graduated steps into
highly perfect and complex organs; the marvellous facts of Instinct; the
whole question of Hybridity; and, lastly, the absence in our known
geological formations of innumerable links connecting all allied species.
Although some of these difficulties are of great weight, we shall see that
many of them are explicable on the theory of natural selection, and are
otherwise inexplicable.
In scientific investigations it is permitted to invent any hypothesis, and
if it explains various large and independent classes of facts it rises to
the rank of a well-grounded theory. The undulations of the ether and even
its existence are hypothetical, yet every one now admits the undulatory
theory of light. The principle of natural selection may be looked at as a
mere hypothesis, but rendered in some degree probable by what we positively
know of the variability of organic beings in a state of nature,--by what we
positively know of the struggle for existence, and the consequent almost
inevitable preservation of favourable variations,--and from the analogical
formation of domestic races. Now this hypothesis may be tested,--and this
seems to me the only fair and legitimate manner of considering the whole
question,--by trying whether it explains several large and independent
classes of facts; such as the geological succession of organic beings,
their distribution in past and present times, and their mutual affinities
and homologies. If the principle of natural selection does explain these
and other large bodies of facts, it ought to be received. On the ordinary
view of each species having been independently created, we gain no
scientific explanation of any one of these facts. We can only say that it
has so pleased the Creator to command that the past and present inhabitants
of the world should appear in a certain order and in certain areas; that He
has impressed on them the most extraordinary resemblances, and has classed
them in groups subordinate to groups. But by such statements we gain no new
knowledge; we do not connect together facts and laws; we explain nothing.
It was the consideration of such large groups of facts as these which first
led me to take up the present subject. When I visited during the voyage of
H.M.S. "Beagle," the Galapagos Archipelago, situated in the Pacific Ocean
about 500 miles from South America, I found myself surrounded by peculiar
species of birds, reptiles, and plants, existing nowhere else in the world.
Yet they nearly all bore an American stamp. In the song of the mocking-
thrush, in the harsh cry of the carrion-hawk, in the great candlestick-like
opuntias, I clearly perceived the neighbourhood of America, though the
islands were separated by so many miles of ocean from the mainland, and
differed much in their geological constitution and climate. Still more
surprising was the fact that most of the inhabitants of each separate
island in this small archipelago were specifically different, though most
closely related to each other. The archipelago, with its innumerable
craters and bare streams of lava, appeared to be of recent origin; and thus
I fancied myself brought near to the very act of creation. I often asked
myself how these many peculiar animals and plants had been produced: the
simplest answer seemed to be that the inhabitants of the several islands
had descended from each other, undergoing modification in the course of
their descent; and that all the inhabitants of the archipelago were
descended from those of the nearest land, namely America, whence colonists
would naturally have been derived. But it long remained to me an
inexplicable problem how the necessary degree of modification could have
been effected, and it would have thus remained for ever, had I not studied
domestic productions, and thus acquired a just idea of the power of
Selection. As soon as I had fully realised this idea, I saw, on reading
Malthus on Population, that Natural Selection was the inevitable result of
the rapid increase of all organic beings; for I was prepared to appreciate
the struggle for existence by having long studied the habits of animals.
Before visiting the Galapagos I had collected many animals whilst
travelling from north to south on both sides of America, and everywhere,
under conditions of life as different as it is possible to conceive,
American forms were met with--species replacing species of the same
peculiar genera. Thus it was when the Cordilleras were ascended, or the
thick tropical forests penetrated, or the fresh waters of America searched.
Subsequently I visited other countries, which in all their conditions of
life were incomparably more like parts of South America, than the different
parts of that continent are to each other; yet in these countries, as in
Australia or Southern Africa, the traveller cannot fail to be struck with
the entire difference of their productions. Again the reflection was forced
on me that community of descent from the early inhabitants of South America
would alone explain the wide prevalence of American types throughout that
immense area.
To exhume with one's own hands the bones of extinct and gigantic quadrupeds
brings the whole question of the succession of species vividly before one's
mind; and I found in South America great pieces of tesselated armour
exactly like, but on a magnificent scale, that covering the pigmy
armadillo; I had found great teeth like those of the living sloth, and
bones like those of the cavy. An analogous succession of allied forms had
been previously observed in Australia. Here then we see the prevalence, as
if by descent, in time as in space, of the same types in the same areas;
and in neither the case does the similarity of the conditions by any means
seem sufficient to account for the similarity of the forms of life. It is
notorious that the fossil remains of closely consecutive formations are
closely allied in structure, and we can at once understand the fact if they
are closely allied by descent. The succession of the many distinct species
of the same genus throughout the long series of geological formations seems
to have been unbroken or continuous. New species come in gradually one by
one. Ancient and extinct forms of life are often intermediate in character,
like the words of a dead language with respect to its several offshoots or
living tongues. All these facts seemed to me to point to descent with
modification as the means of production of new species.
The innumerable past and present inhabitants of the world are connected
together by the most singular and complex affinities, and can be classed in
groups under groups, in the same manner as varieties can be classed under
species and sub-varieties under varieties, but with much higher grades of
difference. These complex affinities and the rules for classification,
receive a rational explanation on the theory of descent, combined with the
principle of natural selection, which entails divergence of character and
the extinction of intermediate forms. How inexplicable is the similar
pattern of the hand of a man, the foot of a dog, the wing of a bat, the
flipper of a seal, on the doctrine of independent acts of creation! how
simply explained on the principle of the natural selection of successive
slight variations in the diverging descendants from a single progenitor! So
it is with certain parts or organs in the same individual animal or plant,
for instance, the jaws and legs of a crab, or the petals, stamens, and
pistils of a flower. During the many changes to which in the course of time
organic beings have been subjected, certain organs or parts have
occasionally become at first of little use and ultimately superfluous; and
the retention of such parts in a rudimentary and useless condition is
intelligible on the theory of descent. It can be shown that modifications
of structure are generally inherited by the offspring at the same age at
which each successive variation appeared in the parents; it can further be
shown that variations do not commonly supervene at a very early period of
embryonic growth, and on these two principles we can understand that most
wonderful fact in the whole circuit of natural history, namely, the close
similarity of the embryos within the same great class--for instance, those
of mammals, birds, reptiles, and fish.
It is the consideration and explanation of such facts as these which has
convinced me that the theory of descent with modification by means of
natural selection is in the main true. These facts have as yet received no
explanation on the theory of independent Creation; they cannot be grouped
together under one point of view, but each has to be considered as an
ultimate fact. As the first origin of life on this earth, as well as the
continued life of each individual, is at present quite beyond the scope of
science, I do not wish to lay much stress on the greater simplicity of the
view of a few forms or of only one form having been originally created,
instead of innumerable miraculous creations having been necessary at
innumerable periods; though this more simple view accords well with
Maupertuis's philosophical axiom of "least action."
In considering how far the theory of natural selection may be extended,
--that is, in determining from how many progenitors the inhabitants of the
world have descended,--we may conclude that at least all the members of the
same class have descended from a single ancestor. A number of organic
beings are included in the same class, because they present, independently
of their habits of life, the same fundamental type of structure, and
because they graduate into each other. Moreover, members of the same class
can in most cases be shown to be closely alike at an early embryonic age.
These facts can be explained on the belief of their descent from a common
form; therefore it may be safely admitted that all the members of the same
class are descended from one progenitor. But as the members of quite
distinct classes have something in common in structure and much in common
in constitution, analogy would lead us one step further, and to infer as
probable that all living creatures are descended from a single prototype.
I hope that the reader will pause before coming to any final and hostile
conclusion on the theory of natural selection. The reader may consult my
'Origin of Species' for a general sketch of the whole subject; but in that
work he has to take many statements on trust. In considering the theory of
natural selection, he will assuredly meet with weighty difficulties, but
these difficulties relate chiefly to subjects--such as the degree of
perfection of the geological record, the means of distribution, the
possibility of transitions in organs, etc.--on which we are confessedly
ignorant; nor do we know how ignorant we are. If we are much more ignorant
than is generally supposed, most of these difficulties wholly disappear.
Let the reader reflect on the difficulty of looking at whole classes of
facts from a new point of view. Let him observe how slowly, but surely, the
noble views of Lyell on the gradual changes now in progress on the earth's
surface have been accepted as sufficient to account for all that we see in
its past history. The present action of natural selection may seem more or
less probable; but I believe in the truth of the theory, because it
collects, under one point of view, and gives a rational explanation of,
many apparently independent classes of facts. (Introduction/4. In treating
the several subjects included in the present and my other works I have
continually been led to ask for information from many zoologists,
botanists, geologists, breeders of animals, and horticulturists, and I have
invariably received from them the most generous assistance. Without such
aid I could have effected little. I have repeatedly applied for information
and specimens to foreigners, and to British merchants and officers of the
Government residing in distant lands, and, with the rarest exceptions, I
have received prompt, open-handed, and valuable assistance. I cannot
express too strongly my obligations to the many persons who have assisted
me, and who, I am convinced, would be equally willing to assist others in
any scientific investigation.)
CHAPTER 1.I.
DOMESTIC DOGS AND CATS.
ANCIENT VARIETIES OF THE DOG.
RESEMBLANCE OF DOMESTIC DOGS IN VARIOUS COUNTRIES TO NATIVE CANINE SPECIES.
ANIMALS NOT ACQUAINTED WITH MAN AT FIRST FEARLESS.
DOGS RESEMBLING WOLVES AND JACKALS.
HABIT OF BARKING ACQUIRED AND LOST.
FERAL DOGS.
TAN-COLOURED EYE-SPOTS.
PERIOD OF GESTATION.
OFFENSIVE ODOUR.
FERTILITY OF THE RACES WHEN CROSSED.
DIFFERENCES IN THE SEVERAL RACES IN PART DUE TO DESCENT FROM DISTINCT
SPECIES.
DIFFERENCES IN THE SKULL AND TEETH.
DIFFERENCES IN THE BODY, IN CONSTITUTION.
FEW IMPORTANT DIFFERENCES HAVE BEEN FIXED BY SELECTION.
DIRECT ACTION OF CLIMATE.
WATER-DOGS WITH PALMATED FEET.
HISTORY OF THE CHANGES WHICH CERTAIN ENGLISH RACES OF THE DOG HAVE
GRADUALLY UNDERGONE THROUGH SELECTION.
EXTINCTION OF THE LESS IMPROVED SUB-BREEDS.
CATS, CROSSED WITH SEVERAL SPECIES.
DIFFERENT BREEDS FOUND ONLY IN SEPARATED COUNTRIES.
DIRECT EFFECTS OF THE CONDITIONS OF LIFE.
FERAL CATS.
INDIVIDUAL VARIABILITY.
The first and chief point of interest in this chapter is, whether the
numerous domesticated varieties of the dog have descended from a single
wild species, or from several. Some authors believe that all have descended
from the wolf, or from the jackal, or from an unknown and extinct species.
Others again believe, and this of late has been the favourite tenet, that
they have descended from several species, extinct and recent, more or less
commingled together. We shall probably never be able to ascertain their
origin with certainty. Palaeontology (1/1. Owen 'British Fossil Mammals'
pages 123 to 133. Pictet 'Traite de Pal.' 1853 tome 1 page 202. De
Blainville in his 'Osteographie, Canidae' page 142 has largely discussed
the whole subject, and concludes that the extinct parent of all
domesticated dogs came nearest to the wolf in organisation, and to the
jackal in habits. See also Boyd Dawkins, 'Cave Hunting' 1874 page 131 etc.
and his other publications. Jeitteles has discussed in great detail the
character of the breeds of pre-historic dogs: 'Die vorgeschichtlichen
Alterthumer der Stadt Olmutz' II. Theil, 1872 page 44 to end.) does not
throw much light on the question, owing, on the one hand, to the close
similarity of the skulls of extinct as well as living wolves and jackals,
and owing, on the other hand, to the great dissimilarity of the skulls of
the several breeds of the domestic dogs. It seems, however, that remains
have been found in the later tertiary deposits more like those of a large
dog than of a wolf, which favours the belief of De Blainville that our dogs
are the descendants of a single extinct species. On the other hand, some
authors go so far as to assert that every chief domestic breed must have
had its wild prototype. This latter view is extremely improbable: it allows
nothing for variation; it passes over the almost monstrous character of
some of the breeds; and it almost necessarily assumes that a large number
of species have become extinct since man domesticated the dog; whereas we
plainly see that wild members of the dog-family are extirpated by human
agency with much difficulty; even so recently as 1710 the wolf existed in
so small an island as Ireland.
The reasons which have led various authors to infer that our dogs have
descended from more than one wild species are as follows. (1/2. Pallas, I
believe, originated this doctrine in 'Act. Acad. St. Petersburgh' 1780 Part
2. Ehrenberg has advocated it, as may be seen in De Blainville's
'Osteographie' page 79. It has been carried to an extreme extent by Col.
Hamilton Smith in the 'Naturalist Library' volumes 9 and 10. Mr. W.C.
Martin adopts it in his excellent 'History of the Dog' 1845; as does Dr.
Morton, as well as Nott and Gliddon, in the United States. Prof. Low, in
his 'Domesticated Animals' 1845 page 666, comes to this same conclusion. No
one has argued on this side with more clearness and force than the late
James Wilson, of Edinburgh, in various papers read before the Highland
Agricultural and Wernerian Societies. Isidore Geoffroy Saint-Hilaire
('Hist. Nat. Gen.' 1860 tome 3 page 107), though he believes that most dogs
have descended from the jackal, yet inclines to the belief that some are
descended from the wolf. Prof. Gervais ('Hist. Nat. Mamm.' 1855 tome 2 page
69, referring to the view that all the domestic races are the modified
descendants of a single species, after a long discussion, says, "Cette
opinion est, suivant nous du moins, la moins probable.") Firstly, the great
difference between the several breeds; but this will appear of
comparatively little weight, after we shall have seen how great are the
differences between the several races of various domesticated animals which
certainly have descended from a single parent-form. Secondly, the more
important fact, that, at the most anciently known historical periods,
several breeds of the dog existed, very unlike each other, and closely
resembling or identical with breeds still alive.
We will briefly run back through the historical records. The materials are
remarkably deficient between the fourteenth century and the Roman classical
period. (1/3. Berjeau 'The Varieties of the Dog; in old Sculptures and
Pictures' 1863. 'Der Hund' von Dr. F.L. Walther, Giessen 1817 s. 48: this
author seems carefully to have studied all classical works on the subject.
See also Volz 'Beitrage zur Kulturgeschichte' Leipzig 1852 s. 115, 'Youatt
on the Dog' 1845 page 6. A very full history is given by De Blainville in
his 'Osteographie, Canidae.') At this latter period various breeds, namely
hounds, house-dogs, lapdogs, etc, existed; but, as Dr. Walther has
remarked, it is impossible to recognise the greater number with any
certainty. Youatt, however, gives a drawing of a beautiful sculpture of two
greyhound puppies from the Villa of Antoninus. On an Assyrian monument,
about 640 B.C.,an enormous mastiff (1/4. I have seen drawings of this dog
from the tomb of the son of Esar Haddon, and clay models in the British
Museum. Nott and Gliddon, in their 'Types of Mankind' 1854 page 393, give a
copy of these drawings. This dog has been called a Thibetan mastiff, but
Mr. H.A. Oldfield, who is familiar with the so-called Thibet mastiff, and
has examined the drawings in the British Museum, informs me that he
considers them different.) is figured; and according to Sir H. Rawlinson
(as I was informed at the British Museum), similar dogs are still imported
into this same country. I have looked through the magnificent works of
Lepsius and Rosellini, and on the Egyptian monuments from the fourth to the
twelfth dynasties (i.e. from about 3400 B.C. to 2100 B.C.) several
varieties of the dog are represented; most of them are allied to
greyhounds; at the later of these periods a dog resembling a hound is
figured, with drooping ears, but with a longer back and more pointed head
than in our hounds. There is, also, a turnspit, with short and crooked
legs, closely resembling the existing variety; but this kind of monstrosity
is so common with various animals, as with the ancon sheep, and even,
according to Rengger, with jaguars in Paraguay, that it would be rash to
look at the monumental animal as the parent of all our turnspits: Colonel
Sykes (1/5. 'Proc. Zoolog. Soc.' July 12, 1831.) also has described an
Indian pariah dog as presenting the same monstrous character. The most
ancient dog represented on the Egyptian monuments is one of the most
singular; it resembles a greyhound, but has long pointed ears and a short
curled tail: a closely allied variety still exists in Northern Africa; for
Mr. E. Vernon Harcourt (1/6. 'Sporting in Algeria' page 51.) states that
the Arab boar-hound is "an eccentric hieroglyphic animal, such as Cheops
once hunted with, somewhat resembling the rough Scotch deer-hound; their
tails are curled tight round on their backs, and their ears stick out at
right angles." With this most ancient variety a pariah-like dog coexisted.
We thus see that, at a period between four and five thousand years ago,
various breeds, viz. pariah dogs, greyhounds, common hounds, mastiffs,
house-dogs, lapdogs, and turnspits, existed, more or less closely
resembling our present breeds. But there is not sufficient evidence that
any of these ancient dogs belonged to the same identical sub-varieties with
our present dogs. (1/7. Berjeau gives facsimiles of the Egyptian drawings.
Mr. C.L. Martin in his 'History of the Dog' 1845 copies several figures
from the Egyptian monuments, and speaks with much confidence with respect
to their identity with still living dogs. Messrs. Nott and Gliddon ('Types
of Mankind' 1854 page 388) give still more numerous figures. Mr. Gliddon
asserts that a curl-tailed greyhound, like that represented on the most
ancient monuments, is common in Borneo; but the Rajah, Sir J. Brooke,
informs me that no such dog exists there.) As long as man was believed to
have existed on this earth only about 6000 years, this fact of the great
diversity of the breeds at so early a period was an argument of much weight
that they had proceeded from several wild sources, for there would not have
been sufficient time for their divergence and modification. But now that we
know, from the discovery of flint tools embedded with the remains of
extinct animals in districts which have since undergone great geographical
changes, that man has existed for an incomparably longer period, and
bearing in mind that the most barbarous nations possess domestic dogs, the
argument from insufficient time falls away greatly in value.
Long before the period of any historical record the dog was domesticated in
Europe. In the Danish Middens of the Neolithic or Newer Stone period, bones
of a canine animal are embedded, and Steenstrup ingeniously argues that
these belonged to a domestic dog; for a very large proportion of the bones
of birds preserved in the refuse consists of long bones, which it was found
on trial dogs cannot devour. (1/8. These, and the following facts on the
Danish remains, are taken from M. Morlot's most interesting memoir in 'Soc.
Vaudoise des Sc. Nat.' tome 6 1860 pages 281, 299, 320.) This ancient dog
was succeeded in Denmark during the Bronze period by a larger kind,
presenting certain differences, and this again during the Iron period, by a
still larger kind. In Switzerland, we hear from Prof. Rutimeyer (1/9. 'Die
Fauna der Pfahlbauten' 1861 s. 117, 162.), that during the Neolithic period
a domesticated dog of middle size existed, which in its skull was about
equally remote from the wolf and jackal, and partook of the characters of
our hounds and setters or spaniels (Jagdhund und Wachtelhund). Rutimeyer
insists strongly on the constancy of form during a very long period of time
of this the most ancient known dog. During the Bronze period a larger dog
appeared, and this closely resembled in its jaw a dog of the same age in
Denmark. Remains of two notably distinct varieties of the dog were found by
Schmerling in a cave (1/10. De Blainville 'Osteographie, Canidae.'); but
their age cannot be positively determined.
The existence of a single race, remarkably constant in form during the
whole Neolithic period, is an interesting fact in contrast with what we see
of the changes which the races underwent during the period of the
successive Egyptian monuments, and in contrast with our existing dogs. The
character of this animal during the Neolithic period, as given by
Rutimeyer, supports De Blainville's view that our varieties have descended
from an unknown and extinct form. But we should not forget that we know
nothing with respect to the antiquity of man in the warmer parts of the
world. The succession of the different kinds of dogs in Switzerland and
Denmark is thought to be due to the immigration of conquering tribes
bringing with them their dogs; and this view accords with the belief that
different wild canine animals were domesticated in different regions.
Independently of the immigration of new races of man, we know from the
wide-spread presence of bronze, composed of an alloy of tin, how much
commerce there must have been throughout Europe at an extremely remote
period, and dogs would then probably have been bartered. At the present
time, amongst the savages of the interior of Guiana, the Taruma Indians are
considered the best trainers of dogs, and possess a large breed which they
barter at a high price with other tribes. (1/11. Sir R. Schomburgk has
given me information on this head. See also 'Journal of R. Geographical
Soc.' volume 13 1843 page 65.)
The main argument in favour of the several breeds of the dog being the
descendants of distinct wild stocks, is their resemblance in various
countries to distinct species still existing there. It must, however, be
admitted that the comparison between the wild and domesticated animal has
been made but in few cases with sufficient exactness. Before entering on
details, it will be well to show that there is no a priori difficulty in
the belief that several canine species have been domesticated. Members of
the dog family inhabit nearly the whole world; and several species agree
pretty closely in habits and structure with our several domesticated dogs.
Mr. Galton has shown (1/12. 'Domestication of Animals' Ethnological Soc.
December 22, 1863.) how fond savages are of keeping and taming animals of
all kinds. Social animals are the most easily subjugated by man, and
several species of Canidae hunt in packs. It deserves notice, as bearing on
other animals as well as on the dog, that at an extremely ancient period,
when man first entered any country, the animals living there would have
felt no instinctive or inherited fear of him, and would consequently have
been tamed far more easily than at present. For instance, when the Falkland
Islands were first visited by man, the large wolf-like dog (Canis
antarcticus) fearlessly came to meet Byron's sailors, who, mistaking this
ignorant curiosity for ferocity, ran into the water to avoid them: even
recently a man, by holding a piece of meat in one hand and a knife in the
other, could sometimes stick them at night. On a island in the Sea of Aral,
when first discovered by Butakoff, the saigak antelopes, which are
"generally very timid and watchful, did not fly from us, but on the
contrary looked at us with a sort of curiosity." So, again, on the shores
of the Mauritius, the manatee was not at first in the least afraid of man,
and thus it has been in several quarters of the world with seals and the
morse. I have elsewhere shown (1/13. 'Journal of Researches' etc. 1845 page
393. With respect to Canis antarcticus, see page 193. For the case of the
antelope, see 'Journal Royal Geographical Soc.' volume 23 page 94.) how
slowly the native birds of several islands have acquired and inherited a
salutary dread of man: at the Galapagos Archipelago I pushed with the
muzzle of my gun hawks from a branch, and held out a pitcher of water for
other birds to alight on and drink. Quadrupeds and birds which have seldom
been disturbed by man, dread him no more than do our English birds, the
cows, or horses grazing in the fields.
It is a more important consideration that several canine species evince (as
will be shown in a future chapter) no strong repugnance or inability to
breed under confinement; and the incapacity to breed under confinement is
one of the commonest bars to domestication. Lastly, savages set the highest
value, as we shall see in the chapter on Selection, on dogs: even half-
tamed animals are highly useful to them: the Indians of North America cross
their half-wild dogs with wolves, and thus render them even wilder than
before, but bolder: the savages of Guiana catch and partially tame and use
the whelps of two wild species of Canis, as do the savages of Australia
those of the wild Dingo. Mr. Philip King informs me that he once trained a
wild Dingo puppy to drive cattle, and found it very useful. From these
several considerations we see that there is no difficulty in believing that
man might have domesticated various canine species in different countries.
It would indeed have been a strange fact if one species alone had been
domesticated throughout the world.
We will now enter into details. The accurate and sagacious Richardson says,
"The resemblance between the Northern American wolves (Canis lupus, var.
occidentalis) and the domestic dogs of the Indians is so great that the
size and strength of the wolf seems to be the only difference. I have more
than once mistaken a band of wolves for the dogs of a party of Indians; and
the howl of the animals of both species is prolonged so exactly in the same
key that even the practised ear of the Indian fails at times to
discriminate them.' He adds that the more northern Esquimaux dogs are not
only extremely like the grey wolves of the Arctic circle in form and
colour, but also nearly equal them in size. Dr. Kane has often seen in his
teams of sledge-dogs the oblique eye (a character on which some naturalists
lay great stress), the drooping tail, and scared look of the wolf. In
disposition the Esquimaux dogs differ little from wolves, and, according to
Dr. Hayes, they are capable of no attachment to man, and are so savage that
when hungry they will attack even their masters. According to Kane they
readily become feral. Their affinity is so close with wolves that they
frequently cross with them, and the Indians take the whelps of wolves "to
improve the breed of their dogs." The half-bred wolves sometimes (Lamare-
Picquot) cannot be tamed, "though this case is rare;" but they do not
become thoroughly well broken in till the second or third generation. These
facts show that there can be but little, if any, sterility between the
Esquimaux dog and the wolf, for otherwise they would not be used to improve
the breed. As Dr. Hayes says of these dogs, "reclaimed wolves they
doubtless are." (1/14. The authorities for the foregoing statements are as
follow:--Richardson in 'Fauna Boreali-Americana' 1829 pages 64, 75; Dr.
Kane 'Arctic Explorations' 1856 volume 1 pages 398, 455; Dr. Hayes 'Arctic
Boat Journey' 1860 page 167. Franklin's 'Narrative' volume 1 page 269,
gives the case of three whelps of a black wolf being carried away by the
Indians. Parry, Richardson, and others, give accounts of wolves and dogs
naturally crossing in the eastern parts of North America. Seeman in his
'Voyage of H.M.S. "Herald"' 1853 volume 2 page 26, says the wolf is often
caught by the Esquimaux for the purpose of crossing with their dogs, and
thus adding to their size and strength. M. Lamare-Picquot in 'Bull. de la
Soc. d'Acclimat.' tome 7 1860 page 148, gives a good account of the half-
bred Esquimaux dogs.)
North America is inhabited by a second kind of wolf, the prairie-wolf
(Canis latrans), which is now looked at by all naturalists as specifically
distinct from the common wolf; and is, according to Mr. J.K. Lord, in some
respects intermediate in habits between a wolf and a fox. Sir J.
Richardson, after describing the Hare Indian dog, which differs in many
respects from the Esquimaux dog, says, "It bears the same relation to the
prairie-wolf that the Esquimaux dog does to the great grey wolf." He could,
in fact, detect no marked difference between them; and Messrs. Nott and
Gliddon give additional details showing their close resemblance. The dogs
derived from the above two aboriginal sources cross together and with the
wild wolves, at least with the C. occidentalis, and with European dogs. In
Florida, according to Bartram, the black wolf-dog of the Indians differs in
nothing from the wolves of that country except in barking. (1/15. 'Fauna
Boreali-Americana' 1829 pages 73, 78, 80. Nott and Gliddon, 'Types of
Mankind' page 383. The naturalist and traveller Bartram is quoted by
Hamilton Smith in 'Naturalist Lib.' volume 10 page 156. A Mexican domestic
dog seems also to resemble a wild dog of the same country; but this may be
the prairie-wolf. Another capable judge, Mr. J.K. Lord ('The Naturalist in
Vancouver Island' 1866 volume 2 page 218), says that the Indian dog of the
Spokans, near the Rocky Mountains, "is beyond all question nothing more
than a tamed Cayote or prairie-wolf," or Canis latrans.)
Turning to the southern parts of the new world, Columbus found two kinds of
dogs in the West Indies; and Fernandez (1/16. I quote this from Mr. R.
Hill's excellent account of the Alco or domestic dog of Mexico, in Gosse's
'Naturalist's Sojourn in Jamaica' 1851 page 329.) describes three in
Mexico: some of these native dogs were dumb--that is, did not bark. In
Guiana it has been known since the time of Buffon that the natives cross
their dogs with an aboriginal species, apparently the Canis cancrivorus.
Sir R. Schomburgk, who has so carefully explored these regions, writes to
me, "I have been repeatedly told by the Arawaak Indians, who reside near
the coast, that they cross their dogs with a wild species to improve the
breed, and individual dogs have been shown to me which certainly resembled
the C. cancrivorus much more than the common breed. It is but seldom that
the Indians keep the C. cancrivorus for domestic purposes, nor is the Ai,
another species of wild dog, and which I consider to be identical with the
Dusicyon silvestris of H. Smith, now much used by the Arecunas for the
purpose of hunting. The dogs of the Taruma Indians are quite distinct, and
resemble Buffon's St. Domingo greyhound." It thus appears that the natives
of Guiana have partially domesticated two aboriginal species, and still
cross their dogs with them; these two species belong to a quite different
type from the North American and European wolves. A careful observer,
Rengger (1/17. 'Naturgeschichte der Saugethiere von Paraguay' 1830 s.
151.), gives reasons for believing that a hairless dog was domesticated
when America was first visited by Europeans: some of these dogs in Paraguay
are still dumb, and Tschudi (1/18. Quoted in Humboldt 'Aspects of Nature'
(English translation) volume 1 page 108.) states that they suffer from cold
in the Cordillera. This naked dog is, however quite distinct from that
found preserved in the ancient Peruvian burial-places, and described by
Tschudi, under the name of Canis ingae, as withstanding cold well and as
barking. It is not known whether these two distinct kinds of dog are the
descendants of native species, and it might be argued that when man first
migrated into America he brought with him from the Asiatic continent dogs
which had not learned to bark; but this view does not seem probable, as the
natives along the line of their march from the north reclaimed, as we have
seen, at least two N. American species of Canidae.
Turning to the Old World, some European dogs closely resemble the wolf;
thus the shepherd dog of the plains of Hungary is white or reddish-brown,
has a sharp nose, short, erect ears, shaggy coat, and bushy tail, and so
much resembles a wolf that Mr. Paget, who gives this description, says he
has known a Hungarian mistake a wolf for one of his own dogs. Jeitteles,
also, remarks on the close similarity of the Hungarian dog and wolf.
Shepherd dogs in Italy must anciently have closely resembled wolves, for
Columella (vii. 12) advises that white dogs be kept, adding, "pastor album
probat, ne pro lupo canem feriat." Several accounts have been given of dogs
and wolves crossing naturally; and Pliny asserts that the Gauls tied their
female dogs in the woods that they might cross with wolves. (1/19. Paget
'Travels in Hungary and Transylvania' volume 1 page 501. Jeitteles 'Fauna
Hungariae Superioris' 1862 s. 13. See Pliny 'History of the World' (English
translation) 8th book ch. 40 about the Gauls crossing their dogs. See also
Aristotle 'Hist. Animal.' Lib. 8 c. 28. For good evidence about wolves and
dogs naturally crossing near the Pyrenees, see M. Mauduyt 'Du Loup et de
ses Races' Poitiers, 1851; also Pallas in 'Acta Acad. St. Petersburgh' 1780
part 2 page 94.) The European wolf differs slightly from that of North
America, and has been ranked by many naturalists as a distinct species. The
common wolf of India is also by some esteemed as a third species, and here
again we find a marked resemblance between the pariah dogs of certain
districts of India and the Indian wolf. (1/20. I give this on excellent
authority, namely Mr. Blyth (under the signature of Zoophilus) in the
'Indian Sporting Review' October 1856 page 134. Mr. Blyth states that he
was struck with the resemblance between a brush-tailed race of pariah-dogs,
north-west of Cawnpore, and the Indian wolf. He gives corroborative
evidence with respect to the dogs of the valley of the Nerbudda.)
With respect to Jackals, Isidore Geoffroy Saint-Hilaire (1/21. For numerous
and interesting details on the resemblance of dogs and jackals see Isid.
Geoffroy St.-Hilaire 'Hist. Nat. Gen.' 1860 tome 3 page 101. See also
'Hist. Nat. des Mammiferes' par Prof. Gervais, 1855 tome 2 page 60.) says
that not one constant difference can be pointed out between their structure
and that of the smaller races of dogs. They agree closely in habits:
jackals, when tamed and called by their master, wag their tails, lick his
hands, crouch, and throw themselves on their backs; they smell at the tails
of other dogs, and void their urine sideways; they roll on carrion or on
animals which they have killed; and, lastly, when in high spirits, they run
round in circles or in a figure of eight, with their tails between their
legs. (1/22. Also Guldenstadt 'Nov. Comment. Acad. Petrop.' tome 20 pro
anno 1775 page 449. Also Salvin in 'Land and Water' October 1869.) A number
of excellent naturalists, from the time of Guldenstadt to that of
Ehrenberg, Hemprich, and Cretzschmar, have expressed themselves in the
strongest terms with respect to the resemblance of the half-domestic dogs
of Asia and Egypt to jackals. M. Nordmann, for instance, says, "Les chiens
d'Awhasie ressemblent etonnamment a des chacals." Ehrenberg (1/23. Quoted
by De Blainville in his 'Osteographie, Canidae' pages 79, 98.) asserts that
the domestic dogs of Lower Egypt, and certain mummied dogs, have for their
wild type a species of wolf (C. lupaster) of the country; whereas the
domestic dogs of Nubia and certain other mummied dogs have the closest
relation to a wild species of the same country, viz. C. sabbar, which is
only a form of the common jackal. Pallas asserts that jackals and dogs
sometimes naturally cross in the East; and a case is on record in Algeria.
(1/24. See Pallas in 'Act. Acad. St. Petersburgh' 1780 part 2 page 91. For
Algeria, see Isid. Geoffroy St.-Hilaire 'Hist. Nat. Gen.' tome 3 page 177.
In both countries it is the male jackal which pairs with female domestic
dogs.) The greater number of naturalists divide the jackals of Asia and
Africa into several species, but some few rank them all as one.
I may add that the domestic dogs on the coast of Guinea are fox-like
animals, and are dumb. (1/25. John Barbut 'Description of the Coast of
Guinea in 1746.') On the east coast of Africa, between latitude 4 deg and 6
deg south, and about ten days' journey in the interior, a semi-domestic
dog, as the Rev. S. Erhardt informs me, is kept, which the natives assert
is derived from a similar wild animal. Lichtenstein (1/26. 'Travels in
South Africa' volume 2 page 272.) says that the dogs of the Bosjemans
present a striking resemblance even in colour (excepting the black stripe
down the back) with the C. mesomelas of South Africa. Mr. E. Layard informs
me that he has seen a Caffre dog which closely resembled an Esquimaux dog.
In Australia the Dingo is both domesticated and wild; though this animal
may have been introduced aboriginally by man, yet it must be considered as
almost an endemic form, for its remains have been found in a similar state
of preservation and associated with extinct mammals, so that its
introduction must have been ancient. (1/27. Selwyn, Geology of Victoria;
'Journal of Geolog. Soc.' volume 14 1858 page 536 and volume 16 1860 page
148; and Prof. M'Coy in 'Annals and Mag. of Nat. Hist.' (3rd series) volume
9 1862 page 147. The Dingo differs from the dogs of the central Polynesian
islands. Dieffenbach remarks ('Travels' volume 2 page 45) that the native
New Zealand dog also differs from the Dingo.)
From this resemblance of the half-domesticated dogs in several countries to
the wild species still living there,--from the facility with which they can
often be crossed together,--from even half-tamed animals being so much
valued by savages,--and from the other circumstances previously remarked on
which favour their domestication, it is highly probable that the domestic
dogs of the world are descended from two well-defined species of wolf (viz.
C. lupus and C. latrans), and from two or three other doubtful species
(namely, the European, Indian, and North African wolves); from at least one
or two South American canine species; from several races or species of
jackal; and perhaps from one or more extinct species. Although it is
possible or even probable that domesticated dogs, introduced into any
country and bred there for many generations, might acquire some of the
characters proper to the aboriginal Canidae of the country, we can hardly
thus account for introduced dogs having given rise to two breeds in the
same country, resembling two of its aboriginal species, as in the above-
given cases of Guiana and of North America. (1/28. These latter remarks
afford, I think, a sufficient answer to some criticisms by Mr. Wallace, on
the multiple origin of dogs, given in Lyell's 'Principles of Geology' 1872
volume 2 page 295.)
It cannot be objected to the view of several canine species having been
anciently domesticated, that these animals are tamed with difficulty: facts
have been already given on this head, but I may add that the young of the
Canis primaevus of India were tamed by Mr. Hodgson (1/29. 'Proceedings
Zoological Soc.' 1833 page 112. See also on the taming of the common wolf,
L. Lloyd 'Scandinavian Adventures' 1854 volume 1 page 460. With respect to
the jackal, see Prof. Gervais 'Hist. Nat. Mamm.' tome 2 page 61. With
respect to the aguara of Paraguay see Rengger's work.), and became as
sensible of caresses, and manifested as much intelligence, as any sporting
dog of the same age. There is not much difference, as we have already shown
and shall further see, in habits between the domestic dogs of the North
American Indians and the wolves of that country, or between the Eastern
pariah dogs and jackals, or between the dogs which have run wild in various
countries and the several natural species of the family. The habit of
barking, however, which is almost universal with domesticated dogs, forms
an exception, as it does not characterise a single natural species of the
family, though I am assured that the Canis latrans of North America utters
a noise which closely approaches a bark. But this habit is soon lost by
dogs when they become feral and is soon reacquired when they are again
domesticated. The case of the wild dogs on the island of Juan Fernandez
having become dumb has often been quoted, and there is reason to believe
(130. Roulin, in 'Mem. present. par divers Savans' tome 6 page 341.) that
the dumbness ensued in the course of thirty-three years; on the other hand,
dogs taken from this island by Ulloa slowly reacquired the habit of
barking. The Mackenzie-river dogs, of the Canis latrans type, when brought
to England, never learned to bark properly; but one born in the Zoological
Gardens (1/31. Martin 'History of the Dog' page 14.) "made his voice sound
as loudly as any other dog of the same age and size." According to
Professor Nillson (1/32. Quoted by L. Lloyd in 'Field Sports of North of
Europe' volume 1 page 387.), a wolf-whelp reared by a bitch barks. I.
Geoffroy Saint-Hilaire exhibited a jackal which barked with the same tone
as any common dog. (1/33. Quatrefages 'Soc. d'Acclimat.' May 11, 1863 page
7.) An interesting account has been given by Mr. G. Clarke (1/34. 'Annals
and Mag of Nat. Hist.' volume 15 1845 page 140.) of some dogs run wild on
Juan de Nova, in the Indian Ocean; "they had entirely lost the faculty of
barking; they had no inclination for the company of other dogs, nor did
they acquire their voice" during a captivity of several months. On the
island they "congregate in vast packs, and catch sea-birds with as much
address as foxes could display." The feral dogs of La Plata have not become
dumb; they are of large size, hunt singly or in packs, and burrow holes for
their young. (1/35. Azara 'Voyages dans l'Amer. Merid.' tome 1 page 381;
his account is fully confirmed by Rengger. Quatrefages gives an account of
a bitch brought from Jerusalem to France which burrowed a hole and littered
in it. See 'Discours, Exposition des Races Canines' 1865 page 3.) In these
habits the feral dogs of La Plata resemble wolves and jackals; both of
which hunt either singly or in packs, and burrow holes. (1/36. With respect
to wolves burrowing holes see Richardson 'Fauna Boreali-Americana' page 64;
and Bechstein 'Naturgeschichte Deutschlands' b. 1 s. 617.) These feral dogs
have not become uniform in colour on Juan Fernandez, Juan de Nova, or La
Plata. (1/37. See Poeppig 'Reise in Chile' b. 1 s. 290; Mr. G. Clarke, as
above; and Rengger, s. 155.) In Cuba the feral dogs are described by
Poeppig as nearly all mouse-coloured, with short ears and light-blue eyes.
In St. Domingo, Col. Ham. Smith says (1/38. Dogs, 'Nat. Library' volume 10
page 121; an endemic South American dog seems also to have become feral in
this island. See Gosse 'Jamaica' page 340.) that the feral dogs are very
large, like greyhounds, of a uniform pale blue-ash, with small ears, and
large light-brown eyes. Even the wild Dingo, though so anciently
naturalised in Australia, "varies considerably in colour," as I am informed
by Mr. P.P. King: a half-bred Dingo reared in England (1/39. Low
'Domesticated Animals' page 650.) showed signs of wishing to burrow.
[From the several foregoing facts we see that reversion in the feral state
gives no indication of the colour or size of the aboriginal parent-species.
One fact, however, with respect to the colouring of domestic dogs, I at one
time hoped might have thrown some light on their origin; and it is worth
giving, as showing how colouring follows laws, even in so anciently and
thoroughly domesticated an animal as the dog. Black dogs with tan-coloured
feet, whatever breed they may belong to, almost invariably have a tan-
coloured spot on the upper and inner corners of each eye, and their lips
are generally thus coloured. I have seen only two exceptions to this rule,
namely, in a spaniel and terrier. Dogs of a light-brown colour often have a
lighter, yellowish-brown spot over the eyes; sometimes the spot is white,
and in a mongrel terrier the spot was black. Mr. Waring kindly examined for
me a stud of fifteen greyhounds in Suffolk: eleven of them were black, or
black and white, or brindled, and these had no eye-spots; but three were
red and one slaty-blue, and these four had dark-coloured spots over their
eyes. Although the spots thus sometimes differ in colour, they strongly
tend to be tan-coloured; this is proved by my having seen four spaniels, a
setter, two Yorkshire shepherd dogs, a large mongrel, and some fox-hounds,
coloured black and white, with not a trace of tan-colour, excepting the
spots over the eyes, and sometimes a little on the feet. These latter
cases, and many others, show plainly that the colour of the feet and the
eye-spots are in some way correlated. I have noticed, in various breeds,
every gradation, from the whole face being tan-coloured, to a complete ring
round the eyes, to a minute spot over the inner and upper corners. The
spots occur in various sub-breeds of terriers and spaniels; in setters; in
hounds of various kinds, including the turnspit-like German badger-hound;
in shepherd dogs; in a mongrel, of which neither parent had the spots; in
one pure bulldog, though the spots were in this case almost white; and in
greyhounds,--but true black-and-tan greyhounds are excessively rare;
nevertheless I have been assured by Mr. Warwick, that one ran at the
Caledonian Champion meeting of April 1860, and was "marked precisely like a
black-and-tan terrier." This dog, or another exactly the same colour, ran
at the Scottish National Club on the 21st of March, 1865; and I hear from
Mr. C.M. Browne, that "there was no reason either on the sire or dam side
for the appearance of this unusual colour." Mr. Swinhoe at my request
looked at the dogs in China, at Amoy, and he soon noticed a brown dog with
yellow spots over the eyes. Colonel H. Smith (1/40. 'The Naturalist
Library' Dogs, volume 10 pages 4, 19.) figures the magnificent black
mastiff of Thibet with a tan-coloured stripe over the eyes, feet, and
chaps; and what is more singular, he figures the Alco, or native domestic
dog of Mexico, as black and white, with narrow tan-coloured rings round the
eyes; at the Exhibition of dogs in London, May 1863, a so-called forest dog
from North-West Mexico was shown, which had pale tan-coloured spots over
the eyes. The occurrence of these tan-coloured spots in dogs of such
extremely different breeds, living in various parts of the world, makes the
fact highly remarkable.
We shall hereafter see, especially in the chapter on Pigeons, that coloured
marks are strongly inherited, and that they often aid us in discovering the
primitive forms of our domestic races. Hence, if any wild canine species
had distinctly exhibited the tan-coloured spots over the eyes, it might
have been argued that this was the parent-form of nearly all our domestic
races. But after looking at many coloured plates, and through the whole
collection of skins in the British Museum, I can find no species thus
marked. It is no doubt possible that some extinct species was thus
coloured. On the other hand, in looking at the various species, there seems
to be a tolerably plain correlation between tan-coloured legs and face; and
less frequently between black legs and a black face; and this general rule
of colouring explains to a certain extent the above-given cases of
correlation between the eye-spots and the colour of the feet. Moreover,
some jackals and foxes have a trace of a white ring round their eyes, as in
C. mesomelas, C. aureus, and (judging from Colonel H. Smith's drawing) in
C. alopex, and C. thaleb. Other species have a trace of a black line over
the corners of the eyes, as in C. variegatus, cinereo-variegatus, and
fulvus, and the wild Dingo. Hence I am inclined to conclude that a tendency
for tan-coloured spots to appear over the eyes in the various breeds of
dogs, is analogous to the case observed by Desmarest, namely, that when any
white appears on a dog the tip of the tail is always white, "de maniere a
rappeler la tache terminale de meme couleur, qui caracterise la plupart des
Canides sauvages." (1/41. Quoted by Prof. Gervais 'Hist. Nat. Mamm.' tome 2
page 66.) This rule, however, as I am assured by Mr. Jesse, does not
invariably hold good.]
It has been objected that our domestic dogs cannot be descended from wolves
or jackals, because their periods of gestation are different. The supposed
difference rests on statements made by Buffon, Gilibert, Bechstein, and
others; but these are now known to be erroneous; and the period is found to
agree in the wolf, jackal, and dog, as closely as could be expected, for it
is often in some degree variable. (1/42. J. Hunter shows that the long
period of seventy-three days given by Buffon is easily explained by the
bitch having received the dog many times during a period of sixteen days
('Phil. Transact.' 1787 page 353). Hunter found that the gestation of a
mongrel from wolf and dog ('Phil. Transact.' 1789 page 160) apparently was
sixty-three days, for she received the dog more than once. The period of a
mongrel dog and jackal was fifty-nine days. Fred. Cuvier found the period
of gestation of the wolf to be ('Dict. Class. d'Hist. Nat.' tome 4 page 8)
two months and a few days, which agrees with the dog. Isid G. St.-Hilaire,
who has discussed the whole subject, and from whom I quote Bellingeri,
states ('Hist. Nat. Gen.' tome 3 page 112) that in the Jardin des Plantes
the period of the jackal has been found to be from sixty to sixty-three
days, exactly as with the dog.) Tessier, who has closely attended to this
subject, allows a difference of four days in the gestation of the dog. The
Rev. W.D. Fox has given me three carefully recorded cases of retrievers, in
which the bitch was put only once to the dog; and not counting this day,
but counting that of parturition, the periods were fifty-nine, sixty-two,
and sixty-seven days. The average period is sixty-three days; but
Bellingeri states that this applies only to large dogs; and that for small
races it is from sixty to sixty-three days; Mr. Eyton of Eyton, who has had
much experience with dogs, also informs me that the time is apt to be
longer with large than with small dogs.
F. Cuvier has objected that the jackal would not have been domesticated on
account of its offensive smell; but savages are not sensitive in this
respect. The degree of odour, also, differs in the different kinds of
jackal (1/43. See Isid. Geoffroy St.-Hilaire 'Hist. Nat. Gen.' tome 3 page
112, on the odour of jackals. Col. Ham. Smith in 'Nat. Lib.' volume 10 page
289.); and Colonel H. Smith makes a sectional division of the group with
one character dependent on not being offensive. On the other hand, dogs--
for instance, rough and smooth terriers--differ much in this respect; and
M. Godron states that the hairless so-called Turkish dog is more
odoriferous than other dogs. Isidore Geoffroy (1/44. Quoted by Quatrefages
in 'Bull. Soc. d'Acclimat.' May 11, 1863.) gave to a dog the same odour as
that from a jackal by feeding it on raw flesh.
The belief that our dogs are descended from wolves, jackals, South American
Canidae, and other species, suggests a far more important difficulty. These
animals in their undomesticated state, judging from a widely-spread
analogy, would have been in some degree sterile if intercrossed; and such
sterility will be admitted as almost certain by all those who believe that
the lessened fertility of crossed forms is an infallible criterion of
specific distinctness. Anyhow these animals keep distinct in the countries
which they inhabit in common. On the other hand, all domestic dogs, which
are here supposed to be descended from several distinct species, are, as
far as is known, mutually fertile together. But, as Broca has well remarked
(1/45. 'Journal de la Physiologie' tome 2 page 385.), the fertility of
successive generations of mongrel dogs has never been scrutinised with that
care which is thought indispensable when species are crossed. The few facts
leading to the conclusion that the sexual feelings and reproductive powers
differ in the several races of the dog when crossed are (passing over mere
size as rendering propagation difficult) as follows: the Mexican Alco
(1/46. See Mr. R. Hill's excellent account of this breed in Gosse's
'Jamaica' page 338; Rengger 'Saugethiere von Paraguay' s. 153. With respect
to Spitz dogs, see Bechstein's 'Naturgesch. Deutschlands' 1801 b. 1 s. 638.
With respect to Dr. Hodgkin's statement made before Brit. Assoc. see 'The
Zoologist' volume 4 for 1845-46 page 1097.) apparently dislikes dogs of
other kinds, but this perhaps is not strictly a sexual feeling; the
hairless endemic dog of Paraguay, according to Rengger, mixes less with the
European races than these do with each other; the Spitz dog in Germany is
said to receive the fox more readily than do other breeds; and Dr. Hodgkin
states that a female Dingo in England attracted the male wild foxes. If
these latter statements can be trusted, they prove some degree of sexual
difference in the breeds of the dog. But the fact remains that our domestic
dogs, differing so widely as they do in external structure, are far more
fertile together than we have reason to believe their supposed wild parents
would have been. Pallas assumes (1/47. 'Acta Acad. St. Petersburgh' 1780
part 2 pages 84, 100.) that a long course of domestication eliminates that
sterility which the parent-species would have exhibited if only lately
captured; no distinct facts are recorded in support of this hypothesis; but
the evidence seems to me so strong (independently of the evidence derived
from other domesticated animals) in favour of our domestic dogs having
descended from several wild stocks, that I am inclined to admit the truth
of this hypothesis.
There is another and closely allied difficulty consequent on the doctrine
of the descent of our domestic dogs from several wild species, namely, that
they do not seem to be perfectly fertile with their supposed parents. But
the experiment has not been quite fairly tried; the Hungarian dog, for
instance, which in external appearance so closely resembles the European
wolf, ought to be crossed with this wolf: and the pariah dogs of India with
Indian wolves and jackals; and so in other cases. That the sterility is
very slight between certain dogs and wolves and other Canidae is shown by
savages taking the trouble to cross them. Buffon got four successive
generations from the wolf and dog, and the mongrels were perfectly fertile
together. (1/48. M. Broca has shown ('Journal de Physiologie' tome 2 page
353) that Buffon's experiments have been often misrepresented. Broca has
collected (pages 390-395) many facts on the fertility of crossed dogs,
wolves, and jackals.) But more lately M. Flourens states positively as the
result of his numerous experiments that hybrids from the wolf and dog,
crossed inter se, become sterile at the third generation, and those from
the jackal and dog at the fourth generation. (1/49. 'De la Longevite
Humaine' par M. Flourens 1855 page 143. Mr. Blyth says ('Indian Sporting
Review' volume 2 page 137) that he has seen in India several hybrids from
the pariah-dog and jackal; and between one of these hybrids and a terrier.
The experiments of Hunter on the jackal are well-known. See also Isid.
Geoffroy St.-Hilaire, 'Hist. Nat. Gen.' tome 3 page 217, who speaks of the
hybrid offspring of the jackal as perfectly fertile for three generations.)
But these animals were closely confined; and many wild animals, as we shall
see in a future chapter, are rendered by confinement in some degree or even
utterly sterile. The Dingo, which breeds freely in Australia with our
imported dogs, would not breed though repeatedly crossed in the Jardin des
Plantes. (1/50. On authority of F. Cuvier quoted in Bronn's 'Geschichte der
Natur' b. 2 s. 164.) Some hounds from Central Africa, brought home by Major
Denham, never bred in the Town of London (1/51. W.C.L. Martin 'History of
the Dog' 1845 page 203. Mr. Philip P. King, after ample opportunities of
observation, informs me that the Dingo and European dogs often cross in
Australia.); and a similar tendency to sterility might be transmitted to
the hybrid offspring of a wild animal. Moreover, it appears that in M.
Flourens' experiments the hybrids were closely bred in and in for three or
four generations; and this circumstance would most certainly increase the
tendency to sterility. Several years ago I saw confined in the Zoological
Gardens of London a female hybrid from an English dog and jackal, which
even in this the first generation was so sterile that, as I was assured by
her keeper, she did not fully exhibit her proper periods; but this case was
certainly exceptional, as numerous instances have occurred of fertile
hybrids from these two animals. In almost all experiments on the crossing
of animals there are so many causes of doubt, that it is extremely
difficult to come to any positive conclusion. It would, however, appear,
that those who believe that our dogs are descended from several species
will have not only to admit that their offspring after a long course of
domestication generally lose all tendency to sterility when crossed
together; but that between certain breeds of dogs and some of their
supposed aboriginal parents a certain degree of sterility has been retained
or possibly even acquired.
Notwithstanding the difficulties in regard to fertility given in the last
two paragraphs, when we reflect on the inherent improbability of man having
domesticated throughout the world one single species alone of so widely
distributed, so easily tamed, and so useful a group as the Canidae; when we
reflect on the extreme antiquity of the different breeds; and especially
when we reflect on the close similarity, both in external structure and
habits, between the domestic dogs of various countries and the wild species
still inhabiting these same countries, the balance of evidence is strongly
in favour of the multiple origin of our dogs.
DIFFERENCES BETWEEN THE SEVERAL BREEDS OF THE DOG.
If the several breeds have descended from several wild stocks, their
difference can obviously in part be explained by that of their parent
species. For instance, the form of the greyhound may be partly accounted
for by descent from some such animal as the slim Abyssinian Canis simensis
(1/52. Ruppel 'Neue Wirbelthiere von Abyssinien' 1835-40 'Mammif.' s. 39
pl. 14. There is a specimen of this fine animal in the British Museum.),
with its elongated muzzle; that of the larger dogs from the larger wolves,
and the smaller and slighter dogs from the jackals: and thus perhaps we may
account for certain constitutional and climatal differences. But it would
be a great error to suppose that there has not been in addition (1/53. Even
Pallas admits this; see 'Act. Acad. St. Petersburgh' 1780 page 93.) a large
amount of variation. The intercrossing of the several aboriginal wild
stocks, and of the subsequently formed races, has probably increased the
total number of breeds, and, as we shall presently see, has greatly
modified some of them. But we cannot explain by crossing the origin of such
extreme forms as thoroughbred greyhounds, bloodhounds, bulldogs, Blenheim
spaniels, terriers, pugs, etc., unless we believe that forms equally or
more strongly characterised in these different respects once existed in
nature. But hardly any one has been bold enough to suppose that such
unnatural forms ever did or could exist in a wild state. When compared with
all known members of the family of Canidae they betray a distinct and
abnormal origin. No instance is on record of such dogs as bloodhounds,
spaniels, true greyhounds having been kept by savages: they are the product
of long-continued civilisation.
[The number of breeds and sub-breeds of the dog is great; Youatt for
instance, describes twelve kinds of greyhounds. I will not attempt to
enumerate or describe the varieties, for we cannot discriminate how much of
their difference is due to variation, and how much to descent from
different aboriginal stocks. But it may be worth while briefly to mention
some points. Commencing with the skull, Cuvier has admitted (1/54. Quoted
by I. Geoffroy 'Hist. Nat. Gen.' tome 3 page 453.) that in form the
differences are "plus fortes que celles d'aucunes especes sauvages d'un
meme genre naturel." The proportions of the different bones; the curvature
of the lower jaw, the position of the condyles with respect to the plane of
the teeth (on which F. Cuvier founded his classification), and in mastiffs
the shape of its posterior branch; the shape of the zygomatic arch, and of
the temporal fossae; the position of the occiput--all vary considerably.
(1/55. F. Cuvier in 'Annales du Museum' tome 18 page 337; Godron 'De
l'Espece' tome 1 page 342; and Col. H. Smith in 'Nat. Library' volume 9
page 101. See also some observations on the degeneracy of the skull in
certain breeds, by Prof. Bianconi 'La Theorie Darwinienne' 1874 page 279.)
The difference in size between the brains of dogs belonging to large and
small breeds "is something prodigious." "Some dogs' brains are high and
rounded, while others are low, long, and narrow in front." In the latter,
"the olfactory lobes are visible for about half their extent, when the
brain is seen from above, but they are wholly concealed by the hemispheres
in other breeds." (1/56. Dr. Burt Wilder 'American Assoc. Advancement of
Science' 1873 pages 236, 239.) The dog has properly six pairs of molar
teeth in the upper jaw, and seven in the lower; but several naturalists
have seen not rarely an additional pair in the upper jaw (1/57. Isid.
Geoffroy Saint-Hilaire 'Hist. des Anomalies' 1832 tome 1 page 660, Gervais
'Hist. Nat. des Mammiferes' tome 2 1855 page 66. De Blainville
('Osteographie, Canidae' page 137) has also seen an extra molar on both
sides.); and Professor Gervais says that there are dogs "qui ont sept
paires de dents superieures et huit inferieures." De Blainville (1/58.
'Osteographie, Canidae' page 137.) has given full particulars on the
frequency of these deviations in the number of the teeth, and has shown
that it is not always the same tooth which is supernumerary. In short-
muzzled races, according to H. Muller (1/59. Wurzburger 'Medecin.
Zeitschrift' 1860 b. 1 s. 265.), the molar teeth stand obliquely, whilst in
long-muzzled races they are placed longitudinally, with open spaces between
them. The naked, so-called Egyptian or Turkish dog is extremely deficient
in its teeth (1/60. Mr. Yarrell in 'Proc. Zoological Soc.' October 8, 1833.
Mr. Waterhouse showed me a skull of one of these dogs, which had only a
single molar on each side and some imperfect incisors.),--sometimes having
none except one molar on each side; but this, though characteristic of the
breed, must be considered as a monstrosity. M. Girard (1/61. Quoted in 'The
Veterinary' London volume 8 page 415.), who seems to have attended closely
to the subject, says that the period of the appearance of the permanent
teeth differs in different dogs, being earlier in large dogs; thus the
mastiff assumes its adult teeth in four or five months, whilst in the
spaniel the period is sometimes more than seven or eight months. On the
other hand small dogs are mature, and the females have arrived at the best
age for breeding, when one year old, whereas large dogs "are still in their
puppyhood at this time, and take fully twice as long to develop their
proportions." (1/62. This is quoted from Stonehenge, a great authority,
'The Dog' 1867 page 187.)
With respect to minor differences little need be said. Isidore Geoffroy has
shown (1/63. 'Hist. Nat. General' tome 3 page 448.) that in size some dogs
are six times as long (the tail being excluded) as others; and that the
height relatively to the length of the body varies from between one to two,
and one to nearly four. In the Scotch deer-hound there is a striking and
remarkable difference in the size of the male and female. (1/64. W. Scrope
'Art of Deer-Stalking' page 354.) Every one knows how the ears vary in size
in different breeds, and with their great development their muscles become
atrophied. Certain breeds of dogs are described as having a deep furrow
between the nostrils and lips. The caudal vertebrae, according to F.
Cuvier, on whose authority the two last statements rest, vary in number;
and the tail in English cattle and some shepherd dogs is almost absent. The
mammae vary from seven to ten in number; Daubenton, having examined twenty-
one dogs, found eight with five mammae on each side; eight with four on
each side; and the others with an unequal number on the two sides. (1/65.
Quoted by Col. Ham. Smith in 'Nat. Lib.' volume 10 page 79.) Dogs have
properly five toes in front and four behind, but a fifth toe is often
added; and F. Cuvier states that, when a fifth toe is present, a fourth
cuneiform bone is developed; and, in this case, sometimes the great
cuneiform bone is raised, and gives on its inner side a large articular
surface to the astragalus; so that even the relative connection of the
bones, the most constant of all characters, varies. These modifications,
however, in the feet of dogs are not important, because they ought to be
ranked, as De Blainville has shown (1/66. De Blainville 'Osteographie,
Canidae' page 134. F. Cuvier 'Annales du Museum' tome 18 page 342. In
regard to mastiffs, see Col. H. Smith 'Nat. Lib.' volume 10 page 218. For
the Thibet mastiff, see Mr. Hodgson in 'Journal of As. Soc. of Bengal'
volume 1 1832 page 342.) as monstrosities. Nevertheless they are
interesting from being correlated with the size of the body, for they occur
much more frequently with mastiffs and other large breeds than with small
dogs. Closely allied varieties, however, sometimes differ in this respect;
thus Mr. Hodgson states that the black-and-tan Lassa variety of the Thibet
mastiff has the fifth digit, whilst the Mustang sub-variety is not thus
characterised. The extent to which the skin is developed between the toes
varies much; but we shall return to this point. The degree to which the
various breeds differ in the perfection of their senses, dispositions, and
inherited habits is notorious to every one. The breeds present some
constitutional differences: the pulse, says Youatt (1/67. 'The Dog' 1845
page 186. With respect to diseases Youatt asserts (page 167) that the
Italian greyhound is "strongly subject" to polypi in the matrix or vagina.
The spaniel and pug (page 182) are most liable to bronchocele. The
liability to distemper (page 232) is extremely different in different
breeds. On the distemper, see also Col. Hutchinson on 'Dog Breaking' 1850
page 279.) "varies materially according to the breed, as well as to the
size of the animal." Different breeds of dogs are subject in different
degrees to various diseases. They certainly become adapted to different
climates under which they have long existed. It is notorious that most of
our best European breeds deteriorate in India. (1/68. See 'Youatt on the
Dog' page 15; 'The Veterinary' London volume 11 page 235.) The Rev R.
Everest (1/69. 'Journal of As. Soc. of Bengal' volume 3 page 19.) believes
that no one has succeeded in keeping the Newfoundland dog long alive in
India; so it is, according to Lichtenstein (1/70. 'Travels' volume 2 page
15.), even at the Cape of Good Hope. The Thibet mastiff degenerates on the
plains of India, and can live only on the mountains. (1/71. Hodgson in
'Journal of As. Soc. of Bengal' volume 1 page 342.) Lloyd (1/72. 'Field
Sports of the North of Europe' volume 2 page 165.) asserts that our
bloodhounds and bulldogs have been tried, and cannot withstand the cold of
the northern European forests.]
Seeing in how many characters the races of the dog differ from each other,
and remembering Cuvier's admission that their skulls differ more than do
those of the species of any natural genus, and bearing in mind how closely
the bones of wolves, jackals, foxes, and other Canidae agree, it is
remarkable that we meet with the statement, repeated over and over again,
that the races of the dog differ in no important characters. A highly
competent judge, Prof. Gervais (1/73. 'Hist. Nat. des Mammif.' 1855 tome 2
pages 66, 67.), admits "si l'on prenait sans controle les alterations dont
chacun de ces organes est susceptible, on pourrait croire qu'il y a entre
les chiens domestiques des differences plus grandes que celles qui separent
ailleurs les especes, quelquefois meme les genres." Some of the differences
above enumerated are in one respect of comparatively little value, for they
are not characteristic of distinct breeds: no one pretends that such is the
case with the additional molar teeth or with the number of mammae; the
additional digit is generally present with mastiffs, and some of the more
important differences in the skull and lower jaw are more or less
characteristic of various breeds. But we must not forget that the
predominant power of selection has not been applied in any of these cases;
we have variability in important parts, but the differences have not been
fixed by selection. Man cares for the form and fleetness of his greyhounds,
for the size of his mastiffs, and formerly for the strength of the jaw in
his bulldogs, etc.; but he cares nothing about the number of their molar
teeth or mammae or digits; nor do we know that differences in these organs
are correlated with, or owe their development to, differences in other
parts of the body about which man does care. Those who have attended to the
subject of selection will admit that, nature having given variability, man,
if he so chose, could fix five toes to the hinder feet of certain breeds of
dogs, as certainly as to the feet of his Dorking fowls: he could probably
fix, but with much more difficulty, an additional pair of molar teeth in
either jaw, in the same way as he has given additional horns to certain
breeds of sheep; if he wished to produce a toothless breed of dogs, having
the so-called Turkish dog with its imperfect teeth to work on, he could
probably do so, for he has succeeded in making hornless breeds of cattle
and sheep.
With respect to the precise causes and steps by which the several races of
dogs have come to differ so greatly from each other, we are, as in most
other cases, profoundly ignorant. We may attribute part of the difference
in external form and constitution to inheritance from distinct wild stocks,
that is to changes effected under nature before domestication. We must
attribute something to the crossing of the several domestic and natural
races. I shall, however, soon recur to the crossing of races. We have
already seen how often savages cross their dogs with wild native species;
and Pennant gives a curious account (1/74. 'History of Quadrupeds' 1793
volume 1 page 238.) of the manner in which Fochabers, in Scotland, was
stocked "with a multitude of curs of a most wolfish aspect" from a single
hybrid-wolf brought into that district.
It would appear that climate to a certain extent directly modifies the
forms of dogs. We have lately seen that several of our English breeds
cannot live in India, and it is positively asserted that when bred there
for a few generations they degenerate not only in their mental faculties,
but in form. Captain Williamson (1/75. 'Oriental Field Sports' quoted by
Youatt 'The Dog' page 15.), who carefully attended to this subject, states
that "hounds are the most rapid in their decline;" "greyhounds and
pointers, also, rapidly decline." But spaniels, after eight or nine
generations, and without a cross from Europe, are as good as their
ancestors. Dr. Falconer informs me that bulldogs, which have been known,
when first brought into the country, to pin down even an elephant by its
trunk, not only fall off after two or three generations in pluck and
ferocity, but lose the under-hung character of their lower jaws; their
muzzles become finer and their bodies lighter. English dogs imported into
India are so valuable that probably due care has been taken to prevent
their crossing with native dogs; so that the deterioration cannot be thus
accounted for. The Rev. R. Everest informs me that he obtained a pair of
setters, born in India, which perfectly resembled their Scotch parents: he
raised several litters from them in Delhi, taking the most stringent
precautions to prevent a cross, but he never succeeded, though this was
only the second generation in India, in obtaining a single young dog like
its parents in size or make; their nostrils were more contracted, their
noses more pointed, their size inferior, and their limbs more slender. So
again on the coast of Guinea, dogs, according to Bosman, "alter strangely;
their ears grow long and stiff like those of foxes, to which colour they
also incline, so that in three or four years, they degenerate into very
ugly creatures; and in three or four broods their barking turns into a
howl." (1/76. A. Murray gives this passage in his 'Geographical
Distribution of Mammals' 4to 1866 page 8.) This remarkable tendency to
rapid deterioration in European dogs subjected to the climate of India and
Africa, may be largely accounted for by reversion to a primordial condition
which many animals exhibit, as we shall hereafter see, when their
constitutions are in any way disturbed.
Some of the peculiarities characteristic of the several breeds of the dog
have probably arisen suddenly, and, though strictly inherited, may be
called monstrosities; for instance, the shape of the legs and body in the
turnspit of Europe and India; the shape of the head and the under-hanging
jaw in the bull-and pug-dog, so alike in this one respect and so unlike in
all others. A peculiarity suddenly arising, and therefore in one sense
deserving to be called a monstrosity, may, however, be increased and fixed
by man's selection. We can hardly doubt that long-continued training, as
with the greyhound in coursing hares, as with water-dogs in swimming--and
the want of exercise, in the case of lapdogs--must have produced some
direct effect on their structure and instincts. But we shall immediately
see that the most potent cause of change has probably been the selection,
both methodical and unconscious, of slight individual differences,--the
latter kind of selection resulting from the occasional preservation, during
hundreds of generations, of those individual dogs which were the most
useful to man for certain purposes and under certain conditions of life. In
a future chapter on Selection I shall show that even barbarians attend
closely to the qualities of their dogs. This unconscious selection by man
would be aided by a kind of natural selection; for the dogs of savages have
partly to gain their own subsistence: for instance, in Australia, as we
hear from Mr. Nind (1/77. Quoted by Mr. Galton 'Domestication of Animals'
page 13.), the dogs are sometimes compelled by want to leave their masters
and provide for themselves; but in a few days they generally return. And we
may infer that dogs of different shapes, sizes, and habits, would have the
best chance of surviving under different circumstances,--on open sterile
plains, where they have to run down their own prey,--on rocky coasts, where
they have to feed on crabs and fish left in the tidal pools, as in the case
of New Guinea and Tierra del Fuego. In this latter country, as I am
informed by Mr. Bridges, the Catechist to the Mission, the dogs turn over
the stones on the shore to catch the crustaceans which lie beneath, and
they "are clever enough to knock off the shell-fish at a first blow;" for
if this be not done, shell-fish are well-known to have an almost invincible
power of adhesion.
It has already been remarked that dogs differ in the degree to which their
feet are webbed. In dogs of the Newfoundland breed, which are eminently
aquatic in their habits, the skin, according to Isidore Geoffroy (1/78.
'Hist. Nat. Gen.' tome 3 page 450.), extends to the third phalanges whilst
in ordinary dogs it extends only to the second. In two Newfoundland dogs
which I examined, when the toes were stretched apart and viewed on the
under side, the skin extended in a nearly straight line between the outer
margins of the balls of the toes; whereas, in two terriers of distinct sub-
breeds, the skin viewed in the same manner was deeply scooped out. In
Canada there is a dog which is peculiar to the country and common there,
and this has "half-webbed feet and is fond of the water." (1/79. Mr.
Greenhow on the Canadian Dog in Loudon's 'Mag. of Nat. Hist.' volume 6 1833
page 511.) English otter-hounds are said to have webbed feet: a friend
examined for me the feet of two, in comparison with the feet of some
harriers and bloodhounds; he found the skin variable in extent in all, but
more developed in the otter-hounds than in the others. (1/80. See Mr. C.O.
Groom-Napier on the webbing of the hind feet of Otterhounds in 'Land and
Water' October 13, 1866 page 270.) As aquatic animals which belong to quite
different orders have webbed feet, there can be no doubt that this
structure would be serviceable to dogs that frequent the water. We may
confidently infer that no man ever selected his water-dogs by the extent to
which the skin was developed between their toes; but what he does, is to
preserve and breed from those individuals which hunt best in the water, or
best retrieve wounded game, and thus he unconsciously selects dogs with
feet slightly better webbed. The effects of use from the frequent
stretching apart of the toes will likewise aid in the result. Man thus
closely imitates Natural Selection. We have an excellent illustration of
this same process in North America, where, according to Sir J. Richardson
(1/81. 'Fauna Boreali-Americana' 1829 page 62.), all the wolves, foxes, and
aboriginal domestic dogs have their feet broader than in the corresponding
species of the Old World, and "well calculated for running on the snow."
Now, in these Arctic regions, the life or death of every animal will often
depend on its success in hunting over the snow when soft; and this will in
part depend on the feet being broad; yet they must not be so broad as to
interfere with the activity of the animal when the ground is sticky, or
with its power of burrowing holes, or with other necessary habits of life.
As changes in domestic breeds which take place so slowly are not to be
noticed at any one period, whether due to the selection of individual
variations or of differences resulting from crosses, are most important in
understanding the origin of our domestic productions, and likewise in
throwing indirect light on the changes effected under nature, I will give
in detail such cases as I have been able to collect. Lawrence (1/82. 'The
Horse in all his Varieties, etc.' 1829 pages 230, 234.), who paid
particular attention to the history of the foxhound, writing in 1829, says
that between eighty and ninety years before "an entirely new foxhound was
raised through the breeder's art," the ears of the old southern hound being
reduced, the bone and bulk lightened, the waist increased in length, and
the stature somewhat added to. It is believed that this was effected by a
cross with a greyhound. With respect to this latter dog, Youatt (1/83. 'The
Dog' 1845 pages 31, 35; with respect to King Charles' spaniel page 45; for
the setter page 90.), who is generally cautious in his statements, says
that the greyhound within the last fifty years, that is before the
commencement of the present century, "assumed a somewhat different
character from that which he once possessed. He is now distinguished by a
beautiful symmetry of form, of which he could not once boast, and he has
even superior speed to that which he formerly exhibited. He is no longer
used to struggle with deer, but contends with his fellows over a shorter
and speedier course." An able writer (1/84. In the 'Encyclop. of Rural
Sports' page 557.) believes that our English greyhounds are the
descendants, PROGRESSIVELY IMPROVED, of the large rough greyhounds which
existed in Scotland so early as the third century. A cross at some former
period with the Italian greyhound has been suspected; but this seems hardly
probable, considering the feebleness of this latter breed. Lord Orford, as
is well-known, crossed his famous greyhounds, which failed in courage, with
a bulldog--this breed being chosen from being erroneously supposed to be
deficient in the power of scent; "after the sixth or seventh generation,"
says Youatt, "there was not a vestige left of the form of the bulldog, but
his courage and indomitable perseverance remained."
Youatt infers, from a comparison of an old picture of King Charles's
spaniels with the living dog, that "the breed of the present day is
materially altered for the worse:" the muzzle has become shorter, the
forehead more prominent, and the eyes larger; the changes in this case have
probably been due to simple selection. The setter, as this author remarks
in another place, "is evidently the large spaniel improved to his present
peculiar size and beauty, and taught another way of marking his game. If
the form of the dog were not sufficiently satisfactory on this point, we
might have recourse to history:" he then refers to a document dated 1685
bearing on this subject, and adds that the pure Irish setter shows no signs
of a cross with the pointer, which some authors suspect has been the case
with the English setter. The bulldog is an English breed, and as I hear
from Mr. G.R. Jesse (1/85. Author of 'Researches into the History of the
British Dog.), seems to have originated from the mastiff since the time of
Shakspeare; but certainly existed in 1631, as shown by Prestwick Eaton's
letters. There can be no doubt that the fancy bulldogs of the present day,
now that they are not used for bull-baiting, have become greatly reduced in
size, without any express intention on the part of the breeder. Our
pointers are certainly descended from a Spanish breed, as even their
present names, Don, Ponto, Carlos, etc., show; it is said that they were
not known in England before the Revolution in 1688 (1/86. See Col. Hamilton
Smith on the antiquity of the Pointer, in 'Nat. Lib.' volume 10 page 196.);
but the breed since its introduction has been much modified, for Mr.
Borrow, who is a sportsman and knows Spain intimately well, informs me that
he has not seen in that country any breed "corresponding in figure with the
English pointer; but there are genuine pointers near Xeres which have been
imported by English gentlemen." A nearly parallel case is offered by the
Newfoundland dog, which was certainly brought into England from that
country, but which has since been so much modified that, as several writers
have observed, it does not now closely resemble any existing native dog in
Newfoundland. (1/87. The Newfoundland dog is believed to have originated
from a cross between the Esquimaux dog and a large French hound. See Dr.
Hodgkin 'British Assoc.' 1844; Bechstein 'Naturgesch. Deutschland' b. 1 s.
574; 'Nat. Lib.' volume 10 page 132; also Mr. Jukes 'Excursion in and about
Newfoundland.')
These several cases of slow and gradual changes in our English dogs possess
some interest; for though the changes have generally, but not invariably,
been caused by one or two crosses with a distinct breed, yet we may feel
sure, from the well-known extreme variability of crossed breeds, that
rigorous and long-continued selection must have been practised, in order to
improve them in a definite manner. As soon as any strain or family became
slightly improved or better adapted to alter circumstances, it would tend
to supplant the older and less improved strains. For instance, as soon as
the old foxhound was improved by a cross with the greyhound, or by simple
selection, and assumed its present character--and the change was probably
desired owing to the increased fleetness of our hunters--it rapidly spread
throughout the country, and is now everywhere nearly uniform. But the
process of improvement is still going on for every one tries to improve his
strain by occasionally procuring dogs from the best kennels. Through this
process of gradual substitution the old English hound has been lost; and so
it has been with the Irish wolf-dog, the old English bulldog, and several
other breeds, such as the alaunt, as I am informed by Mr. Jesse. But the
extinction of former breeds is apparently aided by another cause; for
whenever a breed is kept in scanty numbers, as at present with the
bloodhound, it is reared with some difficulty, apparently from the evil
effects of long-continued close interbreeding. As several breeds of the dog
have been slightly but sensibly modified within so short a period as the
last one or two centuries, by the selection of the best individuals,
modified in many cases by crosses with other breeds; and as we shall
hereafter see that the breeding of dogs was attended to in ancient times,
as it still is by savages, we may conclude that we have in selection, even
if only occasionally practised, a potent means of modification.
DOMESTIC CATS.
Cats have been domesticated in the East from an ancient period; Mr. Blyth
informs me that they are mentioned in a Sanskrit writing 2000 years old,
and in Egypt their antiquity is known to be even greater, as shown by
monumental drawings and their mummied bodies. These mummies, according to
De Blainville (1/88. De Blainville 'Osteographie, Felis' page 65 on the
character of F. caligulata; pages 85, 89, 90, 175, on the other mummied
species. He quotes Ehrenberg on F. maniculata being mummied.), who has
particularly studied the subject, belong to no less than three species,
namely, F. caligulata, bubastes, and chaus. The two former species are said
to be still found, both wild and domesticated, in parts of Egypt. F.
caligulata presents a difference in the first inferior milk molar tooth, as
compared with the domestic cats of Europe, which makes De Blainville
conclude that it is not one of the parent-forms of our cats. Several
naturalists, as Pallas, Temminck, Blyth, believe that domestic cats are the
descendants of several species commingled: it is certain that cats cross
readily with various wild species, and it would appear that the character
of the domestic breeds has, at least in some cases, been thus affected. Sir
W. Jardine has no doubt that, "in the north of Scotland, there has been
occasional crossing with our native species (F. sylvestris), and that the
result of these crosses has been kept in our houses. I have seen," he adds,
"many cats very closely resembling the wild cat, and one or two that could
scarcely be distinguished from it." Mr. Blyth (1/89. Asiatic Soc. of
Calcutta; Curator's Report, August 1856. The passage from Sir W. Jardine is
quoted from this Report. Mr. Blyth, who has especially attended to the wild
and domestic cats of India, has given in this Report a very interesting
discussion on their origin.) 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
(1/90. 'Fauna Hungariae Sup.' 1862 s. 12.) was assured on trustworthy
authority that a wild male cat crossed with a female domestic cat, and that
the hybrids long lived in a domesticated state. In Algiers the domestic cat
has crossed with the wild cat (F. lybica) of that country. (1/91. Isid.
Geoffroy Saint-Hilaire 'Hist. Nat. Gen.' tome 3 page 177.) In South Africa
as Mr. E. Layard informs me, the domestic cat intermingles freely with the
wild F. caffra; he has seen a pair of hybrids which were quite tame and
particularly attached to the lady who brought them up; and Mr. Fry has
found that these hybrids are fertile. In India the domestic cat, according
to Mr. Blyth, has crossed with four Indian species. With respect to one of
these species, F. chaus, an excellent observer, Sir W. Elliot, informs me
that he once killed, near Madras, a wild brood, which were evidently
hybrids from the domestic cat; these young animals had a thick lynx-like
tail and the broad brown bar on the inside of the forearm characteristic of
F. chaus. Sir W. Elliot adds that he has often observed this same mark on
the forearms of domestic cats in India. Mr. Blyth states that domestic cats
coloured nearly like F. chaus, but not resembling that species in shape,
abound in Bengal; he adds, "such a colouration is utterly unknown in
European cats, and the proper tabby markings (pale streaks on a black
ground, peculiarly and symmetrically disposed), so common in English cats,
are never seen in those of India." Dr. D. Short has assured Mr. Blyth
(1/92. 'Proc. Zoolog. Soc.' 1863 page 184.) that, at Hansi, hybrids between
the common cat and F. ornata (or torquata) occur, "and that many of the
domestic cats of that part of India were undistinguishable from the wild F.
ornata." Azara states, but only on the authority of the inhabitants, that
in Paraguay the cat has crossed with two native species. From these several
cases we see that in Europe, Asia, Africa, and America, the common cat,
which lives a freer life than most other domesticated animals, has crossed
with various wild species; and that in some instances the crossing has been
sufficiently frequent to affect the character of the breed.
Whether domestic cats have descended from several distinct species, or have
only been modified by occasional crosses, their fertility, as far as is
known, is unimpaired. The large Angora or Persian cat is the most distinct
in structure and habits of all the domestic breeds; and is believed by
Pallas, but on no distinct evidence, to be descended from the F. manul of
middle Asia; and I am assured by Mr. Blyth that the Angora cat breeds
freely with Indian cats, which, as we have already seen, have apparently
been much crossed with F. chaus. In England half-bred Angora cats are
perfectly fertile with one another.
Within the same country we do not meet with distinct races of the cat, as
we do of dogs and of most other domestic animals; though the cats of the
same country present a considerable amount of fluctuating variability. The
explanation obviously is that, from their nocturnal and rambling habits,
indiscriminate crossing cannot without much trouble be prevented. Selection
cannot be brought into play to produce distinct breeds, or to keep those
distinct which have been imported from foreign lands. On the other hand, in
islands and in countries completely separated from each other, we meet with
breeds more or less distinct; and these cases are worth giving, showing
that the scarcity of distinct races in the same country is not caused by a
deficiency of variability in the animal. The tailless cats of the Isle of
Man are said to differ from common cats not only in the want of a tail, but
in the greater length of their hind legs, in the size of their heads, and
in habits. The Creole cat of Antigua, as I am informed by Mr. Nicholson, is
smaller, and has a more elongated head, than the British cat. In Ceylon, as
Mr. Thwaites writes to me, every one at first notices the different
appearance of the native cat from the English animal; it is of small size,
with closely lying hairs; its head is small, with a receding forehead; but
the ears are large and sharp; altogether it has what is there called a
"low-caste" appearance. Rengger (1/93. 'Saugethiere von Paraguay' 1830 s.
212.) says that the domestic cat, which has been bred for 300 years in
Paraguay, presents a striking difference from the European cat; it is
smaller by a fourth, has a more lanky body, its hair is short, shining,
scanty and lies close, especially on the tail: he adds that the change has
been less at Ascension, the capital of Paraguay, owing to the continual
crossing with newly imported cats; and this fact well illustrates the
importance of separation. The conditions of life in Paraguay appear not to
be highly favourable to the cat, for, though they have run half-wild, they
do not become thoroughly feral, like so many other European animals. In
another part of South America, according to Roulin (1/94. 'Mem. presentes
par divers Savans: Acad. Roy. des Sciences' tome 6 page 346. Gomara first
noticed this fact in 1554.), the introduced cat has lost the habit of
uttering its hideous nocturnal howl. The Rev. W.D. Fox purchased a cat in
Portsmouth, which he was told came from the coast of Guinea; its skin was
black and wrinkled, fur bluish-grey and short, its ears rather bare, legs
long, and whole aspect peculiar. This "negro" cat was fertile with common
cats. On the opposite coast of Africa, at Mombas, Captain Owen, R.N. (1/95.
'Narrative of Voyages' volume 2 page 180.) states that all the cats are
covered with short stiff hair instead of fur: he gives a curious account of
a cat from Algoa Bay, which had been kept for some time on board and could
be identified with certainty; this animal was left for only eight weeks at
Mombas, but during that short period it "underwent a complete
metamorphosis, having parted with its sandy-coloured fur." A cat from the
Cape of Good Hope has been described by Desmarest as remarkable from a red
stripe extending along the whole length of its back. Throughout an immense
area, namely, the Malayan archipelago, Siam, Pegu, and Burmah, all the cats
have truncated tails about half the proper length (1/96. J. Crawfurd
'Descript. Dict. of the Indian Islands' page 255. The Madagascar cat is
said to have a twisted tail; see Desmarest in 'Encyclop. Nat. Mamm.' 1820
page 233, for some of the other breeds.), 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. (1/97. Admiral Lutke's Voyage volume 3 page 308.) In
China a breed has drooping ears. At Tobolsk, according to Gmelin, there is
a red-coloured breed. In Asia, also, we find the well-known Angora or
Persian breed.
The domestic cat has run wild in several countries, and everywhere assumes,
as far as can be judged by the short recorded descriptions, a uniform
character. Near Maldonado, in La Plata, I shot one which seemed perfectly
wild; it was carefully examined by Mr. Waterhouse (1/98. 'Zoology of the
Voyage of the Beagle, Mammalia' page 20. Dieffenbach 'Travels in New
Zealand' volume 2 page 185. Ch. St. John 'Wild Sports of the Highlands'
1846 page 40.), who found nothing remarkable in it, excepting its great
size. In New Zealand according to Dieffenbach, the feral cats assume a
streaky grey colour like that of wild cats; and this is the case with the
half-wild cats of the Scotch Highlands.
We have seen that distant countries possess distinct domestic races of the
cat. The differences may be in part due to descent from several aboriginal
species, or at least to crosses with them. In some cases, as in Paraguay,
Mombas, and Antigua, the differences seem due to the direct action of
different conditions of life. In other cases some slight effect may
possibly be attributed to natural selection, as cats in many cases have
largely to support themselves and to escape diverse dangers. But man, owing
to the difficulty of pairing cats, has done nothing by methodical
selection; and probably very little by unintentional selection; though in
each litter he generally saves the prettiest, and values most a good breed
of mouse- or rat-catchers. Those cats which have a strong tendency to prowl
after game, generally get destroyed by traps. As cats are so much petted, a
breed bearing the same relation to other cats, that lapdogs bear to larger
dogs, would have been much valued; and if selection could have been
applied, we should certainly have had many breeds in each long-civilised
country, for there is plenty of variability to work upon.
We see in this country considerable diversity in size, some in the
proportions of the body, and extreme variability in colouring. I have only
lately attended to this subject, but have already heard of some singular
cases of variation; one of a cat born in the West Indies toothless, and
remaining so all its life. Mr. Tegetmeier has shown me the skull of a
female cat with its canines so much developed that they protruded uncovered
beyond the lips; the tooth with the fang being .95, and the part projecting
from the gum .6 of an inch in length. I have heard of several families of
six-toed cats, in one of which the peculiarity had been transmitted for at
least three generations. The tail varies greatly in length; I have seen a
cat which always carried its tail flat on its back when pleased. The ears
vary in shape, and certain strains, in England, inherit a pencil-like tuft
of hairs, above a quarter of an inch in length, on the tips of their ears;
and this same peculiarity, according to Mr. Blyth, characterises some cats
in India. The great variability in the length of the tail and the lynx-like
tufts of hairs on the ears are apparently analogous to differences in
certain wild species of the genus. A much more important difference,
according to Daubenton (1/99. Quoted by Isid. Geoffroy 'Hist. Nat. Gen.'
tome 3 page 427.), is that the intestines of domestic cats are wider, and a
third longer, than in wild cats of the same size; and this apparently has
been by their less strictly carnivorous diet.
CHAPTER 1.II.
HORSES AND ASSES.
HORSE.
DIFFERENCES IN THE BREEDS.
INDIVIDUAL VARIABILITY OF.
DIRECT EFFECTS OF THE CONDITIONS OF LIFE.
CAN WITHSTAND MUCH COLD.
BREEDS MUCH MODIFIED BY SELECTION.
COLOURS OF THE HORSE.
DAPPLING.
DARK STRIPES ON THE SPINE, LEGS, SHOULDERS, AND FOREHEAD.
DUN-COLOURED HORSES MOST FREQUENTLY STRIPED.
STRIPES PROBABLY DUE TO REVERSION TO THE PRIMITIVE STATE OF THE HORSE.
ASSES.
BREEDS OF.
COLOUR OF.
LEG- AND SHOULDER-STRIPES.
SHOULDER-STRIPES SOMETIMES ABSENT, SOMETIMES FORKED.
The history of the Horse is lost in antiquity. Remains of this animal in a
domesticated condition have been found in the Swiss lake-dwellings,
belonging to the Neolithic period. (2/1. Rutimeyer 'Fauna der Pfahlbauten'
1861 s. 122.) At the present time the number of breeds is great, as may be
seen by consulting any treatise on the Horse. (2/2. See 'Youatt on the
Horse': J. Lawrence on the Horse 1829; W.C.L. Martin 'History of the Horse'
1845: Col. H. Smith in 'Nat. Library, Horses' 1841 volume 12: Prof. Veith
'Die naturgesch. Haussaugethiere' 1856.) Looking only to the native ponies
of Great Britain, those of the Shetland Isles, Wales, the New Forest, and
Devonshire are distinguishable; and so it is, amongst other instances, with
each separate island in the great Malay archipelago. (2/3. Crawfurd
'Descript. Dict. of Indian Islands' 1856 page 153. "There are many
different breeds, every island having at least one peculiar to it." Thus in
Sumatra there are at least two breeds; in Achin and Batubara one; in Java
several breeds; one in Bali, Lomboc, Sumbawa (one of the best breeds),
Tambora, Bima, Gunung-api, Celebes, Sumba, and Philippines. Other breeds
are specified by Zollinger in the 'Journal of the Indian Archipelago'
volume 5 page 343 etc.) Some of the breeds present great differences in
size, shape of ears, length of mane, proportions of the body, form of the
withers and hind quarters, and especially in the head. Compare the race-
horse, dray-horse, and a Shetland pony in size, configuration, and
disposition; and see how much greater the difference is than between the
seven or eight other living species of the genus Equus.
Of individual variations not known to characterise particular breeds, and
not great or injurious enough to be called monstrosities, I have not
collected many cases. Mr. G. Brown, of the Cirencester Agricultural
College, who has particularly attended to the dentition of our domestic
animals, writes to me that he has "several times noticed eight permanent
incisors instead of six in the jaw." Male horses only should have canines,
but they are occasionally found in the mare, though a small size. (2/4.
'The Horse' etc. by John Lawrence 1829 page 14.) The number of ribs on each
side is properly eighteen, but Youatt (2/5. 'The Veterinary' London volume
5 page 543.) asserts that not unfrequently there are nineteen, the
additional one being always the posterior rib. It is a remarkable fact that
the ancient Indian horse is said in the Rig-Veda to have only seventeen
ribs; and M. Pietrement (2/6. 'Memoire sur les chevaux a trente-quatre
cotes' 1871.), who has called attention to this subject, gives various
reasons for placing full trust in this statement, more especially as during
former times the Hindoos carefully counted the bones of animals. I have
seen several notices of variations in the bones of the leg; thus Mr. Price
(2/7. 'Proc. Veterinary Assoc.' in 'The Veterinary' volume 13 page 42.)
speaks of an additional bone in the hock, and of certain abnormal
appearances between the tibia and astragalus, as quite common in Irish
horses, and not due to disease. Horses have often been observed, according
to M. Gaudry (2/8. 'Bulletin de la Soc. Geolog.' tome 22 1866 page 22.), to
possess a trapezium and a rudiment of a fifth metacarpal bone, so that "one
sees appearing by monstrosity, in the foot of the horse, structures which
normally exist in the foot of the Hipparion,"--an allied and extinct
animal. In various countries horn-like projections have been observed on
the frontal bones of the horse: in one case described by Mr. Percival they
arose about two inches above the orbital processes, and were "very like
those in a calf from five to six months old," being from half to three-
quarters of an inch in length. (2/9. Mr. Percival of the Enniskillen
Dragoons in 'The Veterinary' volume 1 page 224: see Azara, 'Des Quadrupedes
du Paraguay' tome 2 page 313. The French translator of Azara refers to
other cases mentioned by Huzard as having occurred in Spain.) Azara has
described two cases in South America in which the projections were between
three and four inches in length: other instances have occurred in Spain.
That there has been much inherited variation in the horse cannot be
doubted, when we reflect on the number of the breeds existing throughout
the world or even within the same country, and when we know that they have
largely increased in number since the earliest known records. (2/10.
Godron, 'De l'Espece' tome 1 page 378.) Even in so fleeting a character as
colour, Hofacker (2/11. 'Ueber die Eigenschaften' etc. 1828 s. 10.) found
that, out of 216 cases in which horses of the same colour were paired, only
eleven pairs produced foals of a quite different colour. As Professor Low
(2/12. 'Domesticated Animals of the British Islands' pages 527, 532. In all
the veterinary treatises and papers which I have read, the writers insist
in the strongest terms on the inheritance by the horse of all good and bad
tendencies and qualities. Perhaps the principle of inheritance is not
really stronger in the horse than in any other animal; but, from its value,
the tendency has been more carefully observed.) has remarked, the English
race-horse offers the best possible evidence of inheritance. The pedigree
of a race-horse is of more value in judging of its probable success than
its appearance: "King Herod" gained in prizes 201,505 pounds sterling, and
begot 497 winners; "Eclipse" begot 334 winners.
Whether the whole amount of difference between the various breeds has
arisen under domestication is doubtful. From the fertility of the most
distinct breeds (2/13. Andrew Knight crossed breeds so different in size as
a dray-horse and Norwegian pony: see A. Walker on 'Intermarriage' 1838 page
205.) when crossed, naturalists have generally looked at all the breeds as
having descended from a single species. Few will agree with Colonel H.
Smith, who believes that they have descended from no less than five
primitive and differently coloured stocks. (2/14. 'Nat. Library, Horses'
volume 12 page 208.) But as several species and varieties of the horse
existed (2/15. Gervais 'Hist. Nat. Mamm.' tome 2 page 143. Owen 'British
Fossil Mammals' page 383.) during the later tertiary periods, and as
Rutimeyer found differences in the size and form of the skull in the
earliest known domesticated horses (2/16. 'Kenntniss der fossilen Pferde'
1863 s. 131.), we ought not to feel sure that all our breeds are descended
from a single species. The savages of North and South America easily
reclaim the feral horses, so that there is no improbability in savages in
various quarters of the world having domesticated more than one native
species or natural race. M. Sanson (2/17. 'Comptes rendus' 1866 page 485
and 'Journal de l'Anat. et de la Phys.' Mai 1868.) thinks that he has
proved that two distinct species have been domesticated, one in the East,
and one in North Africa; and that these differed in the number of their
lumbar vertebra and in various other parts; but M. Sanson seems to believe
that osteological characters are subject to very little variation, which is
certainly a mistake. At present no aboriginal or truly wild horse is
positively known to exist; for it is commonly believed that the wild horses
of the East are escaped domestic animals. (2/18. Mr. W.C.L. Martin, 'The
Horse' 1845 page 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.) If therefore our domestic breeds are descended from
several species or natural races, all have become extinct in the wild
state.
With respect to the causes of the modifications which horses have
undergone, the conditions of life seem to produce a considerable direct
effect. Mr. D. Forbes, who has had excellent opportunities of comparing the
horses of Spain with those of South America, informs me that the horses of
Chile, which have lived under nearly the same conditions as their
progenitors in Andalusia, remain unaltered, whilst the Pampas horses and
the Puno horses are considerably modified. There can be no doubt that
horses become greatly reduced in size and altered in appearance by living
on mountains and islands; and this apparently is due to want of nutritious
or varied food. Every one knows how small and rugged the ponies are on the
Northern islands and on the mountains of Europe. Corsica and Sardinia have
their native ponies; and there were (2/19. 'Transact. Maryland Academy'
volume 1 part 1 page 28.), or still are, on some islands on the coast of
Virginia, ponies like those of the Shetland Islands, which are believed to
have originated through exposure to unfavourable conditions. The Puno
ponies, which inhabit the lofty regions of the Cordillera, are, as I hear
from Mr. D. Forbes, strange little creatures, very unlike their Spanish
progenitors. Further south, in the Falkland Islands, the offspring of the
horses imported in 1764 have already so much deteriorated in size (2/20.
Mr. Mackinnon 'The Falkland Islands' page 25. The average height of the
Falkland horses is said to be 14 hands 2 inches. See also my 'Journal of
Researches.') and strength that they are unfitted for catching wild cattle
with the lasso; so that fresh horses have to be brought for this purpose
from La Plata at a great expense. The reduced size of the horses bred on
both southern and northern islands, and on several mountain-chains, can
hardly have been caused by the cold, as a similar reduction has occurred on
the Virginian and Mediterranean islands. The horse can withstand intense
cold, for wild troops live on the plains of Siberia under lat. 56 deg,
(2/21. Pallas 'Act. Acad. St. Petersburgh' 1777 part 2 page 265. With
respect to the tarpans scraping away the snow see Col. Hamilton Smith in
'Nat. Lib.' volume 12 page 165.) and aboriginally the horses must have
inhabited countries annually covered with snow, for he long retains the
instinct of scraping it away to get at the herbage beneath. The wild
tarpans in the East have this instinct; and so it is, as I am informed by
Admiral Sulivan, with the horses recently and formerly introduced into the
Falkland Islands from La Plata, some of which have run wild; this latter
fact is remarkable, as the progenitors of these horses could not have
followed this instinct during many generations in La Plata. On the other
hand, the wild cattle of the Falklands never scrape away the snow, and
perish when the ground is long covered. In the northern parts of America
the horses descended from those introduced by the Spanish conquerors of
Mexico, have the same habit, as have the native bisons, but not so the
cattle introduced from Europe. (2/22. Franklin 'Narrative' volume 1 page 87
note by Sir J. Richardson.)
The horse can flourish under intense heat as well as under intense cold,
for he is known to come to the highest perfection, though not attaining a
large size, in Arabia and northern Africa. Much humidity is apparently more
injurious to the horse than heat or cold. In the Falkland Islands, horses
suffer much from the dampness; and this circumstance may perhaps partly
account for the singular fact that to the eastward of the Bay of Bengal
(2/23. Mr. J.H. Moor 'Notices of the Indian Archipelago' Singapore 1837
page 189. A pony from Java was sent ('Athenaeum' 1842 page 718) to the
Queen only 28 inches in height. For the Loo Choo Islands, see Beechey
'Voyage' 4th. edition volume 1 page 499.), 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. (2/24. J. Crawfurd,
'History of the Horse' 'Journal of Royal United Service Institution' volume
4.)
With most of our domesticated animals, some breeds are kept on account of
their curiosity or beauty; but the horse is valued almost solely for its
utility. Hence semi-monstrous breeds are not preserved; and probably all
the existing breeds have been slowly formed either by the direct action of
the conditions of life, or through the selection of individual differences.
No doubt semi-monstrous breeds might have been formed: thus Mr. Waterton
records (2/25. 'Essays on Natural History' 2nd series page 161.) the case
of a mare which produced successively three foals without tails; so that a
tailless race might have been formed like the tailless races of dogs and
cats. A Russian breed of horses is said to have curled hair, and Azara
(2/26. 'Quadrupedes du Paraguay' tome 2 page 333. Dr. Canfield informs me
that a breed with curly hair was formed by selection at Los Angeles in
North America.) relates that in Paraguay horses are occasionally born, but
are generally destroyed, with hair like that on the head of a negro; and
this peculiarity is transmitted even to half-breeds: it is a curious case
of correlation that such horses have short manes and tails, and their hoofs
are of a peculiar shape like those of a mule.
It is scarcely possible to doubt that the long-continued selection of
qualities serviceable to man has been the chief agent in the formation of
the several breeds of the horse. Look at a dray-horse, and see how well
adapted he is to draw heavy weights, and how unlike in appearance to any
allied wild animal. The English race-horse is known to be derived from the
commingled blood of Arabs, Turks, and Barbs; but selection, which was
carried on during very early times in England (2/27. See the evidence on
this head in 'Land and Water' May 2, 1868.), together with training, have
made him a very different animal from his parent-stocks. As a writer in
India, who evidently knows the pure Arab well, asks, who now, "looking at
our present breed of race-horses, could have conceived that they were the
result of the union of the Arab horse and African mare?" The improvement is
so marked that in running for the Goodwood Cup "the first descendants of
Arabian, Turkish, and Persian horses, are allowed a discount of 18 pounds
weight; and when both parents are of these countries a discount of 36
pounds (2/28. Prof. Low 'Domesticated Animals' page 546. With respect to
the writer in India see 'India Sporting Review' volume 2 page 181. As
Lawrence has remarked ('The Horse' page 9), "perhaps no instance has ever
occurred of a three-part bred horse (i.e. a horse, one of whose
grandparents was of impure blood) saving his distance in running two miles
with thoroughbred racers." Some few instances are on record of seven-eights
racers having been successful.) It is notorious that the Arabs have long
been as careful about the pedigree of their horses as we are, and this
implies great and continued care in breeding. Seeing what has been done in
England by careful breeding, can we doubt that the Arabs must likewise have
produced during the course of centuries a marked effect on the qualities of
their horses? But we may go much farther back in time, for in the Bible we
hear of studs carefully kept for breeding, and of horses imported at high
prices from various countries. (2/29. Prof. Gervais 'Hist. Nat. Mamm.' tome
2 page 144 has collected many facts on this head. For instance Solomon (1
Kings x. 28) bought horses in Egypt at a high price.) We may therefore
conclude that, whether or not the various existing breeds of the horse have
proceeded from one or more aboriginal stocks, yet that a great amount of
change has resulted from the direct action of the conditions of life, and
probably a still greater amount from the long-continued selection by man of
slight individual differences.
With several domesticated quadrupeds and birds, certain coloured marks are
either strongly inherited or tend to reappear after having been lost for a
long time. As this subject will hereafter be seen to be of importance, I
will give a full account of the colouring of horses. All English breeds,
however unlike in size and appearance, and several of those in India and
the Malay archipelago, present a similar range and diversity of colour. The
English race-horse, however, is said (2/30. 'The Field' July 13, 1861 page
42.) 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"
(2/31. E. Vernon Harcourt 'Sporting in Algeria' page 26.), 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 (2/32. I state this from my own observations made
during several years on the colours of horses. I have seen cream-coloured,
light-dun and mouse-dun horses dappled, which I mention because it has been
stated (Martin 'History of the Horse' page 134) that duns are never
dappled. Martin (page 205) refers to dappled asses. In the 'Farrier'
(London 1828 pages 453, 455) there are some good remarks on the dappling of
horses; and likewise in Col. Hamilton Smith on 'The Horse.'), in the same
manner as is so conspicuous with grey horses. This fact does not throw any
clear light on the colouring of the aboriginal horse, but is a case of
analogous variation, for even asses are sometimes dappled, and I have seen,
in the British Museum, a hybrid from the ass and zebra dappled on its
hinder quarters. By the expression analogous variation (and it is one that
I shall often have occasion to use) I mean a variation occurring in a
species or variety which resembles a normal character in another and
distinct species or variety. Analogous variations may arise, as will be
explained in a future chapter, from two or more forms with a similar
constitution having been exposed to similar conditions,--or from one of two
forms having reacquired through reversion a character inherited by the
other form from their common progenitor,--or from both forms having
reverted to the same ancestral character. We shall immediately see that
horses occasionally exhibit a tendency to become striped over a large part
of their bodies; and as we know that in the varieties of the domestic cat
and in several feline species stripes readily pass into spots and cloudy
marks--even the cubs of the uniformly-coloured lion being spotted with dark
marks on a lighter ground--we may suspect that the dappling of the horse,
which has been noticed by some authors with surprise, is a modification or
vestige of a tendency to become striped.
(FIGURE 1. DUN DEVONSHIRE PONY, with shoulder, spinal, and leg stripes.)
[This tendency in the horse to become striped is in several respects an
interesting fact. Horses of all colours, of the most diverse breeds, in
various parts of the world, often have a dark stripe extending along the
spine, from the mane to the tail; but this is so common that I need enter
into no particulars. (2/33. Some details are given in 'The Farrier' 1828
pages 452, 455. One of the smallest ponies I ever saw, of the colour of a
mouse, had a conspicuous spinal stripe. A small Indian chestnut pony had
the same stripe, as had a remarkably heavy chestnut cart-horse. Race-horses
often have the spinal stripe.) Occasionally horses are transversely barred
on the legs, chiefly on the under side; and more rarely they have a
distinct stripe on the shoulder, like that on the shoulder of the ass, or a
broad dark patch representing a stripe. Before entering on any details I
must premise that the term dun-coloured is vague, and includes three groups
of colours, viz., that between cream-colour and reddish-brown, which
graduates into light-bay or light-chestnut--this, I believe is often called
fallow-dun; secondly, leaden or slate-colour or mouse-dun, which graduates
into an ash-colour; and, lastly, dark-dun, between brown and black. In
England I have examined a rather large, lightly-built, fallow-dun
Devonshire pony (Figure 1), with a conspicuous stripe along the back, with
light transverse stripes on the under sides of its front legs, and with
four parallel stripes on each shoulder. Of these four stripes the posterior
one was very minute and faint; the anterior one, on the other hand, was
long and broad, but interrupted in the middle, and truncated at its lower
extremity, with the anterior angle produced into a long tapering point. I
mention this latter fact because the shoulder-stripe of the ass
occasionally presents exactly the same appearance. I have had an outline
and description sent to me of a small, purely-bred, light fallow-dun Welch
pony, with a spinal stripe, a single transverse stripe on each leg, and
three shoulder-stripes; the posterior stripe corresponding with that on the
shoulder of the ass was the longest, whilst the two anterior parallel
stripes, arising from the mane, decreased in length, in a reversed manner
as compared with the shoulder-stripes on the above-described Devonshire
pony. I have seen a bright fallow-dun cob, with its front legs transversely
barred on the under sides in the most conspicuous manner; also a dark-
leaden mouse-coloured pony with similar leg stripes, but much less
conspicuous; also a bright fallow-dun colt, fully three-parts thoroughbred,
with very plain transverse stripes on the legs; also a chestnut-dun cart-
horse with a conspicuous spinal stripe, with distinct traces of shoulder-
stripes, but none on the legs; I could add other cases. My son made a
sketch for me of a large, heavy, Belgian cart-horse, of a fallow-dun, with
a conspicuous spinal stripe, traces of leg-stripes, and with two parallel
(three inches apart) stripes about seven or eight inches in length on both
shoulders. I have seen another rather light cart-horse, of a dirty dark
cream-colour, with striped legs, and on one shoulder a large ill-defined
dark cloudy patch, and on the opposite shoulder two parallel faint stripes.
All the cases yet mentioned are duns of various tints; but Mr. W.W. Edwards
has seen a nearly thoroughbred chestnut horse which had the spinal stripe,
and distinct bars on the legs; and I have seen two bay carriage-horses with
black spinal stripes; one of these horses had on each shoulder a light
shoulder-stripe, and the other had a broad back ill-defined stripe, running
obliquely half-way down each shoulder; neither had leg-stripes.
The most interesting case which I have met with occurred in a colt of my
own breeding. A bay mare (descended from a dark-brown Flemish mare by a
light grey Turcoman horse) was put to Hercules, a thoroughbred dark bay,
whose sire (Kingston) and dam were both bays. The colt ultimately turned
out brown; but when only a fortnight old it was a dirty bay, shaded with
mouse-grey, and in parts with a yellowish tint: it had only a trace of the
spinal stripe, with a few obscure transverse bars on the legs; but almost
the whole body was marked with very narrow dark stripes, in most parts so
obscure as to be visible only in certain lights, like the stripes which may
be seen on black kittens. These stripes were distinct on the hind-quarters,
where they diverged from the spine, and pointed a little forwards; many of
them as they diverged became a little branched, exactly in the same manner
as in some zebrine species. The stripes were plainest on the forehead
between the ears, where they formed a set of pointed arches, one under the
other, decreasing in size downwards towards the muzzle; exactly similar
marks may be seen on the forehead of the quagga and Burchell's zebra. When
this foal was two or three months old all the stripes entirely disappeared.
I have seen similar marks on the forehead of a fully grown, fallow-dun,
cob-like horse, having a conspicuous spinal stripe, and with its front legs
well barred.
In Norway the colour of the native horse or pony is dun, varying from
almost cream-colour to dark-mouse dun; and an animal is not considered
purely bred unless it has the spinal and leg-stripes. (2/34. I have
received information, through the kindness of the Consul-General, Mr. J.R.
Crowe, from Prof. Boeck, Rasck, and Esmarck, on the colours of the
Norwegian ponies. See also 'The Field' 1861 page 431.) My son estimated
that about a third of the ponies which he saw there had striped legs; he
counted seven stripes on the fore-legs and two on the hind-legs of one
pony; only a few of them exhibited traces of shoulder stripes; but I have
heard of a cob imported from Norway which had the shoulder as well as the
other stripes well developed. Colonel H. Smith (2/35. Col. Hamilton Smith
'Nat. Lib.' volume 12 page 275.) alludes to dun-horses with the spinal
stripe in the Sierras of Spain; and the horses originally derived from
Spain, in some parts of South America, are now duns. Sir W. Elliot informs
me that he inspected a herd of 300 South American horses imported into
Madras, and many of these had transverse stripes on the legs and short
shoulder-stripes; the most strongly marked individual, of which a coloured
drawing was sent me, was a mouse-dun, with the shoulder-stripes slightly
forked.
In the North-Western parts of India striped horses of more than one breed
are apparently commoner than in any other part of the world; and I have
received information respecting them from several officers, especially from
Colonel Poole, Colonel Curtis, Major Campbell, Brigadier St. John, and
others. The Kattywar horses are often fifteen or sixteen hands in height,
and are well but lightly built. They are of all colours, but the several
kinds of duns prevail; and these are so generally striped, that a horse
without stripes is not considered pure. Colonel Poole believes that all the
duns have the spinal stripe, the leg-stripes are generally present, and he
thinks that about half the horses have the shoulder-stripe; this stripe is
sometimes double or treble on both shoulders. Colonel Poole has often seen
stripes on the cheeks and sides of the nose. He has seen stripes on the
grey and bay Kattywars when first foaled, but they soon faded away. I have
received other accounts of cream-coloured, bay, brown, and grey Kattywar
horses being striped. Eastward of India, the Shan (north of Burmah) ponies,
as I am informed by Mr. Blyth, have spinal, leg, and shoulder stripes. Sir
W. Elliot informs me that he saw two bay Pegu ponies with leg-stripes.
Burmese and Javanese ponies are frequently dun-coloured, and have the three
kinds of stripes, "in the same degree as in England." (2/36. Mr. G. Clark
in 'Annal and Mag. of Nat. History' 2nd series volume 2 1848 page 363. Mr.
Wallace informs me that he saw in Java a dun and clay-coloured horse with
spinal and leg stripes.) Mr. Swinhoe informs me that he examined two light-
dun ponies of two Chinese breeds, viz., those of Shanghai and Amoy; both
had the spinal stripe, and the latter an indistinct shoulder-stripe.
We thus see that in all parts of the world breeds of the horse as different
as possible, when of a dun-colour (including under this term a wide range
of tint from cream to dusty black), and rarely when almost white tinged
with yellow, grey, bay, and chestnut, have the several above-specified
stripes. Horses which are of a yellow colour with white mane and tail, and
which are sometimes called duns, I have never seen with stripes. (2/37. See
also on this point 'The Field' July 27, 1861 page 91.)
From reasons which will be apparent in the chapter on Reversion, I have
endeavoured, but with poor success, to discover whether duns, which are so
much oftener striped than other coloured horses, are ever produced from the
crossing of two horses, neither of which are duns. Most persons to whom I
have applied believe that one parent must be dun; and it is generally
asserted that, when this is the case, the dun-colour and the stripes are
strongly inherited. (2/38. 'The Field' 1861 pages 431, 493, 545.) One case,
however, has fallen under my own observation of a foal from a black mare by
a bay horse, which when fully grown was a dark fallow-dun and had a narrow
but plain spinal stripe. Hofacker (2/39. 'Ueber die Eigenschaften' etc.
1828 s. 13, 14.) gives two instances of mouse-duns (Mausrapp) being
produced from two parents of different colours and neither duns.
The stripes of all kinds are generally plainer in the foal than in the
adult horse, being commonly lost at the first shedding of the hair. (2/40.
Von Nathusius 'Vortrage uber Viehzucht' 1872 135.) Colonel Poole believes
that "the stripes in the Kattywar breed are plainest when the colt is first
foaled; they then become less and less distinct till after the first coat
is shed, when they come out as strongly as before; but certainly often fade
away as the age of the horse increases." Two other accounts confirm this
fading of the stripes in old horses in India. One writer, on the other
hand, states that colts are often born without stripes, but that they
appear as the colt grows older. Three authorities affirm that in Norway the
stripes are less plain in the foal than in the adult. In the case described
by me of the young foal which was narrowly striped over nearly all its
body, there was no doubt about the early and complete disappearance of the
stripes. Mr. W.W. Edwards examined for me twenty-two foals of race-horses,
and twelve had the spinal stripe more or less plain; this fact, and some
other accounts which I have received, lead me to believe that the spinal
stripe often disappears in the English race-horse when old. With natural
species, the young often exhibit characters which disappear at maturity.]
The stripes are variable in colour, but are always darker than the rest of
the body. They do not by any means always coexist on the different parts of
the body: the legs may be striped without any shoulder-stripe, or the
converse case, which is rarer, may occur; but I have never heard of either
shoulder or leg-stripes without the spinal stripe. The latter is by far the
commonest of all the stripes, as might have been expected, as it
characterises the other seven or eight species of the genus. It is
remarkable that so trifling a character as the shoulder-stripe being double
or triple should occur in such different breeds as Welch and Devonshire
ponies, the Shan pony, heavy cart-horses, light South American horses, and
the lanky Kattywar breed. Colonel Hamilton Smith believes that one of his
five supposed primitive stocks was dun-coloured and striped; and that the
stripes in all the other breeds result from ancient crosses with this one
primitive dun; but it is extremely improbable that different breeds living
in such distant quarters of the world should all have been crossed with any
one aboriginally distinct stock. Nor have we any reason to believe that the
effects of a cross at a very remote period would be propagated for so many
generations as is implied on this view.
With respect to the primitive colour of the horse having been dun, Colonel
Hamilton Smith (2/41. 'Nat. Library' volume 12 1841 pages 109, 156 to 163,
280, 281. Cream-colour, passing into Isabella (i.e. the colour of the dirty
linen of Queen Isabella), seems to have been common in ancient times. See
also Pallas's account of the wild horses of the East, who speaks of dun and
brown as the prevalent colours. In the Icelandic sagas, which were
committed to writing in the twelfth century, dun-coloured horses with a
black spinal stripe are mentioned; see Dasent's translation volume 1 page
169.) has collected a large body of evidence showing that this tint was
common in the East as far back as the time of Alexander, and that the wild
horses of Western Asia and Eastern Europe now are, or recently were, of
various shades of dun. It seems that not very long ago a wild breed of dun-
coloured horses with a spinal stripe was preserved in the royal parks in
Prussia. I hear from Hungary that the inhabitants of that country look at
the duns with a spinal stripe as the aboriginal stock, and so it is in
Norway. Dun-coloured ponies are not rare in the mountainous parts of
Devonshire, Wales, and Scotland, where the aboriginal breed would have the
best chance of being preserved. In South America in the time of Azara, when
the horse had been feral for about 250 years, 90 out of 100 horses were
"bai-chatains," and the remaining ten were "zains," that is brown; not more
than one in 2000 being black. In North America the feral horses show a
strong tendency to become roans of various shades; but in certain parts, as
I hear from Dr. Canfield, they are mostly duns and striped. (2/42. Azara
'Quadrupedes du Paraguay' tome 2 page 307. In North America Catlin (volume
2 page 57) describes the wild horses, believed to have descended from the
Spanish horses of Mexico, as of all colours, black, grey, roan, and roan
pied with sorrel. F. Michaux 'Travels in North America' English translation
page 235, describes two wild horses from Mexico as roan. In the Falkland
Islands, where the horse has been feral only between 60 and 70 years, I was
told that roans and iron-greys were the prevalent colours. These several
facts show that horses do not soon revert to any uniform colour.)
In the following chapters on the Pigeon we shall see that a blue bird is
occasionally produced by pure breeds of various colours and that when this
occurs certain black marks invariably appear on the wings and tail; so
again, when variously coloured breeds are crossed, blue birds with the same
black marks are frequently produced. We shall further see that these facts
are explained by, and afford strong evidence in favour of, the view that
all the breeds are descended from the rock-pigeon, or Columba livia, which
is thus coloured and marked. But the appearance of the stripes on the
various breeds of the horse, when of a dun colour, does not afford nearly
such good evidence of their descent from a single primitive stock as in the
case of the pigeon: because no horse certainly wild is known as a standard
of comparison; because the stripes when they appear are variable in
character; because there is far from sufficient evidence that the crossing
of distinct breeds produces stripes, and lastly, because all the species of
the genus Equus have the spinal stripe, and several species have shoulder
and leg stripes. Nevertheless the similarity in the most distinct breeds in
their general range of colour, in their dappling, and in the occasional
appearance, especially in duns, of leg-stripes and of double or triple
shoulder-stripes, taken together, indicate the probability of the descent
of all the existing races from a single, dun-coloured, more or less
striped, primitive stock, to which our horses occasionally revert.
THE ASS.
Four species of Asses, besides three zebras, have been described by
naturalists. There is now little doubt that our domesticated animal is
descended from the Equus taeniopus of Abyssinia. (2/43. Dr. Sclater in
'Proc. Zoolog. Soc.' 1862 page 164. Dr. Hartmann says ('Annalen der Landw.'
b. 44 page 222) that this animal in its wild state is not always striped
across the legs.) The ass is sometimes advanced as an instance of an animal
domesticated, as we know by the Old Testament, from an ancient period,
which has varied only in a very slight degree. But this is by no means
strictly true; for in Syria alone there are four breeds (2/44. W.C. Martin
'History of the Horse' 1845 page 207.); first, a light and graceful animal,
with an agreeable gait, used by ladies; secondly, an Arab breed reserved
exclusively for the saddle; thirdly, a stouter animal used for ploughing
and various purposes; and lastly, the large Damascus breed, with a
peculiarly long body and ears. In the South of France also there are
several breeds, and one of extraordinary size, some individuals being as
tall as full-sized horses. Although the ass in England is by no means
uniform in appearance, distinct breeds have not been formed. This may
probably be accounted for by the animal being kept chiefly by poor persons,
who do not rear large numbers, nor carefully match and select the young.
For, as we shall see in a future chapter, the ass can with ease be greatly
improved in size and strength by careful selection, combined no doubt with
good food; and we may infer that all its other characters would be equally
amenable to selection. The small size of the ass in England and Northern
Europe is apparently due far more to want of care in breeding than to cold;
for in Western India, where the ass is used as a beast of burden by some of
the lower castes, it is not much larger than a Newfoundland dog, "being
generally not more than from twenty to thirty inches high." (2/45. Col.
Sykes Cat. of Mammalia 'Proc. Zoolog. Soc.' July 12, 1831. Williamson
'Oriental Field Sports' volume 2 quoted by Martin page 206.)
The ass varies greatly in colour; and its legs, especially the fore-legs,
both in England and other countries--for instance, in China--are
occasionally barred more plainly than those of dun-coloured horses.
Thirteen or fourteen transverse stripes have been counted on both the fore
and hind legs. With the horse the occasional appearance of leg-stripes was
accounted for by reversion to a supposed parent-form, and in the case of
the ass we may confidently believe in this explanation, as E. taeniopus is
known to be barred, though only in a slight degree, and not quite
invariably. The stripes are believed to occur most frequently and to be
plainest on the legs of the domestic ass during early youth (2/46. Blyth in
'Charlesworth's Mag. of Nat. Hist.' vol 4 1840 page 83. I have also been
assured by a breeder that this is the case.), as likewise occurs with the
horse. The shoulder-stripe, which is so eminently characteristic of the
species, is nevertheless variable in breadth, length, and manner of
termination. I have measured one four times as broad as another, and some
more than twice as long as others. In one light-grey ass the shoulder-
stripe was only six inches in length, and as thin as a piece of string; and
in another animal of the same colour there was only a dusky shade
representing a stripe. I have heard of three white asses, not albinoes,
with no trace of shoulder or spinal stripes (2/47. One case is given by
Martin 'The Horse' page 205.); and I have seen nine other asses with no
shoulder-stripe, and some of them had no spinal stripe. Three of the nine
were light-greys, one a dark-grey, another grey passing into reddish-roan,
and the others were brown, two being tinted on parts of their bodies with a
reddish or bay shade. If therefore grey and reddish-brown asses had been
steadily selected and bred from, the shoulder stripe would probably have
been lost almost as generally and completely as in the case of the horse.
The shoulder stripe on the ass is sometimes double, and Mr. Blyth has seen
even three or four parallel stripes. (2/48. 'Journal As. Soc. of Bengal'
volume 28 1860 page 231. Martin on the Horse page 205.) I have observed in
ten cases shoulder-stripes abruptly truncated at the lower end, with the
anterior angle produced into a tapering point, precisely as in the above
dun Devonshire pony. I have seen three cases of the terminal portion
abruptly and angularly bent; and have seen and heard of four cases of a
distinct though slight forking of the stripe. In Syria, Dr. Hooker and his
party observed for me no less than five similar instances of the shoulder-
stripe plainly bifurcating over the fore leg. In the common mule it
likewise sometimes bifurcates. When I first noticed the forking and angular
bending of the shoulder-stripe, I had seen enough of the stripes in the
various equine species to feel convinced that even a character so
unimportant as this had a distinct meaning, and was thus led to attend to
the subject. I now find that in the E. burchellii and quagga, the stripe
which corresponds with the shoulder-stripe of the ass, as well as some of
the stripes on the neck, bifurcate, and that some of those near the
shoulder have their extremities bent angularly backwards. The bifurcation
and angular bending of the stripes on the shoulders apparently are
connected with the nearly upright stripes on the sides of the body and neck
changing their direction and becoming transverse on the legs. Finally, we
see that the presence of shoulder, leg, and spinal stripes in the horse,--
their occasional absence in the ass,--the occurrence of double and triple
shoulder-stripes in both animals, and the similar manner in which these
stripes terminate downwards,--are all cases of analogous variation in the
horse and ass. These cases are probably not due to similar conditions
acting on similar constitutions, but to a partial reversion in colour to
the common progenitor of the genus. We shall hereafter return to this
subject, and discuss it more fully.
CHAPTER 1.III.
PIGS--CATTLE--SHEEP--GOATS.
PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICUS.
TORFSCHWEIN.
JAPAN PIGS.
FERTILITY OF CROSSED PIGS.
CHANGES IN THE SKULL OF THE HIGHLY CULTIVATED RACES.
CONVERGENCE OF CHARACTER.
GESTATION.
SOLID-HOOFED SWINE.
CURIOUS APPENDAGES TO THE JAWS.
DECREASE IN SIZE OF THE TUSKS.
YOUNG PIGS LONGITUDINALLY STRIPED.
FERAL PIGS.
CROSSED BREEDS.
CATTLE.
ZEBU A DISTINCT SPECIES.
EUROPEAN CATTLE PROBABLY DESCENDED FROM THREE WILD FORMS.
ALL THE RACES NOW FERTILE TOGETHER.
BRITISH PARK CATTLE.
ON THE COLOUR OF THE ABORIGINAL SPECIES.
CONSTITUTIONAL DIFFERENCES.
SOUTH AFRICAN RACES.
SOUTH AMERICAN RACES.
NIATA CATTLE.
ORIGIN OF THE VARIOUS RACES OF CATTLE.
SHEEP.
REMARKABLE RACES OF.
VARIATIONS ATTACHED TO THE MALE SEX.
ADAPTATIONS TO VARIOUS CONDITIONS.
GESTATION OF.
CHANGES IN THE WOOL.
SEMI-MONSTROUS BREEDS.
GOATS.
REMARKABLE VARIATIONS OF.
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 Rutimeyer in his celebrated Fauna of the ancient Swiss lake-dwellings.
(3/1. Hermann von Nathusius 'Die Racen des Schweines' Berlin 1860; and
'Vorstudien fur Geschichte' etc. 'Schweineschadel' Berlin 1864. Rutimeyer
'Die Fauna der Pfahlbauten' Basel 1861.) Nathusius has shown that all the
known breeds may be divided into two great groups: one resembling in all
important respects and no doubt descended from the common wild boar; so
that this may be called the Sus scrofa group. The other group differs in
several important and constant osteological characters; its wild parent-
form is unknown; the name given to it by Nathusius, according to the law of
priority, is Sus indicus, of Pallas. This name must now be followed, though
an unfortunate one, as the wild aboriginal does not inhabit India, and the
best-known domesticated breeds have been imported from Siam and China.
First for the Sus scrofa breeds, or those resembling the common wild boar.
These still exist, according to Nathusius ('Schweineschadel' s. 75), in
various parts of central and northern Europe; formerly every kingdom (3/2.
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), and almost every province in Britain, possessed its own
native breed; but these are now everywhere rapidly disappearing, being
replaced by improved breeds crossed with the S. indicus form. The skull in
the breeds of the S. scrofa type resembles, in all important respects, that
of the European wild boar; but it has become ('Schweineschadel' s. 63-68)
higher and broader relatively to its length; and the hinder part is more
upright. The differences, however, are all variable in degree. The breeds
which thus resemble S. scrofa in their essential skull characters differ
conspicuously from each other in other respects, as in the length of the
ears and legs, curvature of the ribs, colour, hairiness, size and
proportions of the body.
The wild Sus scrofa has a wide range, namely, Europe, North Africa, as
identified by osteological characters by Rutimeyer, and Hindostan, as
similarly identified by Nathusius. But the wild boars inhabiting these
several countries differ so much from each other in external characters,
that they have been ranked by some naturalists as specifically distinct.
Even within Hindostan these animals, according to Mr. Blyth, form very
distinct races in the different districts; in the N. Western provinces, as
I am informed by the Rev. R. Everest, the boar never exceeds 36 inches in
height, whilst in Bengal one has been measured 44 inches in height. In
Europe, Northern Africa, and Hindostan, domestic pigs have been known to
cross with the wild native species (3/3. For Europe see Bechstein
'Naturgesch. Deutschlands' 1801 b. 1 s. 505. Several accounts have been
published on the fertility of the offspring from wild and tame swine. See
Burdach 'Physiology' and Godron 'De l'Espece' tome 1 page 370. For Africa
'Bull. de la Soc. d'Acclimat.' tome 4 page 389. For India see Nathusius
'Schweineschadel' s. 148.); and in Hindostan an accurate observer (3/4. Sir
W. Elliot Catalogue of Mammalia 'Madras Journal of Lit. and Science' volume
10 page 219.), Sir Walter Elliot, after describing the differences between
wild Indian and wild German boars, remarks that "the same differences are
perceptible in the domesticated individuals of the two countries." We may
therefore conclude that the breeds of the Sus scrofa type are descended
from, or have been modified by crossing with, forms which may be ranked as
geographical races, but which, according to some naturalists, ought to be
ranked as distinct species.
Pigs of the Sus indicus type are best known to Englishmen under the form of
the Chinese breed. The skull of S. indicus, as described by Nathusius,
differs from that of S. scrofa in several minor respects, as in its greater
breadth and in some details in the teeth; but chiefly in the shortness of
the lachrymal bones, in the greater width of the fore part of the palate-
bones, and in the divergence of the premolar teeth. It deserves especial
notice that these latter characters are not gained, even in the least
degree, by the domesticated forms of S. scrofa. After reading the remarks
and descriptions given by Nathusius, it seems to me to be merely playing
with words to doubt whether S. indicus ought to be ranked as a species; for
the above-specified differences are more strongly marked than any that can
be pointed out between, for instance, the fox and the wolf, or the ass and
the horse. As already stated, S. indicus is not known in a wild state; but
its domesticated forms, according to Nathusius, come near to S. vittatus of
Java and some allied species. A pig found wild in the Aru islands
('Schweineschadel' s. 169) is apparently identical with S. indicus; but it
is doubtful whether this is a truly native animal. The domesticated breeds
of China, Cochin-China, and Siam belong to this type. The Roman or
Neapolitan breed, the Andalusian, the Hungarian, and the "Krause" swine of
Nathusius, inhabiting south-eastern Europe and Turkey, and having fine
curly hair, and the small Swiss "Bundtnerschwein" of Rutimeyer, all agree
in their more important skull-characters with S. indicus, and, as is
supposed, have all been largely crossed with this form. Pigs of this type
have existed during a long period on the shores of the Mediterranean, for a
figure ('Schweineschadel' s. 142) closely resembling the existing
Neapolitan pig was found in the buried city of Herculaneum.
Rutimeyer has made the remarkable discovery that there lived
contemporaneously in Switzerland, during the Neolithic period, two
domesticated forms, the S. scrofa, and the S. scrofa palustris or
Torfschwein. Rutimeyer perceived that the latter approached the Eastern
breeds, and, according to Nathusius, it certainly belongs to the S. indicus
group; but Rutimeyer has subsequently shown that it differs in some well-
marked characters. This author was formerly convinced that his Torfschwein
existed as a wild animal during the first part of the Stone period, and was
domesticated during a later part of the same period. (3/5. 'Pfahlbauten' s.
163 et passim.) Nathusius, whilst he fully admits the curious fact first
observed by Rutimeyer, that the bones of domesticated and wild animals can
be distinguished by their different aspect, yet, from special difficulties
in the case of the bones of the pig ('Schweineschadel' s. 147), is not
convinced of the truth of the above conclusion; and Rutimeyer himself seems
now to feel some doubt. Other naturalists have also argued strongly on the
same side as Nathusius. (3/6. See J.W. Schutz' interesting essay 'Zur
Kenntniss des Torfschweins' 1868. This author believes that the Torfschwein
is descended from a distinct species, the S. sennariensis of Central
Africa.)
Several breeds, differing in the proportions of the body, in the length of
the ears, in the nature of the hair, in colour, etc., come under the S.
indicus type. Nor is this surprising, considering how ancient the
domestication of this form has been both in Europe and in China. In this
latter country the date is believed by an eminent Chinese scholar (3/7.
Stan. Julien quoted by de Blainville 'Osteographie' page 163.) to go back
at least 4900 years from the present time. This same scholar alludes to the
existence of many local varieties of the pig in China; and at the present
time the Chinese take extraordinary pains in feeding and tending their
pigs, not even allowing them to walk from place to place. (3/8. Richardson
'Pigs, their Origin' etc. page 26.) Hence these pigs, as Nathusius has
remarked (3/9. 'Die Racen des Schweines' s. 47, 64.), display in an eminent
degree the characters of a highly-cultivated race, and hence, no doubt,
their high value in the improvement of our European breeds. Nathusius makes
a remarkable statement ('Schweineschadel' s. 138), that the infusion of the
1/32nd, or even of the 1/64th, part of the blood of S. indicus into a breed
of S. scrofa, is sufficient plainly to modify the skull of the latter
species. This singular fact may perhaps be accounted for by several of the
chief distinctive characters of S. indicus, such as the shortness of the
lachrymal bones, etc., being common to several species of the genus; for in
crosses characters which are common to many species apparently tend to be
prepotent over those appertaining to only a few species.
(FIGURE 2. HEAD OF JAPAN OR MASKED PIG. (Copied from Mr. Bartlett's paper
in 'Proc. Zoolog. Soc.' 1861 page 263.))
The Japan pig (S. pliciceps of Gray), which was formerly exhibited in the
Zoological Gardens, has an extraordinary appearance from its short head,
broad forehead and nose, great fleshy ears, and deeply furrowed skin.
Figure 2 is copied from that given by Mr. Bartlett. (3/10. 'Proc. Zoolog.
Soc.' 1861 page 263.) Not only is the face furrowed, but thick folds of
skin, which are harder than the other parts, almost like the plates on the
Indian rhinoceros, hang about the shoulders and rump. It is coloured black,
with white feet, and breeds true. That it has long been domesticated there
can be little doubt; and this might have been inferred even from the fact
that its young are not longitudinally striped; for this is a character
common to all the species included within the genus Sus and the allied
genera whilst in their natural state. (3/11. Sclater in 'Proc. Zoolog.
Soc.' February 26, 1861.) Dr. Gray (3/12. 'Proc. Zoolog. Soc.' 1862 page
13. The skull has since been described much more fully by Professor Lucae
in a very interesting essay, 'Der Schadel des Maskenschweines' 1870. He
confirms the conclusion of von Nathusius on the relationship of this kind
of pig.) has described the skull of this animal, which he ranks not only as
a distinct species, but places it in a distinct section of the genus.
Nathusius, however, after his careful study of the whole group, states
positively ('Schweineschadel' s. 153-158). that the skull in all essential
characters closely resembles that of the short-eared Chinese breed of the
S. indicus type. Hence Nathusius considers the Japan pig as only a
domesticated variety of S. indicus: if this really be the case, it is a
wonderful instance of the amount of modification which can be effected
under domestication.
Formerly there existed in the central islands of the Pacific Ocean a
singular breed of pigs. These are described by the Rev. D. Tyerman and G.
Bennett (3/13. 'Journal of Voyages and Travels from 1821 to 1829' volume 1
page 300.) as of small size, hump-backed, with a disproportionately long
head, with short ears turned backwards, with a bushy tail not more than two
inches in length, placed as if it grew from the back. Within half a century
after the introduction of European and Chinese pigs into these islands, the
native breed, according to the above authors, became almost completely lost
by being repeatedly crossed with them. Secluded islands, as might have been
expected, seem favourable for the production or retention of peculiar
breeds; thus, in the Orkney Islands, the hogs have been described as very
small, with erect and sharp ears, and "with an appearance altogether
different from the hogs brought from the south." (3/14. Rev. G. Low 'Fauna
Orcadensis' page 10. See also Dr. Hibbert's account of the pig of the
Shetland Islands.)
Seeing how different the Chinese pigs, belonging to the Sus indicus type,
are in their osteological characters and in external appearance from the
pigs of the S. scrofa type, so that they must be considered specifically
distinct, it is a fact well deserving attention, that Chinese and common
pigs have been repeatedly crossed in various manners, with unimpaired
fertility. One great breeder who had used pure Chinese pigs assured me that
the fertility of the half-breeds inter se and of their recrossed progeny
was actually increased; and this is the general belief of agriculturists.
Again, the Japan pig or S. pliciceps of Gray is so distinct in appearance
from all common pigs, that it stretches one's belief to the utmost to admit
that it is simply a domestic variety; yet this breed has been found
perfectly fertile with the Berkshire breed; and Mr. Eyton informs me that
he paired a half-bred brother and sister and found them quite fertile
together.
(FIGURE 3. HEAD OF WILD BOAR, AND OF "GOLDEN DAYS," a pig of the Yorkshire
Large Breed; the latter from a photograph. (Copied from Sidney's edition of
'The Pig' by Youatt.))
The modification of the skull in the most highly cultivated races is
wonderful. To appreciate the amount of change, Nathusius' work, with its
excellent figures, should be studied. The whole of the exterior in all its
parts has been altered: the hinder surface, instead of sloping backwards,
is directed forwards, entailing many changes in other parts; the front of
the head is deeply concave; the orbits have a different shape; the auditory
meatus has a different direction and shape; the incisors of the upper and
lower jaws do not touch each other, and they stand in both jaws beyond the
plane of the molars; the canines of the upper jaw stand in front of those
of the lower jaw, and this is a remarkable anomaly: the articular surfaces
of the occipital condyles are so greatly changed in shape, that, as
Nathusius remarks (s. 133), no naturalist, seeing this important part of
the skull by itself, would suppose that it belonged to the genus Sus. These
and various other modifications, as Nathusius observes, can hardly be
considered as monstrosities, for they are not injurious, and are strictly
inherited. The whole head is much shortened; thus, whilst in common breeds
its length to that of the body is as 1 to 6, in the "cultur-racen" the
proportion is as 1 to 9, and even recently as 1 to 11. (3/15. 'Die Racen
des Schweines' s. 70.) Figure 3 (3/16. These woodcuts are copied from
engravings given in Mr. S. Sidney's excellent edition of 'The Pig' by
Youatt 1860. See pages 1, 16, 19.) of the head of a wild boar and of a sow
from a photograph of the Yorkshire Large Breed, may aid in showing how
greatly the head in a highly cultivated race has been modified and
shortened.
Nathusius has well discussed the causes of the remarkable changes in the
skull and shape of the body which the highly cultivated races have
undergone. These modifications occur chiefly in the pure and crossed races
of the S. indicus type; but their commencement may be clearly detected in
the slightly improved breeds of the S. scrofa type. (3/17.
'Schweineschadel' s. 74, 135.) Nathusius states positively (s. 99, 103), as
the result of common experience and of his experiments, that rich and
abundant food, given during youth, tends by some direct action to make the
head broader and shorter; and that poor food works a contrary result. He
lays much stress on the fact that all wild and semi-domesticated pigs, in
ploughing up the ground with their muzzles, have,
whilst young, to exert the powerful muscles fixed to the hinder part of the
head. In highly cultivated races this habit is no longer followed, and
consequently the back of the skull becomes modified in shape, entailing
other changes in other parts. There can hardly be a doubt that so great a
change in habits would affect the skull; but it seems rather doubtful how
far this will account for the greatly reduced length of the skull and for
its concave front. It is well known (Nathusius himself advancing many
cases, s. 104) that there is a strong tendency in many domestic animals--in
bull- and pug-dogs, in the niata cattle, in sheep, in Polish fowls, short-
faced tumbler pigeons, and in one variety of the carp--for the bones of the
face to become greatly shortened. In the case of the dog, as H. Muller has
shown, this seems caused by an abnormal state of the primordial cartilage.
We may, however, readily admit that abundant and rich food supplied during
many generations would give an inherited tendency to increased size of
body, and that, from disuse, the limbs would become finer and shorter.
(3/18. Nathusius 'Die Racen des Schweines' s. 71.) We shall in a future
chapter see also that the skull and limbs are apparently in some manner
correlated, so that any change in the one tends to affect the other.
Nathusius has remarked, and the observation is an interesting one, that the
peculiar form of the skull and body in the most highly cultivated races is
not characteristic of any one race, but is common to all when improved up
to the same standard. Thus the large-bodied, long-eared, English breeds
with a convex back, and the small-bodied, short-eared, Chinese breeds with
a concave back, when bred to the same state of perfection, nearly resemble
each other in the form of the head and body. This result, it appears, is
partly due to similar causes of change acting on the several races, and
partly to man breeding the pig for one sole purpose, namely, for the
greatest amount of flesh and fat; so that selection has always tended
towards one and the same end. With most domestic animals the result of
selection has been divergence of character, here it has been convergence.
(3/19. 'Die Racen des Schweines' s. 47. 'Schweineschadel' s. 104. Compare
also the figures of the old Irish and the improved Irish breeds in
Richardson on 'The Pig' 1847.)
The nature of the food supplied during many generations has apparently
affected the length of the intestines; for, according to Cuvier (3/20.
Quoted by Isid. Geoffroy 'Hist. Nat. Gen.' tome 3 page 441.), their length
to that of the body in the wild boar is as 9 to 1,--in the common domestic
boar as 13.5 to 1,--and in the Siam breed as 16 to 1. In this latter breed
the greater length may be due either to descent from a distinct species or
to more ancient domestication. The number of mammae vary, as does the
period of gestation. The latest authority says (3/21. S. Sidney 'The Pig'
page 61.) that "the period averages from 17 to 20 weeks," but I think there
must be some error in this statement: in M. Tessier's observations on 25
sows it varied from 109 to 123 days. The Rev. W.D. Fox has given me ten
carefully recorded cases with well-bred pigs, in which the period varied
from 101 to 116 days. According to Nathusius the period is shortest in the
races which come early to maturity; but the course of their development
does not appear to be actually shortened, for the young animal is born,
judging from the state of the skull, less fully developed, or in a more
embryonic condition (3/22. 'Schweineschadel' s. 2, 20.) than in the case of
common swine. In the highly cultivated and early matured races the teeth,
also, are developed earlier.
The difference in the number of the vertebrae and ribs in different kinds
of pigs, as observed by Mr. Eyton (3/23. 'Proc. Zoolog. Soc.' 1837 page 23.
I have not given the caudal vertebrae, as Mr. Eyton says some might
possibly have been lost. I have added together the dorsal and lumbar
vertebrae, owing to Prof. Owen's remarks ('Journal Linn. Soc.' volume 2
page 28) on the difference between dorsal and lumbar vertebrae depending
only on the development of the ribs. Nevertheless the difference in the
number of the ribs in pigs deserves notice. M. Sanson gives the number of
lumbar vertebrae in various pigs; 'Comptes Rendus' 93 page 843.), and as
given in the following table, has often been quoted. The African sow
probably belongs to the S. scrofa type; and Mr. Eyton informs me that,
since the publication of this paper, cross-bred animals from the African
and English races were found by Lord Hill to be perfectly fertile.
TABLE 2: NUMBER OF VERTEBRAE IN VARIOUS PIGS:
ENGLISH LONG-LEGGED MALE.
Dorsal 15. Lumbar 6. Dorsal plus Lumbar 21. Sacral 5. Total 26.
AFRICAN FEMALE.
Dorsal 13. Lumbar 6. Dorsal plus Lumbar 19. Sacral 5. Total 24.
CHINESE MALE.
Dorsal 15. Lumbar 4. Dorsal plus Lumbar 19. Sacral 4. Total 23.
WILD BOAR FROM CUVIER.
Dorsal 14. Lumbar 5. Dorsal plus Lumbar 19. Sacral 4. Total 23.
FRENCH DOMESTIC BOAR, FROM CUVIER.
Dorsal 14. Lumbar 5. Dorsal plus Lumbar 19. Sacral 4. Total 23.
Some semi-monstrous breeds deserve notice. From the time of Aristotle to
the present time solid-hoofed swine have occasionally been observed in
various parts of the world. Although this peculiarity is strongly
inherited, it is hardly probable that all the animals with solid hoofs have
descended from the same parents; it is more probable that the same
peculiarity has reappeared at various times and places. Dr. Struthers has
lately described and figured (3/24. 'Edinburgh New Philosoph. Journal'
April 1863. See also De Blainville 'Osteographie' page 128 for various
authorities on this subject.) the structure of the feet; in both front and
hind feet the distal phalanges of the two greater toes are represented by a
single, great, hoof-bearing phalanx; and in the front feet, the middle
phalanges are represented by a bone which is single towards the lower end,
but bears two separate articulations towards the upper end. From other
accounts it appears that an intermediate toe is likewise sometimes
superadded.
(FIGURE 4. OLD IRISH PIG, with jaw appendages. (Copied from H.D. Richardson
on Pigs.))
Another curious anomaly is offered by the appendages, described by M.
Eudes-Deslongchamps as often characterizing the Normandy pigs. These
appendages are always attached to the same spot, to the corners of the jaw;
they are cylindrical, about three inches in length, covered with bristles,
and with a pencil of bristles rising out of a sinus on one side: they have
a cartilaginous centre, with two small longitudinal muscles they occur
either symmetrically on both sides of the face or on one side alone.
Richardson figures them on the gaunt old "Irish Greyhound pig;" and
Nathusius states that they occasionally appear in all the long eared races,
but are not strictly inherited, for they occur or fail in animals of the
same litter. (3/25. Eudes-Deslongchamps 'Memoires de la Soc. Linn. de
Normandie' volume 7 1842 page 41. Richardson 'Pigs, their Origin, etc.'
1847 page 30. Nathusius 'Die Racen des Schweines' 1863 s. 54.) As no wild
pigs are known to have analogous appendages, we have at present no reason
to suppose that their appearance is due to reversion; and if this be so, we
are forced to admit that a somewhat complex, though apparently useless,
structure may be suddenly developed without the aid of selection.
It is a remarkable fact that the boars of all domesticated breeds have much
shorter tusks than wild boars. Many facts show that with many animals the
state of the hair is much affected by exposure to, or protection from,
climate; and as we see that the state of the hair and teeth are correlated
in Turkish dogs (other analogous facts will be hereafter given), may we not
venture to surmise that the reduction of the tusks in the domestic boar is
related to his coat of bristles being diminished from living under shelter?
On the other hand, as we shall immediately see, the tusks and bristles
reappear with feral boars, which are no longer protected from the weather.
It is not surprising that the tusks should be more affected than the other
teeth; as parts developed to serve as secondary sexual characters are
always liable to much variation.
It is a well-known fact that the young of wild European and Indian pigs
(3/26. D. Johnson 'Sketches of Indian Field Sports' page 272. Mr. Crawfurd
informs me that the same fact holds good with the wild pigs of the Malay
peninsula.), for the first six months, are longitudinally banded with
light-coloured stripes. This character generally disappears under
domestication. The Turkish domestic pigs, however, have striped young, as
have those of Westphalia, "whatever may be their hue" (3/27. For Turkish
pigs see Desmarest 'Mammalogie' 1820 page 391. For those of Westphalia see
Richardson 'Pigs, their Origin, etc.' 1847 page 41.); whether these latter
pigs belong to the same curly-haired race as the Turkish swine, I do not
know. The pigs which have run wild in Jamaica and the semi-feral pigs of
New Granada, both those which are black and those which are black with a
white band across the stomach, often extending over the back, have resumed
this aboriginal character and produce longitudinally-striped young. This is
likewise the case, at least occasionally, with the neglected pigs in the
Zambesi settlement on the coast of Africa. (3/28. With respect to the
several foregoing and following statements on feral pigs see Roulin in
'Mem. presentes par divers Savans a l'Acad.' etc. Paris tome 6 1835 page
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 'Sojourn in
Jamaica' 1851 page 386; and Col Hamilton Smith in 'Nat. Library' volume 9
page 93. With respect to Africa see Livingstone 'Expedition to the Zambesi'
1865 page 153. The most precise statement with respect to the tusks of the
West Indian feral boars is by P. Labat quoted by Roulin; but this author
attributes the state of these pigs to descent from a domestic stock which
he saw in Spain. Admiral Sulivan, R.N., had ample opportunities of
observing the wild pigs on Eagle Islet in the Falklands; and he informs me
that they resembled wild boars with bristly ridged backs and large tusks.
The pigs which have run wild in the province of Buenos Ayres (Rengger
'Saugethiere' s. 331) have not reverted to the wild type. De Blainville
'Osteographie' page 132 refers to two skulls of domestic pigs sent from
Patagonia by Al. d'Orbigny, and he states that they have the occipital
elevation of the wild European boar, but that the head altogether is "plus
courte et plus ramassee." He refers also to the skin of a feral pig from
North America, and says "il ressemble tout a fait a un petit sanglier, mais
il est presque tout noir, et peut-etre un peu plus ramasse dans ses
formes."
The common belief that all domesticated animals, when they run wild, revert
completely to the character of their parent-stock, is chiefly founded, as
far as I can discover, on feral pigs. But even in this case the belief is
not grounded on sufficient evidence; for the two main types, namely, S.
scrofa and indicus, have not been distinguished. The young, as we have just
seen, reacquire their longitudinal stripes, and the boars invariably
reassume their tusks. They revert also in the general shape of their
bodies, and in the length of their legs and muzzles, to the state of the
wild animal, as might have been expected from the amount of exercise which
they are compelled to take in search of food. In Jamaica the feral pigs do
not acquire the full size of the European wild boar, "never attaining a
greater height than 20 inches at the shoulder." In various countries they
reassume their original bristly covering, but in different degrees,
dependent on the climate; thus, according to Roulin, the semi-feral pigs in
the hot valleys of New Granada are very scantily clothed; whereas, on the
Paramos, at the height of 7000 to 8000 feet, they acquire a thick covering
of wool lying under the bristles, like that on the truly wild pigs of
France. These pigs on the Paramos are small and stunted. The wild boar of
India is said to have the bristles at the end of its tail arranged like the
plumes of an arrow, whilst the European boar has a simple tuft; and it is a
curious fact that many, but not all, of the feral pigs in Jamaica, derived
from a Spanish stock, have a plumed tail. (3/29. Gosse 'Jamaica' page 386
with a quotation from Williamson 'Oriental Field Sports.' Also Col.
Hamilton Smith in 'Naturalist Library' volume 9 page 94.) With respect to
colour, feral pigs generally revert to that of the wild boar; but in
certain parts of S. America, as we have seen, some of the semi-feral pigs
have a curious white band across their stomachs; and in certain other hot
places the pigs are red, and this colour has likewise occasionally been
observed in the feral pigs of Jamaica. From these several facts we see that
with pigs when feral there is a strong tendency to revert to the wild type;
but that this tendency is largely governed by the nature of the climate,
amount of exercise, and other causes of change to which they have been
subjected.
The last point worth notice is that we have unusually good evidence of
breeds of pigs now keeping perfectly true, which have been formed by the
crossing of several distinct breeds. The Improved Essex pigs, for instance,
breed very true; but there is no doubt that they largely owe their present
excellent qualities to crosses originally made by Lord Western with the
Neapolitan race, and to subsequent crosses with the Berkshire breed (this
also having been improved by Neapolitan crosses), and likewise, probably,
with the Sussex breed. (3/30. S. Sidney's edition of 'Youatt on the Pig'
1860 pages 7, 26, 27, 29, 30.) In breeds thus formed by complex crosses,
the most careful and unremitting selection during many generations has been
found to be indispensable. Chiefly in consequence of so much crossing, some
well-known breeds have undergone rapid changes; thus, according to
Nathusius (3/31. 'Schweineschadel' s 140.), the Berkshire breed of 1780 is
quite different from that of 1810; and, since this latter period, at least
two distinct forms have borne the same name.
CATTLE.
Domestic cattle are certainly the descendants of more than one wild form,
in the same manner as has been shown to be the case with our dogs and pigs.
Naturalists have generally made two main divisions of cattle: the humped
kinds inhabiting tropical countries, called in India Zebus, to which the
specific name of Bos indicus has been given; and the common non-humped
cattle, generally included under the name of Bos taurus. The humped cattle
were domesticated, as may be seen on the Egyptian monuments, at least as
early as the twelfth dynasty, that is 2100 B.C. They differ from common
cattle in various osteological characters, even in a greater degree,
according to Rutimeyer (3/32. 'Die Fauna der Pfahlbauten' 1861 s. 109, 149,
222. See also Geoffroy Saint-Hilaire in 'Mem. du Mus. d'Hist. Nat.' tome 10
page 172; and his son Isidore in 'Hist. Nat. Gen.' tome 3 page 69. Vasey in
his 'Delineations of the Ox Tribe' 1851 page 127, says the zebu has four,
and common ox five, sacral vertebrae. Mr. Hodgson found the ribs either
thirteen or fourteen in number; see a note in 'Indian Field' 1858 page 62.)
than do the fossil and prehistoric European species, namely, Bos
primigenius and longifrons, from each other. They differ, also, as Mr.
Blyth (3/33. 'The Indian Field' 1858 page 74 where Mr. Blyth gives his
authorities with respect to the feral humped cattle. Pickering also in his
'Races of Man' 1850 page 274 notices the peculiar grunt-like character of
the voice of the humped cattle.), who has particularly attended to this
subject, remarks, in general configuration, in the shape of their ears, in
the point where the dewlap commences, in the typical curvature of their
horns, in their manner of carrying their heads when at rest, in their
ordinary variations of colour, especially in the frequent presence of
"nilgau-like markings on their feet," and "in the one being born with teeth
protruding through the jaws, and the other not so." They have different
habits, and their voice is entirely different. The humped cattle in India
"seldom seek shade, and never go into the water and there stand knee-deep,
like the cattle of Europe." They have run wild in parts of Oude and
Rohilcund, and can maintain themselves in a region infested by tigers. They
have given rise to many races differing greatly in size, in the presence of
one or two humps, in length of horns, and other respects. Mr. Blyth sums up
emphatically that the humped and humpless cattle must be considered as
distinct species. When we consider the number of points in external
structure and habits, independently of important osteological differences,
in which they differ from each other; and that many of these points are not
likely to have been affected by domestication, there can hardly be a doubt,
notwithstanding the adverse opinion of some naturalists, that the humped
and non-humped cattle must be ranked as specifically distinct.
The European breeds of humpless cattle are numerous. Professor Low
enumerates 19 British breeds, only a few of which are identical with those
on the Continent. Even the small Channel islands of Guernsey, Jersey, and
Alderney possess their own sub-breeds (3/34. Mr. H.E. Marquand in 'The
Times' June 23, 1856.); and these again differ from the cattle of the other
British islands, such as Anglesea, and the western isles of Scotland.
Desmarest, who paid attention to the subject, describes 15 French races,
excluding sub-varieties and those imported from other countries. In other
parts of Europe there are several distinct races, such as the pale-coloured
Hungarian cattle, with their light and free step, and enormous horns
sometimes measuring above five feet from tip to tip (3/35. Vasey
'Delineations of the Ox-Tribe' page 124. Brace 'Hungary' 1851 page 94. The
Hungarian cattle descend according to Rutimeyer 'Zahmen Europ. Rindes' 1866
s. 13 from Bos primigenius.): the Podolian cattle also are remarkable from
the height of their fore-quarters. In the most recent work on Cattle (3/36.
Moll and Gayot 'La Connaissance Gen. du Boeuf' Paris 1860. Fig. 82 is that
of the Podolian breed.), engravings are given of fifty-five European
breeds; it is, however, probable that several of these differ very little
from each other, or are merely synonyms. It must not be supposed that
numerous breeds of cattle exist only in long-civilised countries, for we
shall presently see that several kinds are kept by the savages of Southern
Africa.
[With respect to the parentage of the several European breeds, we already
know much from Nilsson's Memoir (3/37. A translation appeared in three
parts in the 'Annals and Mag. of Nat. Hist.' 2nd series volume 4 1849.),
and more especially from Rutimeyer's works and those of Boyd Dawkins. Two
or three species or forms of Bos, closely allied to still living domestic
races, have been found in the more recent tertiary deposits or amongst
prehistoric remains in Europe. Following Rutimeyer, we have:-
Bos primigenius.
This magnificent, well known species was domesticated in Switzerland during
the Neolithic period; even at this early period it varied a little, having
apparently been crossed with other races. Some of the larger races on the
Continent, as the Friesland, etc., and the Pembroke race in England,
closely resemble in essential structure B. primigenius, and no doubt are
its descendants. This is likewise the opinion of Nilsson. Bos primigenius
existed as a wild animal in Caesar's time, and is now semi-wild, though
much degenerated in size, in the park of Chillingham; for I am informed by
Professor Rutimeyer, to whom Lord Tankerville sent a skull, that the
Chillingham cattle are less altered from the true primigenius type than any
other known breed. (3/38. See also Rutimeyer 'Beitrage pal. Gesch. der
Wiederkauer' Basel 1865 s. 54.)
Bos trochoceros.
This form is not included in the three species above mentioned, for it is
now considered by Rutimeyer to be the female of an early domesticated form
of B. primigenius, and as the progenitor of his frontosus race. I may add
that specific names have been given to four other fossil oxen, now believed
to be identical with B. primigenius. (3/39. Pictet 'Palaeontologie' tome 1
page 365 2nd edition. With respect to B. trochoceros see Rutimeyer 'Zahmen
Europ. Rindes' 1866 s. 26.)
Bos longifrons (or brachyceros) of Owen.
This very distinct species was of small size, and had a short body with
fine legs. According to Boyd Dawkins (3/40. W. Boyd Dawkins on the British
Fossil Oxen 'Journal of the Geolog. Soc.' August 1867 page 182. Also 'Proc.
Phil. Soc. of Manchester' November 14, 1871 and 'Cave Hunting' 1875 page
27, 138.) it was introduced as a domesticated animal into Britain at a very
early period, and supplied food to the Roman legionaries. (3/41. 'British
Pleistocene Mammalia' by W.B. Dawkins and W.A. Sandford 1866 page 15.) Some
remains have been found in Ireland in certain crannoges, of which the dates
are believed to be from 843-933 A.D. (3/42. W.R. Wilde 'An Essay on the
Animal Remains, etc. Royal Irish Academy' 1860 page 29. Also 'Proc. of R.
Irish Academy' 1858 page 48.) It was also the commonest form in a
domesticated condition in Switzerland during the earliest part of the
Neolithic period. Professor Owen (3/43. 'Lecture: Royal Institution of G.
Britain' May 2, 1856 page 4. 'British Fossil Mammals' page 513.) thinks it
probable that the Welsh and Highland cattle are descended from this form;
as likewise is the case, according to Rutimeyer, with some of the existing
Swiss breeds. These latter are of different shades of colour from light-
grey to blackish-brown, with a lighter stripe along the spine, but they
have no pure white marks. The cattle of North Wales and the Highlands, on
the other hand, are generally black or dark-coloured.
Bos frontosus of Nilsson.
This species is allied to B. longifrons, and, according to the high
authority of Mr. Boyd Dawkins, is identical with it, but in the opinion of
some judges is distinct. Both co-existed in Scania during the same late
geological period (3/44. Nilsson in 'Annals and Mag. of Nat. Hist.' 1849
volume 4 page 354.), and both have been found in the Irish crannoges.
(3/45. See W.R. Wilde ut supra; and Mr. Blyth in 'Proc. Irish Academy'
March 5, 1864.) Nilsson believes that his B. frontosus may be the parent of
the mountain cattle of Norway, which have a high protuberance on the skull
between the base of the horns. As Professor Owen and others believe that
the Scotch Highland cattle are descended from his B. longifrons, it is
worth notice that a capable judge (3/46. Laing 'Tour in Norway' page 110.)
has remarked that he saw no cattle in Norway like the Highland breed, but
that they more nearly resembled the Devonshire breed.]
On the whole we may conclude, more especially from the researches of Boyd
Dawkins, that European cattle are descended from two species; and there is
no improbability in this fact, for the genus Bos readily yields to
domestication. Besides these two species and the zebu, the yak, the gayal,
and the arni (3/47. Isid. Geoffroy Saint-Hilaire 'Hist. Nat. Gen.' tome 3,
96.) (not to mention the buffalo or genus Bubalus) have been domesticated;
making altogether six species of Bos. The zebu and the two European species
are now extinct in a wild state. Although certain races of cattle were
domesticated at a very ancient period in Europe, it does not follow that
they were first domesticated here. Those who place much reliance on
philology argue that they were imported from the East. (3/48. Idem tome 3
pages 82, 91.) It is probable that they originally inhabited a temperate or
cold climate, but not a land long covered with snow; for our cattle, as we
have seen in the chapter on Horses, have not the instinct of scraping away
the snow to get at the herbage beneath. No one could behold the magnificent
wild bulls on the bleak Falkland Islands in the southern hemisphere, and
doubt about the climate being admirably suited to them. Azara has remarked
that in the temperate regions of La Plata the cows conceive when two years
old, whilst in the much hotter country of Paraguay they do not conceive
till three years old; "from which fact," as he adds, "one may conclude that
cattle do not succeed so well in warm countries." (3/49. 'Quadrupedes du
Paraguay' tome 2 page 360.)
Bos primigenius and longifrons have been ranked by nearly all
palaeontologists as distinct species; and it would not be reasonable to
take a different view simply because their domesticated descendants now
intercross with the utmost freedom. All the European breeds have so often
been crossed both intentionally and unintentionally, that, if any sterility
had ensued from such unions, it would certainly have been detected. As
zebus inhabit a distant and much hotter region, and as they differ in so
many characters from our European cattle, I have taken pains to ascertain
whether the two forms are fertile when crossed. The late Lord Powis
imported some zebus and crossed them with common cattle in Shropshire; and
I was assured by his steward that the cross-bred animals were perfectly
fertile with both parent-stocks. Mr. Blyth informs me that in India
hybrids, with various proportions of either blood, are quite fertile; and
this can hardly fail to be known, for in some districts (3/50. Walther 'Das
Rindvieh' 1817 s. 30.) the two species are allowed to breed freely
together. Most of the cattle which were first introduced into Tasmania were
humped, so that at one time thousands of crossed animals existed there; and
Mr. B. O'Neile Wilson, M.A., writes to me from Tasmania that he has never
heard of any sterility having been observed. He himself formerly possessed
a herd of such crossed cattle, and all were perfectly fertile; so much so,
that he cannot remember even a single cow failing to calve. These several
facts afford an important confirmation of the Pallasian doctrine that the
descendants of species which when first domesticated would if crossed have
been in all probability in some degree sterile, become perfectly fertile
after a long course of domestication. In a future chapter we shall see that
this doctrine throws some light on the difficult subject of Hybridism.
I have alluded to the cattle in Chillingham Park, which, according to
Rutimeyer, have been very little changed from the Bos primigenius type.
This park is so ancient that it is referred to in a record of the year
1220. The cattle in their instincts and habits are truly wild. They are
white, with the inside of the ears reddish-brown, eyes rimmed with black,
muzzles brown, hoofs black, and horns white tipped with black. Within a
period of thirty-three years about a dozen calves were born with "brown and
blue spots upon the cheeks or necks; but these, together with any defective
animals, were always destroyed." According to Bewick, about the year 1770
some calves appeared with black ears; but these were also destroyed by the
keeper, and black ears have not since reappeared. The wild white cattle in
the Duke of Hamilton's park, where I have heard of the birth of a black
calf, are said by Lord Tankerville to be inferior to those at Chillingham.
The cattle kept until the year 1780 by the Duke of Queensberry, but now
extinct, had their ears, muzzle, and orbits of the eyes black. Those which
have existed from time immemorial at Chartley, closely resemble the cattle
at Chillingham, but are larger, "with some small difference in the colour
of the ears." "They frequently tend to become entirely black; and a
singular superstition prevails in the vicinity that, when a black calf is
born, some calamity impends over the noble house of Ferrers. All the black
calves are destroyed." The cattle at Burton Constable in Yorkshire, now
extinct, had ears, muzzle, and the tip of the tail black. Those at
Gisburne, also in Yorkshire, are said by Bewick to have been sometimes
without dark muzzles, with the inside alone of the ears brown; and they are
elsewhere said to have been low in stature and hornless. (3/51. 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. Rutimeyer. The fullest
account of the Chillingham cattle is given by Mr. Hindmarsh, together with
a letter by the late Lord Tankerville, in 'Annals and Mag. of Nat. Hist.'
volume 2 1839 page 274. See Bewick 'Quadrupeds' 2nd edition 1791 page 35
note. With respect to those of the Duke of Queensberry see Pennant 'Tour in
Scotland' page 109. For those of Chartley, see Low 'Domesticated Animals of
Britain' 1845 page 238. For those of Gisburne see Bewick 'Quadrupeds' and
'Encyclop. of Rural Sports' page 101.)
The several above-specified differences in the park-cattle, slight though
they be, are worth recording, as they show that animals living nearly in a
state of nature, and exposed to nearly uniform conditions, if not allowed
to roam freely and to cross with other herds, do not keep as uniform as
truly wild animals. For the preservation of a uniform character, even
within the same park, a certain degree of selection--that is, the
destruction of the dark-coloured calves--is apparently necessary.
Boyd Dawkins believes that the park-cattle are descended from anciently
domesticated, and not truly wild animals; and from the occasional
appearance of dark-coloured calves, it is improbable that the aboriginal
Bos primigenius was white. It is curious what a strong, though not
invariable, tendency there is in wild or escaped cattle to become white
with coloured ears, under widely different conditions of life. If the old
writers Boethius and Leslie (3/52. Boethius was born in 1470; 'Annals and
Mag. of Nat. Hist.' volume 2 1839 page 281; and volume 4 1849 page 424.)
can be trusted, the wild cattle of Scotland were white and furnished with a
great mane; but the colour of their ears is not mentioned. In Wales (3/53.
'Youatt on Cattle' 1834 page 48: See also page 242, on shorthorn cattle.
Bell in his 'British Quadrupeds' page 423 states that, after long attending
to the subject, he has found that white cattle invariably have coloured
ears.), during the tenth century, some of the cattle are described as being
white with red ears. Four hundred cattle thus coloured were sent to King
John; and an early record speaks of a hundred cattle with red ears having
been demanded as a compensation for some offence, but, if the cattle were
of a dark or black colour, 150 were to be presented. The black cattle of
North Wales apparently belong, as we have seen, to the small longifrons
type: and as the alternative was offered of either 150 dark cattle, or 100
white cattle with red ears, we may presume that the latter were the larger
beasts, and probably belonged to the primigenius type. Youatt has remarked
that at the present day, whenever cattle of the shorthorn breed are white,
the extremities of their ears are more or less tinged with red.
The cattle which have run wild on the Pampas, in Texas, and in two parts of
Africa, have become of a nearly uniform dark brownish-red. (3/54. Azara
'Quadrupedes du Paraguay' tome 2 page 361. Azara quotes Buffon for the
feral cattle of Africa. For Texas see 'Times' February 18, 1846.) 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." (3/55. Anson's Voyage. See Kerr and Porter
'Collection' volume 12 page 103.) The Falkland Islands, situated far south,
with all the conditions of life as different as it is possible to conceive
from those of the Ladrones, offer a more interesting case. Cattle have run
wild there during eighty or ninety years; and in the southern districts the
animals are mostly white, with their feet, or whole heads, or only their
ears black; but my informant, Admiral Sulivan (3/56. See also Mr.
Mackinnon's pamphlet on the Falkland Islands, page 24.), who long resided
on these islands, does not believe that they are ever purely white. So that
in these two archipelagos we see that the cattle tend to become white with
coloured ears. In other parts of the Falkland Islands other colours
prevail: near Port Pleasant brown is the common tint; round Mount Usborn,
about half the animals in some of the herds were lead- or mouse-coloured,
which elsewhere is an unusual tint. These latter cattle, though generally
inhabiting high land, breed about a month earlier than the other cattle;
and this circumstance would aid in keeping them distinct and in
perpetuating a peculiar colour. It is worth recalling to mind that blue or
lead-coloured marks have occasionally appeared on the white cattle of
Chillingham. So plainly different were the colours of the wild herds in
different parts of the Falkland Islands, that in hunting them, as Admiral
Sulivan informs me, white spots in one district, and dark spots in another
district, were always looked out for on the distant hills. In the
intermediate districts, intermediate colours prevailed. Whatever the cause
may be, this tendency in the wild cattle of the Falkland Islands, which are
all descended from a few brought from La Plata, to break up into herds of
three different colours, is an interesting fact.
Returning to the several British breeds, the conspicuous difference in
general appearance between Shorthorns, Longhorns (now rarely seen),
Herefords, Highland cattle, Alderneys, etc., must be familiar to every one.
A part of this difference may be attributed to descent from primordially
distinct species; but we may feel sure that there has been a considerable
amount of variation. Even during the Neolithic period, the domestic cattle
were to a certain extent variable. Within recent times most of the breeds
have been modified by careful and methodical selection. How strongly the
characters thus acquired are inherited, may be inferred from the prices
realised by the improved breeds; even at the first sale of Colling's
Shorthorns, eleven bulls reached an average of 214 pounds, and lately
Shorthorn bulls have been sold for a thousand guineas, and have been
exported to all quarters of the world.
Some constitutional differences may be here noticed. The Shorthorns arrive
at maturity far earlier than the wilder breeds, such as those of Wales or
the Highlands. This fact has been shown in an interesting manner by Mr.
Simonds (3/57. 'The Age of the Ox, Sheep, Pig' etc. by Prof. James Simonds,
published by order of the Royal Agricult. Soc.) who has given a table of
the average period of their dentition, which proves that there is a
difference of no less than six months in the appearance of the permanent
incisors. The period of gestation, from observations made by Tessier on
1131 cows, varies to the extent of eighty-one days; and what is more
interesting, M. Lefour affirms "that the period of gestation is longer in
the large German cattle than in the smaller breeds." (3/58. 'Ann. Agricult.
France' April 1837 as quoted in 'The Veterinary' volume 12 page 725. I
quote Tessier's observations from 'Youatt on Cattle' page 527.) With
respect to the period of conception, it seems certain that Alderney and
Zetland cows often become pregnant earlier than other breeds. (3/59. 'The
Veterinary' volume 8 page 681 and volume 10 page 268. Low 'Domest. Animals,
etc.' page 297.) Lastly, as four fully developed mammae is a generic
character in the genus Bos (3/60. Mr. Ogleby in 'Proc. Zoolog. Soc.' 1836
page 138, and 1840 page 4. Quatrefages quotes Philippi 'Revue des Cours
Scientifiques' February 12, 1688 page 657, that the cattle of Piacentino
have thirteen dorsal vertebrae and ribs in the place of the ordinary number
of twelve.), it is worth notice that with our domestic cows the two
rudimentary mammae often become fairly well developed and yield milk.
As numerous breeds are generally found only in long-civilised countries, it
may be well to show that in some countries inhabited by barbarous races,
who are frequently at war with each other, and therefore have little free
communication, several distinct breeds of cattle now exist or formerly
existed. At the Cape of Good Hope Leguat observed, in the year 1720, three
kinds. (3/61. Leguat's Voyage quoted by Vasey in his 'Delineations of the
Ox-tribe' page 132.) 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 (3/62. 'Travels in South Africa'
pages 317, 336.) the Damara, Bechuana, and Namaqua cattle; and he informs
me in a letter that the cattle north of Lake Ngami are likewise different,
as Mr. Galton has heard is also the case with the cattle of Benguela. The
Namaqua cattle in size and shape nearly resemble European cattle, and have
short stout horns and large hoofs. The Damara cattle are very peculiar,
being big-boned, with slender legs, and small hard feet; their tails are
adorned with a tuft of long bushy hair nearly touching the ground, and
their horns are extraordinarily large. The Bechuana cattle have even larger
horns, and there is now a skull in London with the two horns 8 ft. 8 1/4
in. long, as measured in a straight line from tip to tip, and no less than
13 ft. 5 in. as measured along their curvature! Mr. Andersson in his letter
to me says that, though he will not venture to describe the differences
between the breeds belonging to the many different sub-tribes, yet such
certainly exist, as shown by the wonderful facility with which the natives
discriminate them.
That many breeds of cattle have originated through variation, independently
of descent from distinct species, we may infer from what we see in South
America, where the genus Bos was not endemic, and where the cattle which
now exist in such vast numbers are the descendants of a few imported from
Spain and Portugal. In Columbia, Roulin (3/63. 'Mem. de 1'Institut present.
par divers Savans' tome 6 1835 page 333. For Brazil see 'Comptes Rendus'
June 15, 1846. See Azara 'Quadrupedes du Paraguay' tome 2 pages 359, 361.)
describes two peculiar breeds, namely, pelones, with extremely thin and
fine hair, and calongos, absolutely naked. According to Castelnau there are
two races in Brazil, one like European cattle, the other different, with
remarkable horns. In Paraguay, Azara describes a breed which certainly
originated in S. America, called chivos, "because they have straight
vertical horns, conical, and very large at the base." He likewise describes
a dwarf race in Corrientes, with short legs and a body larger than usual.
Cattle without horns, and others with reversed hair, have also originated
in Paraguay.
Another monstrous breed, called niatas or natas, of which I saw two small
herds on the northern bank of the Plata, is so remarkable as to deserve a
fuller description. This breed bears the same relation to other breeds, as
bull or pug dogs do to other dogs, or as improved pigs, according to H. von
Nathusius, do to common pigs. (3/64. 'Schweineschadel' 1864 s. 104.
Nathusius states that the form of skull characteristic in the niata cattle
occasionally appears in European cattle; but he is mistaken, as we shall
hereafter see, in supposing that these cattle do not form a distinct race.
Prof. Wyman, of Cambridge, United States, informs me that the common cod-
fish presents a similar monstrosity, called by the fishermen "bull-dog
cod." Prof. Wyman also concluded, after making numerous inquiries in La
Plata, that the niata cattle transmit their peculiarities or form a race.)
Rutimeyer believes that these cattle belong to the primigenius type. (3/65
'Ueber Art des zahmen Europ. Rindes' 1866 s. 28.) The forehead is very
short and broad, with the nasal end of the skull, together with the whole
plane of the upper molar-teeth, curved upwards. The lower jaw projects
beyond the upper, and has a corresponding upward curvature. It is an
interesting fact that an almost similar confirmation characterizes, as I am
informed by Dr. Falconer, the extinct and gigantic Sivatherium of India,
and is not known in any other ruminant. The upper lip is much drawn back,
the nostrils are seated high up and are widely open, the eyes project
outwards, and the horns are large. In walking the head is carried low, and
the neck is short. The hind legs appear to be longer, compared with the
front legs, than is usual. The exposed incisor teeth, the short head and
upturned nostrils, give these cattle the most ludicrous, self-confident air
of defiance. The skull which I presented to the College of Surgeons has
been thus described by Professor Owen (3/66. 'Descriptive Cat. of Ost.
Collect. of College of Surgeons' 1853 page 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. Rutimeyer.) "It is remarkable from the stunted development
of the nasals, premaxillaries, and fore-part of the lower jaw, which is
unusually curved upwards to come into contact with the premaxillaries. The
nasal bones are about one-third the ordinary length, but retain almost
their normal breadth. The triangular vacuity is left between them, the
frontal and lachrymal, which latter bone articulates with the premaxillary,
and thus excludes the maxillary from any junction with the nasal." So that
even the connexion of some of the bones is changed. Other differences might
be added: thus the plane of the condyles is somewhat modified, and the
terminal edge of the premaxillaries forms an arch. In fact, on comparison
with the skull of a common ox, scarcely a single bone presents the same
exact shape, and the whole skull has a wonderfully different appearance.
The first brief published notice of this race was by Azara, between the
years 1783-96; but Don F. Muniz, of Luxan, who has kindly collected
information for me, states that about 1760 these cattle were kept as
curiosities near Buenos Ayres. Their origin is not positively known, but
they must have originated subsequently to the year 1552, when cattle were
first introduced. Senor Muniz informs me that the breed is believed to have
originated with the Indians southward of the Plata. Even to this day those
reared near the Plata show their less civilised nature in being fiercer
than common cattle, and in the cow, if visited too often, easily deserting
her first calf. The breed is very true, and a niata bull and cow invariably
produce niata calves. The breed has already lasted at least a century. A
niata bull crossed with a common cow, and the reverse cross, yield
offspring having an intermediate character, but with the niata character
strongly displayed. According to Senor Muniz, there is the clearest
evidence, contrary to the common belief of agriculturists in analogous
cases, that the niata cow when crossed with a common bull transmits her
peculiarities more strongly than does the niata bull when crossed with a
common cow. When the pasture is tolerably long, these cattle feed as well
as common cattle with their tongue and palate; but during the great
droughts, when so many animals perish on the Pampas, the niata breed lies
under a great disadvantage, and would, if not attended to, become extinct;
for the common cattle, like horses, are able to keep alive by browsing with
their lips on the twigs of trees and on reeds: this the niatas cannot so
well do, as their lips do not join, and hence they are found to perish
before the common cattle. This strikes me as a good illustration of how
little we are able to judge from the ordinary habits of an animal, on what
circumstances, occurring only at long intervals of time, its rarity or
extinction may depend. It shows us, also, how natural selection would have
determined the rejection of the niata modification had it arisen in a state
of nature.
Having described the semi-monstrous niata breed, I may allude to a white
bull, said to have been brought from Africa, which was exhibited in London
in 1829, and which has been well figured by Mr. Harvey. (3/67. Loudon's
'Magazine of Nat. Hist.' volume 1 1829 page 113. Separate figures are given
of the animal, its hoofs, eye, and dewlap.) It had a hump, and was
furnished with a mane. The dewlap was peculiar, being divided between its
fore-legs into parallel divisions. Its lateral hoofs were annually shed,
and grew to the length of five or six inches. The eye was very peculiar,
being remarkably prominent, and "resembled a cup and ball, thus enabling
the animal to see on all sides with equal ease; the pupil was small and
oval, or rather a parallelogram with the ends cut off, and lying
transversely across the ball." A new and strange breed might probably have
been formed by careful breeding and selection from this animal.
I have often speculated on the probable causes through which each separate
district in Great Britain came to possess in former times its own peculiar
breed of cattle; and the question is, perhaps, even more perplexing in the
case of Southern Africa. We now know that the differences may be in part
attributed to descent from distinct species; but this cause is far from
sufficient. Have the slight differences in climate and in the nature of the
pasture, in the different districts of Britain, directly induced
corresponding differences in the cattle? We have seen that the semi-wild
cattle in the several British parks are not identical in colouring or size,
and that some degree of selection has been requisite to keep them true. It
is almost certain that abundant food given during many generations directly
affects the size of a breed. (3/68. Low 'Domesticated Animals of the
British Isles' page 264.) That climate directly affects the thickness of
the skin and the hair is likewise certain: thus Roulin asserts (69 'Mem. de
l'Institut present. Par divers Savans' tome 6 1835 page 332.) that the
hides of the feral cattle on the hot Llanos "are always much less heavy
than those of the cattle raised on the high platform of Bogota; and that
these hides yield in weight and in thickness of hair to those of the cattle
which have run wild on the lofty Paramos." The same difference has been
observed in the hides of the cattle reared on the bleak Falkland Islands
and on the temperate Pampas. Low has remarked (3/70. Idem pages 304, 368
etc.) that the cattle which inhabit the more humid parts of Britain have
longer hair and thicker skins than other British cattle. When we compare
highly improved stall-fed cattle with the wilder breeds, or compare
mountain and lowland breeds, we cannot doubt that an active life, leading
to the free use of the limbs and lungs, affects the shape and proportions
of the whole body. It is probable that some breeds, such as the semi-
monstrous niata cattle, and some peculiarities, such as being hornless,
etc., have appeared suddenly owing to what we may call in our ignorance
spontaneous variation; but even in this case a rude kind of selection is
necessary, and the animals thus characterised must be at least partially
separated from others. This degree of care, however, has sometimes been
taken even in little-civilised districts, where we should least have
expected it, as in the case of the niata, chivo, and hornless cattle in S.
America.
That methodical selection has done wonders within a recent period in
modifying our cattle, no one doubts. During the process of methodical
selection it has occasionally happened that deviations of structure, more
strongly pronounced than mere individual differences, yet by no means
deserving to be called monstrosities, have been taken advantage of: thus
the famous Longhorn Bull, Shakespeare, though of the pure Canley stock,
"scarcely inherited a single point of the long-horned breed, his horns
excepted (3/71. 'Youatt on Cattle' page 193. A full account of this bull is
taken from Marshall.); yet in the hands of Mr. Fowler, this bull greatly
improved his race. We have also reason to believe that selection, carried
on so far unconsciously that there was at no one time any distinct
intention to improve or change the breed, has in the course of time
modified most of our cattle; for by this process, aided by more abundant
food, all the lowland British breeds have increased greatly in size and in
early maturity since the reign of Henry VII. (3/72. 'Youatt on Cattle' page
116. Lord Spencer has written on this same subject.) It should never be
forgotten that many animals have to be annually slaughtered; so that each
owner must determine which shall be killed and which preserved for
breeding. In every district, as Youatt has remarked, there is a prejudice
in favour of the native breed; so that animals possessing qualities,
whatever they may be, which are most valued in each district, will be
oftenest preserved; and this unmethodical selection assuredly will in the
long run affect the character of the whole breed. But it may be asked, can
this rude kind of selection have been practised by barbarians such as those
of southern Africa? In a future chapter on Selection we shall see that this
has certainly occurred to some extent. Therefore, looking to the origin of
the many breeds of cattle which formerly inhabited the several districts of
Britain, I conclude that, although slight differences in the nature of the
climate, food, etc., as well as changed habits of life, aided by
correlation of growth, and the occasional appearance from unknown causes of
considerable deviations of structure, have all probably played their parts;
yet that the occasional preservation in each district of those individual
animals which were most valued by each owner has perhaps been even more
effective in the production of the several British breeds. As soon as two
or more breeds were formed in any district, or when new breeds descended
from distinct species were introduced, their crossing, especially if aided
by some selection, will have multiplied the number and modified the
characters of the older breeds.
SHEEP.
I shall treat this subject briefly. Most authors look at our domestic sheep
as descended from several distinct species. Mr. Blyth, who has carefully
attended to the subject, believes that fourteen wild species now exist, but
"that not one of them can be identified as the progenitor of any one of the
interminable domestic races." M. Gervais thinks that there are six species
of Ovis (3/73. Blyth on the genus Ovis in 'Annals and Mag. of Nat. History'
volume 7 1841 page 261. With respect to the parentage of the breeds see Mr.
Blyth's excellent articles in 'Land and Water' 1867 pages 134, 156. Gervais
'Hist. Nat. des Mammiferes' 1855 tome 2 page 191.) but that our domestic
sheep form a distinct genus, now completely extinct. A German naturalist
(3/74. Dr. L. Fitzinger 'Ueber die Racen des Zahmen Schafes' 1860 s. 86.)
believes that our sheep descend from ten aboriginally distinct species, of
which only one is still living in a wild state! Another ingenious observer
(3/75. J. Anderson 'Recreations in Agriculture and Natural History' volume
2 page 264.), though not a naturalist, with a bold defiance of everything
known on geographical distribution, infers that the sheep of Great Britain
alone are the descendants of eleven endemic British forms! Under such a
hopeless state of doubt it would be useless for my purpose to give a
detailed account of the several breeds; but a few remarks may be added.
Sheep have been domesticated from a very ancient period. Rutimeyer (3/76.
'Pfahlbauten' s. 127, 193.) found in the Swiss lake-dwellings the remains
of a small breed, with thin tall legs, and horns like those of a goat, thus
differing somewhat from any kind now known. Almost every country has its
own peculiar breed; and many countries have several breeds differing
greatly from each other. One of the most strongly marked races is an
Eastern one with a long tail, including, according to Pallas, twenty
vertebrae, and so loaded with fat that it is sometimes placed on a truck,
which is dragged about by the living animal. These sheep, though ranked by
Fitzinger as a distinct aboriginal form, bear in their drooping ears the
stamp of long domestication. This is likewise the case with those sheep
which have two great masses of fat on the rump, with the tail in a
rudimentary condition. The Angola variety of the long-tailed race has
curious masses of fat on the back of the head and beneath the jaws. (3/77.
'Youatt on Sheep' page 120.) Mr. Hodgson in an admirable paper (3/78.
'Journal of the Asiatic Soc. of Bengal' volume 16 pages 1007, 1016.) on the
sheep of the Himalaya infers from the distribution of the several races,
"that this caudal augmentation in most of its phases is an instance of
degeneracy in these pre-eminently Alpine animals." The horns present an
endless diversity in character; being not rarely absent, especially in the
female sex, or, on the other hand, amounting to four or even eight in
number. The horns, when numerous, arise from a crest on the frontal bone,
which is elevated in a peculiar manner. It is remarkable that multiplicity
of horns "is generally accompanied by great length and coarseness of the
fleece." (3/79. 'Youatt on Sheep' pages 142-169.) This correlation,
however, is far from being general; for instance, I am informed by Mr. D.
Forbes, that the Spanish sheep in Chile resemble, in fleece and in all
other characters, their parent merino-race, except that instead of a pair
they generally bear four horns. The existence of a pair of mammae 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 Cagias (a sub-Himalayan domestic race) possessed of four
teats." (3/80. 'Journal Asiat. Soc. of Bengal' volume 16 1847 page 1015.)
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 (3/81. 'Hist. Nat. Gen.' tome 3 page 435.) has shown that these
pits or pouches are absent in some breeds.
In sheep there is a strong tendency for characters, which have apparently
been acquired under domestication, to become attached either exclusively to
the male sex, or to be more highly developed in this than in the other sex.
Thus in many breeds the horns are deficient in the ewe, though this
likewise occurs occasionally with the female of the wild musmon. In the
rams of the Wallachian breed, "the horns spring almost perpendicularly from
the frontal bone, and then take a beautiful spiral form; in the ewes they
protrude nearly at right angles from the head, and then become twisted in a
singular manner." (3/82. 'Youatt on Sheep' page 138.) Mr. Hodgson states
that the extraordinarily arched nose or chaffron, which is so highly
developed in several foreign breeds, is characteristic of the ram alone,
and apparently is the result of domestication. (3/83. 'Journal Asiat. Soc.
of Bengal' volume 16 1847 pages 1015, 1016.) 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 (3/84. 'Racen des
Zahmen Schafes' s. 77.) remarks that the mane in the African maned race is
far more developed in the ram than in the ewe.
Different races of sheep, like cattle, present constitutional differences.
Thus the improved breeds arrive at maturity at an early age, as has been
well shown by Mr. Simonds through their early average period of dentition.
The several races have become adapted to different kinds of pasture and
climate: for instance, no one can rear Leicester sheep on mountainous
regions, where Cheviots flourish. As Youatt has remarked, "In all the
different districts of Great Britain we find various breeds of sheep
beautifully adapted to the locality which they occupy. No one knows their
origin; they are indigenous to the soil, climate, pasturage, and the
locality on which they graze; they seem to have been formed for it and by
it." (3/85. 'Rural Economy of Norfolk' volume 2 page 136.) Marshall relates
(3/86. 'Youatt on Sheep' page 312. On same subject, see excellent remarks
in 'Gardener's Chronicle' 1858 page 868. For experiments in crossing
Cheviot sheep with Leicesters see Youatt page 325.) 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." (3/87.
'Youatt on Sheep' note page 491.) There is very good evidence that English
breeds of sheep will not succeed in France. (3/88. M. Malingie-Nouel
'Journal R. Agricult. Soc.' volume 14 1853 page 214 translated and
therefore approved by a great authority, Mr. Pusey.) 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 (3/89. 'The Veterinary' volume 10 page 217.)
that the owner could not keep them; the coarser-skinned sheep never being
affected.
The period of gestation was formerly thought to be of so unalterable a
character, that a supposed difference of this kind between the wolf and the
dog was esteemed a sure sign of specific distinction; but we have seen that
the period is shorter in the improved breeds of the pig, and in the larger
breeds of the ox, than in other breeds of these two animals. And now we
know, on the excellent authority of Hermann von Nathusius (3/90. A
translation of his paper is given in 'Bull. Soc. Imp. d'Acclimat.' tome 9
1862 page 723.), that Merino and Southdown sheep, when both have long been
kept under exactly the same conditions, differ in their average period of
gestation, as is seen in the following Table
Merinos 150.3 days.
Southdowns 144.2 "
Half-bred Merinos and Southdowns 146.3 "
3/4 blood of Southdown 145.5 "
7/8 blood of Southdown 144.2 "
In this graduated difference in cross-bred animals having different
proportions of Southdown blood, we see how strictly the two periods of
gestation have been transmitted. Nathusius remarks that, as Southdowns grow
with remarkable rapidity after birth, it is not surprising that their
foetal development should have been shortened. It is of course possible
that the difference in these two breeds may be due to their descent from
distinct parent-species; but as the early maturity of the Southdowns has
long been carefully attended to by breeders, the difference is more
probably the result of such attention. Lastly, the fecundity of the several
breeds differs much; some generally producing twins or even triplets at a
birth, of which fact the curious Shangai sheep (with their truncated and
rudimentary ears, and great Roman noses), lately exhibited in the
Zoological Gardens, offer a remarkable instance.
Sheep are perhaps more readily affected by the direct action of the
conditions of life to which they have been exposed than almost any other
domestic animal. According to Pallas, and more recently according to Erman,
the fat-tailed Kirghisian sheep, when bred for a few generations in Russia,
degenerate, and the mass of fat dwindles away, "the scanty and bitter
herbage of the steppes seems so essential to their development." Pallas
makes an analogous statement with respect to one of the Crimean breeds.
Burnes states that the Karakool breed, which produces a fine, curled,
black, and valuable fleece, when removed from its own canton near Bokhara
to Persia or to other quarters, loses its peculiar fleece. (3/91. Erman
'Travels in Siberia' English translation volume 1 page 228. For Pallas on
the fat-tailed sheep I quote from Anderson's account of the 'Sheep of
Russia' 1794 page 34. With respect to the Crimean sheep see Pallas
'Travels' English translation volume 2 page 454. For the Karakool sheep see
Burnes 'Travels in Bokhara' volume 3 page 151.) In all such cases, however,
it may be that a change of any kind in the conditions of life causes
variability and consequent loss of character, and not that certain
conditions are necessary for the development of certain characters.
Great heat, however, seems to act directly on the fleece: several accounts
have been published of the change which sheep imported from Europe undergo
in the West Indies. Dr. Nicholson of Antigua informs me that, after the
third generation, the wool disappears from the whole body, except over the
loins; and the animal then appears like a goat with a dirty door-mat on its
back. A similar change is said to take place on the west coast of Africa.
(3/92. See Report of the Directors of the Sierra Leone Company as quoted in
White 'Gradation of Man' page 95. With respect to the change which sheep
undergo in the West Indies see also Dr. Davy in 'Edin. New. Phil. Journal'
January 1852. For the statement made by Roulin see 'Mem. de l'Institut
present. par divers Savans.' tome 6 1835 page 347.) On the other hand, many
wool-bearing sheep live on the hot plains of India. Roulin asserts that in
the lower and heated valleys of the Cordillera, if the lambs are sheared as
soon as the wool has grown to a certain thickness, all goes on afterwards
as usual; but if not sheared, the wool detaches itself in flakes, and short
shining hair like that on a goat is produced ever afterwards. This curious
result seems merely to be an exaggerated tendency natural to the Merino
breed, for as a great authority, namely, Lord Somerville, remarks, "the
wool of our Merino sheep after shear-time is hard and coarse to such a
degree as to render it almost impossible to suppose that the same animal
could bear wool so opposite in quality, compared to that which has been
clipped from it: as the cold weather advances, the fleeces recover their
soft quality." As in sheep of all breeds the fleece naturally consists of
longer and coarser hair covering shorter and softer wool, the change which
it often undergoes in hot climates is probably merely a case of unequal
development; for even with those sheep which like goats are covered with
hair, a small quantity of underlying wool may always be found. (3/93.
'Youatt on Sheep' page 69 where Lord Somerville is quoted. See page 117 on
the presence of wool under the hair. With respect to the fleeces of
Australian sheep page 185. On selection counteracting any tendency to
change see pages 70, 117, 120, 168.) In the wild mountain-sheep (0vis
montana) of North America there is an analogous annual change of coat; "the
wool begins to drop out in early spring, leaving in its place a coat of
hair resembling that of the elk, a change of pelage quite different in
character from the ordinary thickening of the coat or hair, common to all
furred animals in winter,--for instance, in the horse, the cow, etc., which
shed their winter coat in the spring." (3/94. Audubon and Bachman 'The
Quadrupeds of North America' 1846 volume 5 page 365.)
A slight difference in climate or pasture sometimes slightly affects the
fleece, as has been observed even in different districts in England, and is
well shown by the great softness of the wool brought from Southern
Australia. But it should be observed, as Youatt repeatedly insists, that
the tendency to change may generally be counteracted by careful selection.
M. Lasterye, after discussing this subject, sums up as follows: "The
preservation of the Merino race in its utmost purity at the Cape of Good
Hope, in the marshes of Holland, and under the rigorous climate of Sweden,
furnishes an additional support of this my unalterable principle, that
fine-woolled sheep may be kept wherever industrious men and intelligent
breeders exist."
That methodical selection has effected great changes in several breeds of
sheep no one who knows anything on the subject, entertains a doubt. The
case of the Southdowns, as improved by Ellman, offers perhaps the most
striking instance. Unconscious or occasional selection has likewise slowly
produced a great effect, as we shall see in the chapters on Selection. That
crossing has largely modified some breeds, no one who will study what has
been written on this subject--for instance, Mr. Spooner's paper--will
dispute; but to produce uniformity in a crossed breed, careful selection
and "rigorous weeding," as this author expresses it, are indispensable.
(3/95. 'Journal of R. Agricult. Soc. of England' volume 20 part 2, W.C.
Spooner on cross-Breeding.)
In some few instances new breeds have suddenly originated; thus, in 1791, a
ram-lamb was born in Massachusetts, having short crooked legs and a long
back, like a turnspit-dog. From this one lamb the otter or ancon semi-
monstrous breed was raised; as these sheep could not leap over the fences,
it was thought that they would be valuable; but they have been supplanted
by merinos, and thus exterminated. The sheep are remarkable from
transmitting their character so truly that Colonel Humphreys (3/96.
'Philosoph. Transactions' London 1813 page 88.) never heard of "but one
questionable case" of an ancon ram and ewe not producing ancon offspring.
When they are crossed with other breeds the offspring, with rare
exceptions, instead of being intermediate in character, perfectly resemble
either parent; even one of twins has resembled one parent and the second
the other. Lastly, "the ancons have been observed to keep together,
separating themselves from the rest of the flock when put into enclosures
with other sheep."
A more interesting case has been recorded in the Report of the Juries for
the Great Exhibition (1851), namely, the production of a merino ram-lamb on
the Mauchamp farm, in 1828, which was remarkable for its long, smooth,
straight, and silky wool. By the year 1833 M. Graux had raised rams enough
to serve his whole flock, and after a few more years he was able to sell
stock of his new breed. So peculiar and valuable is the wool, that it sells
at 25 per cent above the best merino wool: even the fleeces of half-bred
animals are valuable, and are known in France as the "Mauchamp-merino." It
is interesting, as showing how generally any marked deviation of structure
is accompanied by other deviations, that the first ram and his immediate
offspring were of small size, with large heads, long necks, narrow chests,
and long flanks; but these blemishes were removed by judicious crosses and
selection. The long smooth wool was also correlated with smooth horns; and
as horns and hair are homologous structures, we can understand the meaning
of this correlation. If the Mauchamp and ancon breeds had originated a
century or two ago, we should have had no record of their birth; and many a
naturalist would no doubt have insisted, especially in the case of the
Mauchamp race, that they had each descended from, or been crossed with,
some unknown aboriginal form.
GOATS.
From the recent researches of M. Brandt, most naturalists now believe that
all our goats are descended from the Capra aegagrus of the mountains of
Asia, possibly mingled with the allied Indian species C. falconeri of
India. (3/97. Isidore Geoffroy St. Hilaire 'Hist. Nat. Generale' tome 3
page 87. Mr. Blyth 'Land and Water' 1867 page 37 has arrived at a similar
conclusion, but he thinks that certain Eastern races may perhaps be in part
descended from the Asiatic markhor.) In Switzerland, during the neolithic
period, the domestic goat was commoner than the sheep; and this very
ancient race differed in no respect from that now common in Switzerland.
(3/98. Rutimeyer 'Pfahlbauten' s. 127.) 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 (3/99. Godron 'De l'Espece'
tome 1 page 402.) they are all quite fertile when crossed. So numerous are
the breeds, that Mr. G. Clark (3/100. 'Annals and Mag. of Nat History' 2nd
series volume 2 1848 page 363.) has described eight distinct kinds imported
into the one island of Mauritius. The ears of one kind were enormously
developed, being, as measured by Mr. Clark, no less than 19 inches in
length and 4 3/4 inches in breadth. As with cattle, the mammae 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 (3/101. 'De l'Espece' tome 1 page 406. Mr. Clark also
refers to differences in the shape of the mammae. 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.), the
mammae differ greatly in shape in different breeds, being elongated in the
common goat, hemispherical in the Angora race, and bilobed and divergent in
the goats of Syria and Nubia. According to this same author, the males of
certain breeds have lost their usual offensive odour. In one of the Indian
breeds the males and females have horns of widely-different shapes (3/102.
Mr. Clark 'Annals and Mag. of Nat. Hist.' 2nd series volume 2 1848 page
361.); and in some breeds the females are destitute of horns. (3/103.
Desmarest 'Encyclop. Method. Mammalogie' page 480.) M. Ramu of Nancy
informs me that many of the goats there bear on the upper part of the
throat a pair of hairy appendages, 70 mm. in length and about 10 mm. in
diameter, which in external appearance resemble those above described on
the jaws of pigs. The presence of inter-digital pits or glands on all four
feet has been thought to characterise the genus Ovis, and their absence to
be characteristic of the genus Capra; but Mr. Hodgson has found that they
exist in the front feet of the majority of Himalayan goats. (3/104.
'Journal of Asiatic Soc. of Bengal' volume 16 1847 pages 1020, 1025.) Mr.
Hodgson measured the intestines in two goats of the Dugu race, and he found
that the proportional length of the great and small intestines differed
considerably. In one of these goats the caecum was thirteen inches, and in
the other no less than thirty-six inches in length!
CHAPTER 1.IV.
DOMESTIC RABBITS.
DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RABBIT.
ANCIENT DOMESTICATION.
ANCIENT SELECTION.
LARGE LOP-EARED RABBITS.
VARIOUS BREEDS.
FLUCTUATING CHARACTERS.
ORIGIN OF THE HIMALAYAN BREED.
CURIOUS CASE OF INHERITANCE.
FERAL RABBITS IN JAMAICA AND THE FALKLAND ISLANDS.
PORTO SANTO FERAL RABBITS.
OSTEOLOGICAL CHARACTERS.
SKULL.
SKULL OF HALF-LOP RABBITS.
VARIATIONS IN THE SKULL ANALOGOUS TO DIFFERENCES IN DIFFERENT SPECIES OF
HARES.
VERTEBRAE.
STERNUM.
SCAPULA.
EFFECTS OF USE AND DISUSE ON THE PROPORTIONS OF THE LIMBS AND BODY.
CAPACITY OF THE SKULL AND REDUCED SIZE OF THE BRAIN.
SUMMARY ON THE MODIFICATIONS OF DOMESTICATED RABBITS.
All naturalists, with, as far as I know, a single exception, believe that
the several domestic breeds of the rabbit are descended from the common
wild species; I shall therefore describe them more carefully than in the
previous cases. Professor Gervais (4/1. M.P. Gervais 'Hist. Nat. des
Mammiferes' 1854. tome 1 page 288.) states "that the true wild rabbit is
smaller than the domestic; its proportions are not absolutely the same; its
tail is smaller; its ears are shorter and more thickly clothed with hair;
and these characters, without speaking of colour, are so many indications
opposed to the opinion which unites these animals under the same specific
denomination." Few naturalists will agree with this author that such slight
differences are sufficient to separate as distinct species the wild and
domestic rabbit. How extraordinary it would be, if close confinement,
perfect tameness, unnatural food, and careful breeding, all prolonged
during many generations, had not produced at least some effect! The tame
rabbit has been domesticated from an ancient period. Confucius ranges
rabbits among animals worthy to be sacrificed to the gods, and, as he
prescribes their multiplication, they were probably at this early period
domesticated in China. They are mentioned by several of the classical
writers. In 1631 Gervaise Markham writes, "You shall not, as in other
cattell, looke to their shape, but to their richnesse, onely elect your
buckes, the largest and goodliest conies you can get; and for the richnesse
of the skin, that is accounted the richest which hath the equallest mixture
of blacke and white haire together, yet the blacke rather shadowing the
white; the furre should be thicke, deepe, smooth, and shining;...they are
of body much fatter and larger, and, when another skin is worth two or
three pence, they are worth two shillings." From this full description we
see that silver-grey rabbits existed in England at this period; and what is
far more important, we see that the breeding or selection of rabbits was
then carefully attended to. Aldrovandi, in 1637, describes, on the
authority of several old writers (as Scaliger, in 1557), rabbits of various
colours, some "like a hare," and he adds that P. Valerianus (who died a
very old man in 1558) saw at Verona rabbits four times bigger than ours.
(4/2. U. Aldrovandi 'De Quadrupedibus digitatis' 1637 page 383. For
Confucius and G. Markham see a writer who has studied the subject in
'Cottage Gardener' January 22, 1861 page 250.)
From the fact of the rabbit having been domesticated at an ancient period,
we must look to the northern hemisphere of the Old World, and to the warmer
temperate regions alone, for the aboriginal parent-form; for the rabbit
cannot live without protection in countries as cold as Sweden, and, though
it has run wild in the tropical island of Jamaica, it has never greatly
multiplied there. It now exists, and has long existed, in the warmer
temperate parts of Europe, for fossil remains have been found in several
countries. (4/3. Owen 'British Fossil Mammals' page 212.) The domestic
rabbit readily becomes feral in these same countries, and when variously
coloured kinds are turned out they generally revert to the ordinary grey
colour. (4/4. Bechstein 'Naturgesch. Deutschlands' 1801 b. 1 page 1133. I
have received similar accounts with respect to England and Scotland.) Wild
rabbits, if taken young, can be domesticated, though the process is
generally very troublesome. (4/5. 'Pigeons and Rabbits' by E.S. Delamer
1854 page 133. Sir J. Sebright 'Observations on Instinct' 1836 page 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' tome 2 page 368.) The various domestic races are often
crossed, and are believed to be quite fertile together, and a perfect
gradation can be shown to exist from the largest domestic kinds, having
enormously developed ears, to the common wild kind. The parent-form must
have been a burrowing animal, a habit not common, as far as I can discover,
to any other species in the large genus Lepus. Only one wild species is
known with certainty to exist in Europe; but the rabbit (if it be a true
rabbit) from Mount Sinai, and likewise that from Algeria, present slight
differences; and these forms have been considered by some authors as
specifically distinct. (4/6. Gervais 'Hist. Nat. des Mammiferes' tome 1
page 292.) But such slight differences would aid us little in explaining
the more considerable differences characteristic of the several domestic
races. If the latter are the descendants of two or more closely allied
species, these, with the exception of the common rabbit, have been
exterminated in a wild state; and this is very improbable, seeing with what
pertinacity this animal holds its ground. From these several reasons we may
infer with safety that all the domestic breeds are the descendants of the
common wild species. But from what we hear of the marvellous success in
France in rearing hybrids between the hare and rabbit (4/7. See Dr. P.
Broca's interesting memoir on this subject in Brown-Sequard 'Journ. de.
Phys.' volume 2 page 367.), it is possible, though not probable, from the
great difficulty in making the first cross, that some of the larger races,
which are coloured like the hare, may have been modified by crosses with
this animal. Nevertheless, the chief differences in the skeletons of the
several domestic breeds cannot, as we shall presently see, have been
derived from a cross with the hare.
There are many breeds which transmit their characters more or less truly.
Every one has seen the enormous lop-eared rabbits exhibited at our shows;
various allied sub-breeds are reared on the Continent, such as the so-
called Andalusian, which is said to have a large head with a round
forehead, and to attain a greater size than any other kind; another large
Paris breed is named the Rouennais, and has a square head; the so-called
Patagonian rabbit has remarkably short ears and a large round head.
Although I have not seen all these breeds, I feel some doubt about there
being any marked difference in the shape of their skulls. (4/8. The skulls
of these breeds are briefly described in the 'Journal of Horticulture' May
7, 1861 page 108.) English lop-eared rabbits often weigh 8 pounds or 10
pounds, and one has been exhibited weighing 18 pounds; whereas a full-sized
wild rabbit weighs only about 3 1/4 pounds. The head or skull in all the
large lop-eared rabbits examined by me is much longer relatively to its
breadth than in the wild rabbit. Many of them have loose transverse folds
of skin or dewlaps beneath the throat, which can be pulled out so as to
reach nearly to the ends of the jaws. Their ears are prodigiously
developed, and hang down on each side of their faces. A rabbit was
exhibited in 1867 with its two ears, measured from the tip of one to the
tip of the other, 22 inches in length, and each ear 5 3/8 inches in
breadth. In 1869 one was exhibited with ears, measured in the same manner,
23 1/8 in length and 5 1/2 in breadth; "thus exceeding any rabbit ever
exhibited at a prize show." In a common wild rabbit I found that the length
of two ears, from tip to tip, was 7 5/8 inches, and the breadth only 1 7/8
inch. The weight of body in the larger rabbits, and the development of
their ears, are the qualities which win prizes, and have been carefully
selected.
The hare-coloured, or, as it is sometimes called, the Belgian rabbit,
differs in nothing except colour from the other large breeds; but Mr. J.
Young, of Southampton, a great breeder of this kind, informs me that the
females, in all the specimens examined by him, had only six mammae; and
this certainly was the case with two females which came into my possession.
Mr. B.P. Brent, however, assures me that the number is variable with other
domestic rabbits. The common wild rabbit always has ten mammae. 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. (4/9. 'Journal of Horticulture' 1861 page 380.) Two live rabbits
were brought to me from Moscow, of about the size of the wild species, but
with long soft fur, different from that of the Angora. These Moscow rabbits
had pink eyes and were snow-white, excepting the ears, two spots near the
nose, the upper and under surface of the tail, and the hinder tarsi, which
were blackish-brown. In short, they were coloured nearly like the so-called
Himalayan rabbits, presently to be described, and differed from them only
in the character of their fur. There are two other breeds which come true
to colour, but differ in no other respect, namely silver-greys and
chinchillas. Lastly, the Nicard or Dutch rabbit may be mentioned, which
varies in colour, and is remarkable from its small size, some specimens
weighing only 1 1/4 pounds; rabbits of this breed make excellent nurses for
other and more delicate kinds. (4/10. 'Journal of Horticulture' May 28,
1861 page 169.)
(FIGURE 5. HALF-LOP RABBIT. (Copied from E.S. Delamer's work.)
Certain characters are remarkably fluctuating, or are very feebly
transmitted by domestic rabbits: thus, one breeder tells me that with the
smaller kinds he has hardly ever raised a whole litter of the same colour:
with the large lop-eared breeds "it is impossible," says a great judge
(4/11. 'Journal of Horticulture' 1861 page 327. With respect to the ears
see Delamer on 'Pigeons and Rabbits' 1854 page 141; also 'Poultry
Chronicle' volume 2 page 499 and ditto for 1854 page 586.), "to breed true
to colour, but by judicious crossing a great deal may be done towards it.
The fancier should know how his does are bred, that is, the colour of their
parents." Nevertheless, certain colours, as we shall presently see, are
transmitted truly. The dewlap is not strictly inherited. Lop-eared rabbits,
with their ears hanging down flat on each side of the face, do not transmit
this character at all truly. Mr. Delamer remarks that, "with fancy rabbits,
when both the parents are perfectly formed, have model ears, and are
handsomely marked, their progeny do not invariably turn out the same." When
one parent, or even both, are oar-laps, that is, have their ears sticking
out at right angles, or when one parent or both are half-lops, that is,
have only one ear dependent, there is nearly as good a chance of the
progeny having both ears full-lop, as if both parents had been thus
characterised. But I am informed, if both parents have upright ears, there
is hardly a chance of a full-lop. In some half-lops the ear that hangs down
is broader and longer than the upright ear (4/12. Delamer 'Pigeons and
Rabbits' page 136. See also 'Journal of Horticulture' 1861 page 375.); so
that we have the unusual case of a want of symmetry on the two sides. This
difference in the position and size of the two ears probably indicates that
the lopping results from the great length and weight of the ear, favoured
no doubt by the weakness of the muscles consequent on disuse. Anderson
(4/13. 'An Account of the different Kinds of Sheep in the Russian
Dominions' 1794 page 39.) mentions a breed having only a single ear; and
Professor Gervais another breed destitute of ears.
We come now to the Himalayan breed, which is sometimes called Chinese,
Polish, or Russian. These pretty rabbits are white, or occasionally yellow,
excepting their ears, nose, feet, and the upper side of the tail, which are
all brownish-black; but as they have red eyes, they may be considered as
albinoes. I have received several accounts of their breeding perfectly
true. From their symmetrical marks, they were at first ranked as
specifically distinct, and were provisionally named L. nigripes. (4/14.
'Proc. Zoolog. Soc.' June 23, 1857 page 159.) Some good observers thought
that they could detect a difference in their habits, and stoutly maintained
that they formed a new species. The origin of this breed is so curious,
both in itself and as throwing some light on the complex laws of
inheritance that it is worth giving in detail. But it is first necessary
briefly to describe two other breeds: silver-greys or silver-sprigs
generally have black heads and legs, and their fine grey fur is
interspersed with numerous black and white long hairs. They breed perfectly
true, and have long been kept in warrens. When they escape and cross with
common rabbits, the product, as I hear from Mr. Wyrley Birch, of Wretham
Hall, is not a mixture of the two colours, but about half take after the
one parent, and the other half after the other parent. Secondly,
chinchillas or tame silver-greys (I will use the former name) have short,
paler, mouse or slate-coloured fur, interspersed with long, blackish,
slate-coloured, and white hairs. (4/15. 'Journal of Horticulture' April 9,
1861 page 35.) These rabbits breed perfectly true. A writer stated in 1857
(4/16. 'Cottage Gardener' 1857 page 141.) that he had produced Himalayan
rabbits in the following manner. He had a breed of chinchillas which had
been crossed with the common black rabbit, and their offspring were either
blacks or chinchillas. These latter were again crossed with other
chinchillas (which had also been crossed with silver-greys), and from this
complicated cross Himalayan rabbits were raised. From these and other
similar statements, Mr. Bartlett (4/17. Mr. Bartlett in 'Proc. Zoolog Soc.'
1861 page 40.) was led to make a careful trial in the Zoological Gardens,
and he found that by simply crossing silver-greys with chinchillas he could
always produce some few Himalayans; and the latter, notwithstanding their
sudden origin, if kept separate, bred perfectly true. But I have recently
been assured the pure silver-greys of any sub-breed occasionally produce
Himalayans.
The Himalayans, when first born, are quite white, and are then true
albinoes; but in the course of a few months they gradually assume their
dark ears, nose, feet, and tail. Occasionally, however, as I am informed by
Mr. W.A. Wooler and the Rev. W.D. Fox, the young are born of a very pale
grey colour, and specimens of such fur were sent me by the former
gentleman. The grey tint, however, disappears as the animal comes to
maturity. So that with these Himalayans there is a tendency, strictly
confined to early youth, to revert to the colour of the adult silver-grey
parent-stock. Silver-greys and chinchillas, on the other hand, present a
remarkable contrast with the Himalayans in their colour whilst quite young,
for they are born perfectly black, but soon assume their characteristic
grey or silver tints. The same thing occurs with grey horses, which, as
long as they are foals, are generally of a nearly black colour, but soon
become grey, and get whiter and whiter as they grow older. Hence the usual
rule is that Himalayans are born white and afterwards become in certain
parts of their bodies dark-coloured; whilst silver-greys are born black and
afterwards become sprinkled with white. Exceptions, however, and of a
directly opposite nature, occasionally occur in both cases. For young
silver-greys are sometimes born in warrens, as I hear from Mr. W. Birch, of
a cream-colour, but these young animals ultimately become black. The
Himalayans, on the other hand, sometimes produce, as is stated by an
experienced amateur (4/18. 'Phenomenon in Himalayan Rabbits' in 'Journal of
Horticulture' January 27, 1865 page 102.), a single black young one in a
litter; and this, before two months elapse, becomes perfectly white.
To sum up the whole curious case: wild silver-greys may be considered as
black rabbits which become grey at an early period of life. When they are
crossed with common rabbits, the offspring are said not to have blended
colours, but to take after either parent; and in this respect they resemble
black and albino varieties of most quadrupeds, which often transmit their
colours in this same manner. When they are crossed with chinchillas, that
is, with a paler sub-variety, the young are at first pure albinoes, but
soon become dark-coloured in certain parts of their bodies, and are then
called Himalayans. The young Himalayans, however, are sometimes at first
either pale grey or completely black, in either case changing after a time
to white. In a future chapter I shall advance a large body of facts showing
that, when two varieties are crossed both of which differ in colour from
their parent-stock, there is a strong tendency in the young to revert to
the aboriginal colour; and what is very remarkable, this reversion
occasionally supervenes, not before birth, but during the growth of the
animal. Hence, if it could be shown that silver-greys and chinchillas were
the offspring of a cross between a black and albino variety with the
colours intimately blended--a supposition in itself not improbable, and
supported by the circumstance of silver-greys in warrens sometimes
producing creamy-white young, which ultimately become black--then all the
above given paradoxical facts on the changes of colour in silver-greys and
in their descendants the Himalayans would come under the law of reversion,
supervening at different periods of growth and in different degrees, either
to the original black or to the original albino parent-variety.
It is, also, remarkable that Himalayans, though produced so suddenly; breed
true. But as, whilst young, they are albinoes, the case falls under a very
general rule; albinism being well known to be strongly inherited, for
instance with white mice and many other quadrupeds, and even white flowers.
But why, it may be asked, do the ears, tail, nose, and feet, and no other
part of the body, revert to a black colour? This apparently depends on a
law, which generally holds good, namely, that characters common to many
species of a genus--and this, in fact, implies long inheritance from the
ancient progenitor of the genus--are found to resist variation, or to
reappear if lost, more persistently than the characters which are confined
to the separate species. Now, in the genus Lepus, a large majority of the
species have their ears and the upper surface of the tail tinted black; but
the persistence of these marks is best seen in those species which in
winter become white: thus, in Scotland the L. variabilis (4/19. G.R.
Waterhouse 'Natural History of Mammalia: Rodents' 1846 pages 52, 60, 105.)
in its winter dress has a shade of colour on its nose, and the tips of its
ears are black: in the L. tibetanus the ears are black, the upper surface
of the tail greyish-black, and the soles of the feet brown: in L. glacialis
the winter fur is pure white, except the soles of the feet and the points
of the ears. Even in the variously-coloured fancy rabbits we may often
observe a tendency in these same parts to be more darkly tinted than the
rest of the body. Thus the several coloured marks on the Himalayan rabbits,
as they grow old, are rendered intelligible. I may add a nearly analogous
case: fancy rabbits very often have a white star on their foreheads; and
the common English hare, whilst young, generally has, as I have myself
observed, a similar white star on its forehead.
When variously coloured rabbits are set free in Europe, and are thus placed
under their natural conditions, they generally revert to the aboriginal
grey colour; this may be in part due to the tendency in all crossed
animals, as lately observed, to revert to their primordial state. But this
tendency does not always prevail; thus silver-grey rabbits are kept in
warrens, and remain true though living almost in a state of nature; but a
warren must not be stocked with both silver-greys and common rabbits;
otherwise "in a few years there will be none but common greys surviving."
(4/20. Delamer on 'Pigeons and Rabbits' page 114.) When rabbits run wild in
foreign countries under new conditions of life, they by no means always
revert to their aboriginal colour. In Jamaica the feral rabbits are
described as having been "slate-coloured, deeply tinted with sprinklings of
white on the neck, on the shoulders, and on the back; softening off to
blue-white under the breast and belly." (4/21. Gosse 'Sojourn in Jamaica'
1851 page 441 as described by an excellent observer, Mr. R. Hill. This is
the only known case in which rabbits have become feral in a hot country.
They can be kept, however, at Loanda (see Livingstone 'Travels' page 407).
In parts of India, as I am informed by Mr. Blyth, they breed well.) But in
this tropical island the conditions were not favourable to their increase,
and they never spread widely, and are now extinct, as I hear from Mr. R.
Hill, owing to a great fire which occurred in the woods. Rabbits during
many years have run wild in the Falkland Islands; they are abundant in
certain parts, but do not spread extensively. Most of them are of the
common grey colour; a few, as I am informed by Admiral Sulivan, are hare-
coloured, and many are black, often with nearly symmetrical white marks on
their faces. Hence, M. Lesson described the black variety as a distinct
species, under the name of Lepus magellanicus, but this, as I have
elsewhere shown, is an error. (4/22. Darwin 'Journal of Researches' page
193; and 'Zoology of the Voyage of the Beagle: Mammalia' page 92.) Within
recent times the sealers have stocked some of the small outlying islets in
the Falkland group with rabbits; and on Pebble Islet, as I hear from
Admiral Sulivan, a large proportion are hare-coloured, whereas on Rabbit
Islet a large proportion are of a bluish colour, which is not elsewhere
seen. How the rabbits were coloured which were turned out of these islets
is not known.
The rabbits which have become feral on the island of Porto Santo, near
Madeira, deserve a fuller account. In 1418 or 1419, J. Gonzales Zarco
(4/23. Kerr 'Collection of Voyages' volume 2 page 177: page 205 for Cada
Mosto. According to a work published in Lisbon in 1717 entitled 'Historia
Insulana' written by a Jesuit, the rabbits were turned out in 1420. Some
authors believe that the island was discovered in 1413.) happened to have a
female rabbit on board which had produced young during the voyage, and he
turned them all out on the island. These animals soon increased so rapidly,
that they became a nuisance, and actually caused the abandonment of the
settlement. Thirty-seven years subsequently, Cada Mosto describes them as
innumerable; nor is this surprising, as the island was not inhabited by any
beast of prey or by any terrestrial mammal. We do not know the character of
the mother-rabbit; but it was probably the common domesticated kind. The
Spanish peninsula, whence Zarco sailed, is known to have abounded with the
common wild species at the most remote historical period; and as these
rabbits were taken on board for food, it is improbable that they should
have been of any peculiar breed. That the breed was well domesticated is
shown by the doe having littered during the voyage. Mr. Wollaston, at my
request, brought home two of these feral rabbits in spirits of wine; and,
subsequently, Mr. W. Haywood sent to me three more specimens in brine, and
two alive. These seven specimens, though caught at different periods,
closely resembled each other. They were full grown, as shown by the state
of their bones. Although the conditions of life in Porto Santo are
evidently highly favourable to rabbits, as proved by their extraordinarily
rapid increase, yet they differ conspicuously in their small size from the
wild English rabbit. Four English rabbits, measured from the incisors to
the anus, varied between 17 and 17 3/4 inches in length; whilst two of the
Porto Santo rabbits were only 14 1/2 and 15 inches in length. But the
decrease in size is best shown by weight; four wild English rabbits
averaged 3 pounds 5 ounces, whilst one of the Porto Santo rabbits, which
had lived for four years in the Zoological Gardens, but had become thin,
weighed only 1 pound 9 ounces. A fairer test is afforded by the comparison
of the well-cleaned limb-bones of a Porto Santo rabbit killed on the island
with the same bones of a wild English rabbit of average size, and they
differed in the proportion of rather less than five to nine. So that the
Porto Santo rabbits have decreased nearly three inches in length, and
almost half in weight of body. (4/24. Something of the same kind has
occurred on the island of Lipari, where, according to Spallanzani ('Voyage
dans les deux Siciles' quoted by Godron 'De l'Espece' page 364), a
countryman turned out some rabbits which multiplied prodigiously, but, says
Spallanzani, "les lapins de l'ile de Lipari sont plus petits que ceux qu'on
eleve en domesticite.") The head has not decreased in length proportionally
with the body; and the capacity of the brain case is, as we shall hereafter
see, singularly variable. I prepared four skulls, and these resembled each
other more closely than do generally the skulls of wild English rabbits;
but the only difference in structure which they presented was that the
supra-orbital processes of the frontal bones were narrower.
In colour the Porto Santo rabbit differs considerably from the common
rabbit; the upper surface is redder, and is rarely interspersed with any
black or black-tipped hairs. The throat and certain parts of the under
surface, instead of being pure white, are generally pale grey or leaden
colour. But the most remarkable difference is in the ears and tail; I have
examined many fresh English rabbits, and the large collection of skins in
the British Museum from various countries, and all have the upper surface
of the tail and the tips of the ears clothed with blackish-grey fur; and
this is given in most works as one of the specific characters of the
rabbit. Now in the seven Porto Santo rabbits the upper surface of the tail
was reddish-brown, and the tips of the ears had no trace of the black
edging. But here we meet with a singular circumstance: in June, 1861 I
examined two of these rabbits recently sent to the Zoological Gardens, and
their tails and ears were coloured as just described; but when one of their
dead bodies was sent to me in February, 1865, the ears were plainly edged,
and the upper surface of the tail was covered with blackish-grey fur, and
the whole body was much less red; so that under the English climate this
individual rabbit had recovered the proper colour of its fur in rather less
than four years!
The two little Porto Santo rabbits, whilst alive in the Zoological Gardens,
had a remarkably different appearance from the common kind. They were
extraordinarily wild and active, so that many persons exclaimed on seeing
them that they were more like large rats than rabbits. They were nocturnal
to an unusual degree in their habits, and their wildness was never in the
least subdued; so that the superintendent, Mr. Bartlett, assured me that he
had never had a wilder animal under his charge. This is a singular fact,
considering that they are descended from a domesticated breed. I was so
much surprised at it, that I requested Mr. Haywood to make inquiries on the
spot, whether they were much hunted by the inhabitants, or persecuted by
hawks, or cats, or other animals; but this is not the case, and no cause
can be assigned for their wildness. They live both on the central, higher
rocky land and near the sea-cliffs, and, from being exceedingly shy and
timid, seldom appear in the lower and cultivated districts. They are said
to produce from four to six young at a birth, and their breeding season is
in July and August. Lastly, and this is a highly remarkable fact, Mr.
Bartlett could never succeed in getting these two rabbits, which were both
males, to associate or breed with the females of several breeds which were
repeatedly placed with them.
If the history of these Porto Santo rabbits had not been known, most
naturalists, on observing their much reduced size, their colour, reddish
above and grey beneath, their tails and ears not tipped with black, would
have ranked them as a distinct species. They would have been strongly
confirmed in this view by seeing them alive in the Zoological Gardens, and
hearing that they refused to couple with other rabbits. Yet this rabbit,
which there can be little doubt would thus have been ranked as a distinct
species, as certainly originated since the year 1420. Finally, from the
three cases of the rabbits which have run wild in Porto Santo, Jamaica, and
the Falkland Islands, we see that these animals do not, under new
conditions of life, revert to or retain their aboriginal character, as is
so generally asserted to be the case by most authors.
OSTEOLOGICAL CHARACTERS.
When we remember, on the one hand, how frequently it is stated that
important parts of the structure never vary; and, on the other hand, on
what small differences in the skeleton fossil species have often been
founded, the variability of the skull and of some other bones in the
domesticated rabbit well deserves attention. It must not be supposed that
the more important differences immediately to be described strictly
characterise any one breed; all that can be said is, that they are
generally present in certain breeds. We should bear in mind that selection
has not been applied to fix any character in the skeleton, and that the
animals have not had to support themselves under uniform habits of life. We
cannot account for most of the differences in the skeleton; but we shall
see that the increased size of the body, due to careful nurture and
continued selection, has affected the head in a particular manner. Even the
elongation and lopping of the ears have influenced in a small degree the
form of the whole skull. The want of exercise has apparently modified the
proportional length of the limbs in comparison with that of the body.
[As a standard of comparison, I prepared skeletons of two wild rabbits from
Kent, one from the Shetland Islands, and one from Antrim in Ireland. As all
the bones in these four specimens from such distant localities closely
resembled each other, presenting scarcely any appreciable difference, it
may be concluded that the bones of the wild rabbit are generally uniform in
character.
SKULL.
I have carefully examined skulls of ten large lop-eared rabbits, and of
five common domestic rabbits, which latter differ from the lop-eared only
in not having such large bodies or ears, yet both larger than in the wild
rabbit. First for the ten lop-eared rabbits: in all these the skull is
remarkably elongated in comparison with its breadth. In a wild rabbit the
length was 3.15 inches, in a large fancy rabbit 4.3; whilst the breadth of
the cranium enclosing the brain was in both almost exactly the same. Even
by taking as the standard of comparison the widest part of the zygomatic
arch, the skulls of the lop-eared are proportionally to their breadth
three-quarters of an inch too long. The depth of the head has increased
almost in the same proportion with the length; it is the breadth alone
which has not increased. The parietal and occipital bones enclosing the
brain are less arched, both in a longitudinal and transverse line, than in
the wild rabbit, so that the shape of the cranium is somewhat different.
The surface is rougher, less cleanly sculptured, and the lines of sutures
are more prominent.
Although the skulls of the large lop-eared rabbits in comparison with those
of the wild rabbit are much elongated relatively to their breadth, yet,
relatively to the size of body, they are far from elongated. The lop-eared
rabbits which I examined were, though not fat, more than twice as heavy as
the wild specimens; but the skull was very far from being twice as long.
Even if we take the fairer standard of the length of body, from the nose to
the anus, the skull is not on an average as long as it ought to be by a
third of an inch. In the small feral Porto Santo rabbit, on the other hand,
the head relatively to the length of body is about a quarter of an inch too
long.
This elongation of the skull relatively to its breadth, I find a universal
character, not only with the large lop-eared rabbits, but in all the
artificial breeds; as is well seen in the skull of the Angora. I was at
first much surprised at the fact, and could not imagine why domestication
could produce this uniform result; but the explanation seems to lie in the
circumstance that during a number of generations the artificial races have
been closely confined, and have had little occasion to exert either their
senses, or intellect, or voluntary muscles; consequently the brain, as we
shall presently more fully see, has not increased relatively with the size
of body. As the brain has not increased, the bony case enclosing it has not
increased, and this has evidently affected through correlation the breadth
of the entire skull from end to end.
(FIGURE 6. SKULL OF WILD RABBIT, of natural size.
FIGURE 7. SKULL OF LARGE LOP-EARED RABBIT, of natural size.
FIGURE 8. PART OF ZYGOMATIC ARCH, showing the projecting end of the malar
bone of the auditory meatus: of natural size. Upper figure, Wild Rabbit.
Lower figure, Lop-eared, hare-coloured Rabbit.
FIGURE 9. POSTERIOR END OF SKULL, of natural size, showing the inter-
parietal bone. A. Wild Rabbit. B. Feral Rabbit from island of P. Santo,
near Madeira. C. Large Lop-eared Rabbit.)
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, figure 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 figure 8; but this circumstance mainly depends on the changed
direction of the meatus. The inter-parietal bone (see figure 9) differs
much in shape in the several skulls; generally it is more oval, that is
more extended in the line of the longitudinal axis of the skull, than in
the wild rabbit. The posterior margin of "the square raised platform"
(4/25. Waterhouse 'Nat. Hist. Mammalia' volume 2 page 36.) of the occiput,
instead of being truncated, or projecting slightly as in the wild rabbit,
is in most lop-eared rabbits pointed, as in figure 9, C. The paramastoids
relatively to the size of the skull are generally much thicker than in the
wild rabbit.
(FIGURE 10. OCCIPITAL FORAMEN, of natural size, in--A. Wild Rabbit; B.
Large Lop-eared Rabbit.)
The occipital foramen (figure 10) presents some remarkable differences: in
the wild rabbit, the lower edge between the condyles is considerably and
almost angularly hollowed out, and the upper edge is deeply and squarely
notched; hence the longitudinal axis exceeds the transverse axis. In the
skulls of the lop-eared rabbits the transverse axis exceeds the
longitudinal; for in none of these skulls was the lower edge between the
condyles so deeply hollowed out; in five of them there was no upper square
notch, in three there was a trace of the notch, and in two alone it was
well developed. These differences in the shape of the foramen are
remarkable, considering that it gives passage to so important a structure
as the spinal marrow, though apparently the outline of the latter is not
affected by the shape of the passage.
(FIGURE 11. SKULL, of natural size, of Half-lop Rabbit, showing the
different direction of the auditory meatus on the two sides, and the
consequent general distortion of the skull. The left ear of the animal (or
right side of figure) lopped forwards.)
In all the skulls of the large lop-eared rabbits, the bony auditory meatus
is conspicuously larger than in the wild rabbit. In a skull 4.3 inches in
length, and which barely exceeded in breadth the skull of a wild rabbit
(which was 3.15 inches in length), the longer diameter of the meatus was
exactly twice as great. The orifice is more compressed, and its margin on
the side nearest the skull stands up higher than the outer side. The whole
meatus is directed more forwards. As in breeding lop-eared rabbits the
length of the ears, and their consequent lopping and lying flat on the
face, are the chief points of excellence, there can hardly be a doubt that
the great change in the size, form, and direction of the bony meatus,
relatively to this same part in the wild rabbit, is due to the continued
selection of individuals having larger and larger ears. The influence of
the external ear on the bony meatus is well shown in the skulls (I have
examined three) of half-lops (see figure 5), in which one ear stands
upright, and the other and longer ear hangs down; for in these skulls there
was a plain difference in the form and direction of the bony meatus on the
two sides. But it is a much more interesting fact, that the changed
direction and increased size of the bony meatus have slightly affected on
the same side the structure of the whole skull. I here give a drawing
(figure 11) of the skull of a half-lop; and it may be observed that the
suture between the parietal and frontal bones does not run strictly at
right angles to the longitudinal axis of the skull; the left frontal bone
projects beyond the right one; both the posterior and anterior margins of
the left zygomatic arch on the side of the lopping ear stand a little in
advance of the corresponding bones on the opposite side. Even the lower jaw
is affected, and the condyles are not quite symmetrical, that on the left
standing a little in advance of that on the right. This seems to me a
remarkable case of correlation of growth. Who would have surmised that by
keeping an animal during many generations under confinement, and so leading
to the disuse of the muscles of the ears, and by continually selecting
individuals with the longest and largest ears, he would thus indirectly
have affected almost every suture in the skull and the form of the lower
jaw!
In the large lop-eared rabbits the only difference in the lower jaw, in
comparison with that of the wild rabbit, is that the posterior margin of
the ascending ramus is broader and more inflected. The teeth in neither jaw
present any difference, except that the small incisors, beneath the large
ones, are proportionately a little longer. The molar teeth have increased
in size proportionately with the increased width of the skull, measured
across the zygomatic arch, and not proportionally with its increased
length. The inner line of the sockets of the molar teeth in the upper jaw
of the wild rabbit forms a perfectly straight line; but in some of the
largest skulls of the lop-eared this line was plainly bowed inwards. In one
specimen there was an additional molar tooth on each side of the upper jaw,
between the molars and premolars; but these two teeth did not correspond in
size; and as no rodent has seven molars, this is merely a monstrosity,
though a curious one.
The five other skulls of common domestic rabbits, some of which approach in
size the above-described largest skulls, whilst the others exceed but
little those of the wild rabbit, are only worth notice as presenting a
perfect gradation in all the above-specified differences between the skulls
of the largest lop-eared and wild rabbits. In all, however, the supra-
orbital plates are rather larger, and in all the auditory meatus is larger,
in conformity with the increased size of the external ears, than in the
wild rabbit. The lower notch in the occipital foramen in some was not so
deep as in the wild rabbit, but in all five skulls the upper notch was well
developed.
The skull of the Angora rabbit, like the latter five skulls, is
intermediate in general proportions, and in most other characters, between
those of the largest lop-eared and wild rabbits. It presents only one
singular character: though considerably longer than the skull of the wild
rabbit, the breadth measured within the posterior supra-orbital fissures is
nearly a third less than in the wild. The skulls of the silver-grey, and
chinchilla and Himalayan rabbits are more elongated than in the wild, with
broader supra-orbital plates, but differ little in any other respect,
excepting that the upper and lower notches of the occipital foramen are not
so deep or so well developed. The skull of the Moscow rabbit scarcely
differs at all from that of the wild rabbit. In the Porto Santo feral
rabbits the supra-orbital plates are generally narrower and more pointed
than in our wild rabbits.
As some of the largest lop-eared rabbits of which I prepared skeletons were
coloured almost like hares, and as these latter animals and rabbits have,
as it is affirmed, been recently crossed in France, it might be thought
that some of the above-described characters had been derived from a cross
at a remote period with the hare. Consequently I examined skulls of the
hare, but no light could thus be thrown on the peculiarities of the skulls
of the larger rabbits. It is, however, an interesting fact, as illustrating
the law that varieties of one species often assume the characters of other
species of the same genus, that I found, on comparing the skulls of ten
species of hares in the British Museum, that they differed from each other
chiefly in the very same points in which domestic rabbits vary,--namely, in
general proportions, in the form and size of the supra-orbital plates, in
the form of the free end of the malar bone, and in the line of suture
separating the occipital and frontal bones. Moreover two eminently variable
characters in the domestic rabbit, namely, the outline of the occipital
foramen and the shape of the "raised platform" of the occiput, were
likewise variable in two instances in the same species of hare.
VERTEBRAE.
The number is uniform in all the skeletons which I have examined, with two
exceptions, namely, in one of the small feral Porto Santo rabbits and in
one of the largest lop-eared kinds; both of these had as usual seven
cervical, twelve dorsal with ribs, but, instead of seven lumbar, both had
eight lumbar vertebrae. This is remarkable, as Gervais gives seven as the
number for the whole genus Lepus. The caudal vertebrae apparently differ by
two or three, but I did not attend to them, and they are difficult to count
with certainty.
(FIGURE 12. ATLAS VERTEBRAE, of natural size; inferior surface viewed
obliquely. Upper figure, Wild Rabbit. Lower figure, Hare-coloured, large,
Lop-eared Rabbit, a, supra-median, atlantoid process; b, infra-median
process.)
In the first cervical vertebra, or atlas, the anterior margin of the neural
arch varies a little in wild specimens, being either nearly smooth, or
furnished with a small supra-median atlantoid process; I have figured a
specimen with the largest process (a) which I have seen; but it will be
observed how inferior this is in size and different in shape to that in a
large lop-eared rabbit. In the latter, the infra-median process (b) is also
proportionally much thicker and longer. The alae are a little squarer in
outline.
(FIGURE 13. THIRD CERVICAL VERTEBRAE, of natural size, of: A. Wild Rabbit;
B. Hare-coloured, large, Lop-eared Rabbit, a, a, inferior surface; b, b,
anterior articular surfaces.)
THIRD CERVICAL VERTEBRA.
In the wild rabbit (figure 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 vertebrae of the wild and lop-eared (A b, B
b) rabbits differ more conspicuously when their anterior articular surfaces
are compared; for the extremities of the antero-dorsal processes in the
wild rabbit are simply rounded, whilst in the lop-eared they are trifid,
with a deep central pit. The canal for the spinal marrow in the lop-eared
(B b) is more elongated in a transverse direction than in the wild rabbit;
and the passages for the arteries are of a slightly different shape. These
several differences in this vertebra seem to me well deserving attention.
FIRST DORSAL VERTEBRA.
Its neural spine varies in length in the wild rabbit; being sometimes very
short, but generally more than half as long as that of the second dorsal;
but I have seen it in two large lop-eared rabbits three-fourths of the
length of that of the second dorsal vertebra.
(FIGURE 14. DORSAL VERTEBRAE, from sixth to tenth inclusive, of natural
size, viewed laterally. A. Wild Rabbit. B. Large, Hare-coloured, so called
Spanish Rabbit.)
NINTH AND TENTH DORSAL VERTEBRAE.
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
vertebrae. In the large lop-eared rabbits the neural spines of the tenth,
ninth, and eighth vertebrae, and even in a slight degree that of the
seventh, are very much thicker, and of somewhat different shape, in
comparison with those of the wild rabbit. So that this part of the
vertebral column differs considerably in appearance from the same part in
the wild rabbit, and closely resembles in an interesting manner these same
vertebrae in some species of hares. In the Angora, Chinchilla, and
Himalayan rabbits, the neural spines of the eighth and ninth vertebrae are
in a slight degree thicker than in the wild. On the other hand, in one of
the feral Porto Santo rabbits, which in most of its characters deviates
from the common wild rabbit, in a direction exactly opposite to that
assumed by the large lop-eared rabbits, the neural spines of the ninth and
tenth vertebrae were not at all larger than those of the several anterior
vertebra. In this same Porto Santo specimen there was no trace in the ninth
vertebra of the anterior lateral processes (see figure 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 (4/26. 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), a haemal spine was
moderately well developed on the under side of the twelfth dorsal vertebra,
and I have seen this in no other specimen.
LUMBAR VERTEBRAE.
I have stated that in two cases there were eight instead of seven lumbar
vertebrae. The third lumbar vertebrae in one skeleton of a wild British
rabbit, and in one of the Porto Santo feral rabbits, had a haemal spine;
whilst in four skeletons of large lop-eared rabbits, and in the Himalayan
rabbit, this same vertebra had a well developed haemal spine.
PELVIS.
In four wild specimens this bone was almost absolutely identical in shape;
but in several domesticated breeds shades of differences could be
distinguished. In the large lop-eared rabbits, the whole upper part of the
ilium is straighter, or less splayed outwards, than in the wild rabbit; and
the tuberosity on the inner lip of the anterior and upper part of the ilium
is proportionally more prominent.
(FIGURE 15. TERMINAL BONE OF STERNUM, of natural size, A. Wild Rabbit. B.
Hare-coloured, Lop-eared Rabbit. C. Hare-coloured Spanish Rabbit. (N.B. The
left-hand angle of the upper articular extremity of B was broken, and has
been accidentally thus represented.))
STERNUM.
The posterior end of the posterior sternal bone in the wild rabbit (figure
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.
(FIGURE 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 (figure 16, A) varies a little in shape and size,
as does the apex of the acromion in sharpness, and the part just below the
rectangular bar in breadth. But the variations in these respects in the
wild rabbit are very slight: whilst in the large lop-eared rabbits they are
considerable. Thus in some specimens (B) the oblique terminal knob is
developed into a short bar, forming an obtuse angle with the rectangular
bar. In another specimen (C) these two unequal bars form nearly a straight
line. The apex of the acromion varies much in breadth and sharpness, as may
be seen by comparing figures B, C, and D.
LIMBS.
In these I could detect no variation; but the bones of the feet were too
troublesome to compare with much care.]
I have now described all the differences in the skeletons which I have
observed. It is impossible not to be struck with the high degree of
variability or plasticity of many of the bones. We see how erroneous the
often-repeated statement is, that only the crests of the bones which give
attachment to muscles vary in shape, and that only parts of slight
importance become modified under domestication. No one will say, for
instance, that the occipital foramen, or the atlas, or the third cervical
vertebra is a part of slight importance. If the several vertebrae of the
wild and lop-eared rabbits, of which figures have been given, had been
found fossil, palaeontologists would have declared without hesitation that
they had belonged to distinct species.
[THE EFFECTS OF THE USE AND DISUSE OF PARTS.
In the large lop-eared rabbits the relative proportional length of the
bones of the same leg, and of the front and hind legs compared with each
other, have remained nearly the same as in the wild rabbit; but in weight,
the bones of the hind legs apparently have not increased in due proportion
with the front legs. The weight of the whole body in the large rabbits
examined by me was from twice to twice and a half as great as that of the
wild rabbit; and the weight of the bones of the front and hind limbs taken
together (excluding the feet, on account of the difficulty of cleaning so
many small bones) has increased in the large lop-eared rabbits in nearly
the same proportion; consequently in due proportion to the weight of body
which they have to support. If we take the length of the body as the
standard of comparison, the limbs of the large rabbits have not increased
in length in due proportion by one inch and a half. Again, if we take as
the standard of comparison the length of the skull, which, as we have
before seen, has not increased in length in due proportion to the length of
body, the limbs will be found to be, proportionally with those of the wild
rabbit, from half to three-quarters of an inch too short. Hence, whatever
standard of comparison be taken, the limb-bones of the large lop-eared
rabbits have not increased in length, though they have in weight, in full
proportion to the other parts of the frame; and this, I presume, may be
accounted for by the inactive life which during many generations they have
spent. Nor has the scapula increased in length in due proportion to the
increased length of the body.
The capacity of the osseous case of the brain is a more interesting point,
to which I was led to attend by finding, as previously stated, that with
all domesticated rabbits the length of the skull relatively to its breadth
has greatly increased in comparison with that of the wild rabbits. If we
had possessed a large number of domesticated rabbits of nearly the same
size with the wild rabbits, it would have been a simple task to have
measured and compared the capacities of their skulls. But this is not the
case: almost all the domestic breeds have larger bodies than wild rabbits,
and the lop-eared kinds are more than double their weight. As a small
animal has to exert its senses, intellect, and instincts equally with a
large animal, we ought not by any means to expect an animal twice or thrice
as large as another to have a brain of double or treble the size. (4/27.
See Prof. Owen's remarks on this subject in his paper on the 'Zoological
Significance of the Brain, etc., of Man, etc.' read before Brit.
Association 1862: with respect to Birds see 'Proc. Zoolog. Soc.' January
11, 1848 page 8.) Now, after weighing the bodies of four wild rabbits, and
of four large but not fattened lop-eared rabbits, I find that on an average
the wild are to the lop-eared in weight as 1 to 2.17; in average length of
body as 1 to 1.41; whilst in capacity of skull they are as 1 to 1.15. Hence
we see that the capacity of the skull, and consequently the size of the
brain, has increased but little, relatively to the increased size of the
body; and this fact explains the narrowness of the skull relatively to its
length in all domestic rabbits.
TABLE 3: MEASUREMENTS OF WILD AND SEMI-WILD RABBITS.
I. Length of Skull (inches).
II. Length of Body from Incisors to Anus (inches).
III. Weight of whole Body (pounds and ounces).
IV. Capacity of Skull measured by Small Shot (grains).
V. Capacity calculated according to Length of Skull relatively to that of
No. 1 (Wild Rabbit, Kent) (grains).
VI. Difference between actual and calculated capacities of Skulls (grains).
VII. Showing how much per cent the Brain, by calculation according to the
length of the Skull is too light (-) or too heavy (+), relatively to the
Brain of the Wild Rabbit No. 1.
NAME OF BREED: I. II. III. IV. V. VI.
VII.
WILD AND SEMI-WILD RABBITS:
1. Wild Rabbit, Kent: 3.15 17.4 3,5 972 .. ..
+2
2. Wild Rabbit, Shetland Islands: 3.15 .. .. 979 .. ..
+2
3. Wild Rabbit, Ireland: 3.15 .. .. 992 .. ..
+2
4. Domestic rabbit, run wild,
Sandon: 3.15 18.5 .. 977 .. ..
+2
5. Wild, common variety,
small specimen, Kent: 2.96 17.0 2,14 875 913 38 -
4
6. Wild, fawn-coloured variety,
Scotland: 3.1 .. .. 918 950 32 -
3
7. Silver-grey, small specimen,
Thetford warren: 2.95 15.5 2,11 938 910 28
+3
8. Feral rabbit, Porto Santo: 2.83 .. .. 893 873 20
+2
9. Feral rabbit, Porto Santo: 2.85 .. .. 756 879 123 -
16
10.Feral rabbit, Porto Santo: 2.95 .. .. 835 910 75 -
9
Average of three Porto Santo
rabbits: 2.88 .. .. 828 888 60 -
7
DOMESTIC RABBITS:
11.Himalayan: 3.5 20.5 .. 963 1080 117 -
12
12.Moscow: 3.25 17.0 3,8 803 1002 199 -
24
13.Angora: 3.5 19.5 3,1 697 1080 383 -
54
14.Chinchilla: 3.65 22.0 .. 995 1126 131 -
13
15.Large lop-eared: 4.1 24.5 7,0 1065 1265 200 -
18
16.Large lop-eared: 4.1 25.0 7,13 1153 1265 112 -
9
17.Large lop-eared: 4.07 .. .. 1037 1255 218 -
21
18.Large lop-eared: 4.1 25.0 7,4 1208 1265 57 -
4
19.Large lop-eared: 4.3 .. .. 1232 1326 94 -
7
20.Large lop-eared: 4.25 .. .. 1124 1311 187 -
16
21.Large hare-coloured: 3.86 24.0 6,14 1131 1191 60 -
5
22.Average of above seven large
lop-eared rabbits: 4.11 24.62 7,4 1136 1268 132 -
11
23.Hare (L. timidus)
English specimen: 3.61 7,0 1315
24.Hare (L. timidus)
German specimen: 3.82 7,0 1455
In the upper half of Table 3 I have given the measurements of the skull of
ten wild rabbits; and in the lower half, of eleven thoroughly domesticated
kinds. As these rabbits differ so greatly in size, it is necessary to have
some standard by which to compare the capacities of their skulls. I have
selected the length of skull as the best standard, for in the larger
rabbits it has not, as already stated, increased in length so much as the
body; but as the skull, like every other part, varies in length, neither it
nor any other part affords a perfect standard.
In the first column of figures the extreme length of the skull is given in
inches and decimals. I am aware that these measurements pretend to greater
accuracy than is possible; but I have found it the least trouble to record
the exact length which the compass gave. The second and third columns give
the length and weight of body, whenever these observations were made. The
fourth column gives the capacity of the skull by the weight of small shot
with which the skulls were filled; but it is not pretended that these
weights are accurate within a few grains. In the fifth column the capacity
is given which the skull ought to have had by calculation, according to the
length of skull, in comparison with that of the wild rabbit No. 1; in the
sixth column the difference between the actual and calculated capacities,
and in the seventh the percentage of increase or decrease, are given. For
instance, as the wild rabbit No. 5 has a shorter and lighter body than the
wild rabbit No. 1, we might have expected that its skull would have had
less capacity; the actual capacity, as expressed by the weight of shot, is
875 grains, which is 97 grains less than that of the first rabbit. But
comparing these two rabbits by the length of their skulls, we see that in
No. 1 the skull is 3.15 inches in length, and in No. 5 2.96 inches in
length; according to this ratio, the brain of No. 5 ought to have had a
capacity of 913 grains of shot, which is above the actual capacity, but
only by 38 grains. Or, to put the case in another way (as in column VII),
the brain of this small rabbit, No. 5, for every 100 grains of weight is
only 4 grains too light,--that is, it ought, according to the standard
rabbit No. 1, to have been 4 per cent heavier. I have taken the rabbit No.
1 as the standard of comparison because, of the skulls having a full
average length, this has the least capacity; so that it is the least
favourable to the result which I wish to show, namely, that the brain in
all long-domesticated rabbits has decreased in size, either actually, or
relatively to the length of the head and body, in comparison with the brain
of the wild rabbit. Had I taken the Irish rabbit, No. 3, as the standard,
the following results would have been somewhat more striking.
Turning to Table 3: the first four wild rabbits have skulls of the same
length, and these differ but little in capacity. The Sandon rabbit (No. 4)
is interesting, as, though now wild, it is known to be descended from a
domesticated breed, as is still shown by its peculiar colouring and longer
body; nevertheless the skull has recovered its normal length and full
capacity. The next three rabbits are wild, but of small size, and they all
have skulls with slightly lessened capacities. The three Porto Santo feral
rabbits (Nos. 8 to 10) offer a perplexing case; their bodies are greatly
reduced in size, as in a lesser degree are their skulls in length and in
actual capacity, in comparison with the skulls of wild English rabbits. But
when we compare the capacities of the skull in the three Porto Santo
rabbits, we observe a surprising difference, which does not stand in any
relation to the slight difference in the length of their skulls, nor, as I
believe, to any difference in the size of their bodies; but I neglected
weighing separately their bodies. I can hardly suppose that the medullary
matter of the brain in these three rabbits, living under similar
conditions, can differ as much as is indicated by the proportional
difference of capacity in their skulls; nor do I know whether it is
possible that one brain may contain considerably more fluid than another.
Hence I can throw no light on this case.
Looking to the lower half of Table 3, which gives the measurements of
domesticated rabbits, we see that in all the capacity of the skull is less,
but in very various degrees, than might have been anticipated according to
the length of their skulls, relatively to that of the wild rabbit No. 1. In
line 22 the average measurements of seven large lop-eared rabbits are
given. Now the question arises, has the average capacity of the skull in
these seven large rabbits increased as much as might have been expected
from the greatly increased size of body. We may endeavour to answer this
question in two ways: in the upper half of the Table we have measurements
of the skulls of six small wild rabbits (Nos. 5 to 10), and we find that on
an average the skulls are .18 of an inch shorter, and in capacity 91 grains
less, than the average length and capacity of the three first wild rabbits
on the list. The seven large lop-eared rabbits, on an average, have skulls
4.11 inches in length, and 1136 grains in capacity; so that these skulls
have increased in length more than five times as much as the skulls of the
six small wild rabbits have decreased in length; hence we might have
expected that the skulls of the large lop-eared rabbits would have
increased in capacity five times as much as the skulls of the six small
rabbits have decreased in capacity; and this would have given an average
increased capacity of 455 grains, whilst the real average increase is only
155 grains. Again, the large lop-eared rabbits have bodies of nearly the
same weight and size as the common hare, but their heads are longer;
consequently, if the lop-eared rabbits had been wild, it might have been
expected that their skulls would have had nearly the same capacity as that
of the skull of the hare. But this is far from being the case; for the
average capacity of the two hare-skulls (Nos. 23, 24) is so much larger
than the average capacity of the seven lop-eared skulls, that the latter
would have to be increased 21 per cent to come up to the standard of the
hare. (4/23. This standard is apparently considerably too low, for Dr.
Crisp ('Proc. Zoolog. Soc.' 1861 page 86) gives 210 grains as the actual
weight of the brain of a hare which weighed 7 pounds, and 125 grains as the
weight of the brain of a rabbit which weighed 3 pounds 5 ounces, 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 table 972 grains; and according to
Dr. Crisp's ratio of 125 to 210, the skull of the hare ought to have
contained 1632 grains of shot, instead of only (in the largest hare in my
table) 1455 grains.)
I have previously remarked that, if we had possessed many domestic rabbits
of the same average size with the wild rabbit, it would have been easy to
compare the capacity of their skulls. Now the Himalayan, Moscow, and Angora
rabbits (Nos. 11, 12, 13 of Table 3) are only a little larger in body and
have skulls only a little longer, than the wild animal, and we see that the
actual capacity of their skulls is less than in the wild animal, and
considerably less by calculation (column 7), according to the difference in
the length of their skulls. The narrowness of the brain-case in these three
rabbits could be plainly seen and proved by external measurement. The
Chinchilla rabbit (No. 14) is a considerably larger animal than the wild
rabbit, yet the capacity of its skull only slightly exceeds that of the
wild rabbit. The Angora rabbit, No. 13, offers the most remarkable case;
this animal in its pure white colour and length of silky fur bears the
stamp of long domesticity. It has a considerably longer head and body than
the wild rabbit, but the actual capacity of its skull is less than that of
even the little wild Porto Santo rabbits. By the standard of the length of
skull the capacity (see column 7) is only half of what it ought to have
been! I kept this individual animal alive, and it was not unhealthy nor
idiotic. This case of the Angora rabbit so much surprised me, that I
repeated all the measurements and found them correct. I have also compared
the capacity of the skull of the Angora with that of the wild rabbit by
other standards, namely, by the length and weight of the body, and by the
weight of the limb-bones; but by all these standards the brain appears to
be much too small, though in a less degree when the standard of the limb-
bones was used; and this latter circumstance may probably be accounted for
by the limbs of this anciently domesticated breed having become much
reduced in weight, from its long-continued inactive life. Hence I infer
that in the Angora breed, which is said to differ from other breeds in
being quieter and more social, the capacity of the skull has really
undergone a remarkable amount of reduction.]
From the several facts above given,--namely, firstly, that the actual
capacity of the skull in the Himalayan, Moscow, and Angora breeds, is less
than in the wild rabbit, though they are in all their dimensions rather
larger animals; secondly, that the capacity of the skull of the large lop-
eared rabbits has not been increased in nearly the same ratio as the
capacity of the skull of the smaller wild rabbits has been decreased; and
thirdly, that the capacity of the skull in these same large lop-eared
rabbits is very inferior to that of the hare, an animal of nearly the same
size,--I conclude, notwithstanding the remarkable differences in capacity
in the skulls of the small Porto Santo rabbits, and likewise in the large
lop-eared kinds, that in all long-domesticated rabbits the brain has either
by no means increased in due proportion with the increased length of the
head and increased size of the body, or that it has actually decreased in
size, relatively to what would have occurred had these animals lived in a
state of nature. When we remember that rabbits, from having been
domesticated and closely confined during many generations, cannot have
exerted their intellect, instincts, senses, and voluntary movements, either
in escaping from various dangers or in searching for food, we may conclude
that their brains will have been feebly exercised, and consequently have
suffered in development. We thus see that the most important and
complicated organ in the whole organisation is subject to the law of
decrease in size from disuse.
Finally, let us sum up the more important modifications which domestic
rabbits have undergone, together with their causes as far as we can
obscurely see them. By the supply of abundant and nutritious food, together
with little exercise, and by the continued selection of the heaviest
individuals, the weight of the larger breeds has been more than doubled.
The bones of the limbs taken together have increased in weight, in due
proportion with the increased weight of body, but the hind legs have
increased less than the front legs; but in length they have not increased
in due proportion, and this may have been caused by the want of proper
exercise. With the increased size of the body the third cervical has
assumed characters proper to the fourth cervical vertebra; and the eighth
and ninth dorsal vertebrae have similarly assumed characters proper to the
tenth and posterior vertebrae. The skull in the larger breeds has increased
in length, but not in due proportion with the increased length of body; the
brain has not duly increased in dimensions, or has even actually decreased,
and consequently the bony case for the brain has remained narrow, and by
correlation has affected the bones of the face and the entire length of the
skull. The skull has thus acquired its characteristic narrowness. From
unknown causes the supra-orbital process of the frontal bones and the free
end of the malar bones have increased in breadth; and in the larger breeds
the occipital foramen is generally much less deeply notched than in wild
rabbits. Certain parts of the scapula and the terminal sternal bones have
become highly variable in shape. The ears have been increased enormously in
length and breadth through continued selection; their weight, conjoined
probably with the disuse of their muscles, has caused them to lop
downwards; and this has affected the position and form of the bony auditory
meatus; and this again, by correlation, the position in a slight degree of
almost every bone in the upper part of the skull, and even the position of
the condyles of the lower jaw.
CHAPTER 1.V.
DOMESTIC PIGEONS.
ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS.
INDIVIDUAL VARIABILITY.
VARIATIONS OF A REMARKABLE NATURE.
OSTEOLOGICAL CHARACTERS: SKULL, LOWER JAW, NUMBER OF VERTEBRAE.
CORRELATION OF GROWTH: TONGUE WITH BEAK; EYELIDS AND NOSTRILS WITH WATTLED
SKIN.
NUMBER OF WING-FEATHERS, AND LENGTH OF WING.
COLOUR AND DOWN.
WEBBED AND FEATHERED FEET.
ON THE EFFECTS OF DISUSE.
LENGTH OF FEET IN CORRELATION WITH LENGTH OF BEAK.
LENGTH OF STERNUM, SCAPULA, AND FURCULUM.
LENGTH OF WINGS.
SUMMARY ON THE POINTS OF DIFFERENCE IN THE SEVERAL BREEDS.
I have been led to study domestic pigeons with particular care, because the
evidence that all the domestic races are descended from one known source is
far clearer than with any other anciently domesticated animal. Secondly,
because many treatises in several languages, some of them old, have been
written on the pigeon, so that we are enabled to trace the history of
several breeds. And lastly, because, from causes which we can partly
understand, the amount of variation has been extraordinarily great. The
details will often be tediously minute; but no one who really wants to
understand the progress of change in domestic animals, and especially 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
doubt that this minuteness is worth while. Notwithstanding the clear
evidence that all the breeds are the descendants of a single species, I
could not persuade myself until some years had passed that the whole amount
of difference between them, had arisen since man first domesticated the
wild rock-pigeon.
I have kept alive all the most distinct breeds, which I could procure in
England or from the Continent; and have prepared skeletons of all. I have
received skins from Persia, and a large number from India and other
quarters of the world. (5/1. The Hon. C. Murray has sent me some very
valuable specimens from Persia; and H.M. Consul, Mr. Keith Abbott, 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 regarding them. Mr. Blyth has freely
communicated 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.) Since my admission into two of the London pigeon-clubs, I
have received the kindest assistance from many of the most eminent
amateurs. (5/2. Mr. B.P. Brent, well known for his various contributions to
poultry literature, 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 formerly showed me
his unrivalled collection of Pouters, and gave me specimens. I had access
to Mr. Wicking's collection, which contained a greater assortment of 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. Harrison Weir I am
likewise indebted. Nor must I by any means pass over the assistance
received from Mr. J.M. Eaton, Mr. Baker, Mr. Evans, and Mr. J. Baily, jun.,
of Mount-street--to the latter gentleman I have been indebted for some
valuable specimens. To all these gentlemen I beg permission to return my
sincere and cordial thanks.)
The races of the Pigeon which can be distinguished, and which breed true,
are very numerous. MM. Boitard and Corbie (5/3. 'Les Pigeons de Voliere et
de Colombier' Paris 1824. During forty-five years the sole occupation of M.
Corbie was the care of the pigeons belonging to the Duchess of Berry.
Bonizzi has described a large number of coloured varieties in Italy: 'Le
variazioni dei colombi Domestici' Padova 1873.) describe in detail 122
kinds; and I could add several European kinds not known to them. In India,
judging from the skins sent me, there are many breeds unknown here; and Sir
W. Elliot informs me that a collection imported by an Indian merchant into
Madras from Cairo and Constantinople included several kinds unknown in
India. I have no doubt that there exist considerably above 150 kinds which
breed true and have been separately named. But of these the far greater
number differ from each other only in unimportant characters. Such
differences will be here entirely passed over, and I shall confine myself
to the more important points of structure. That many important differences
exist we shall presently see. I have looked through the magnificent
collection of the Columbidae in the British Museum, and, with the exception
of a few forms (such as the Didunculus, Calaenas, Goura, etc.), I do not
hesitate to affirm that some domestic races of the rock-pigeon differ fully
as much from each other in external characters as do the most distinct
natural genera. We may look in vain through the 288 known species (5/4.
'Coup d'Oeil sur l'Ordre des Pigeons' par Prince C.L. Bonaparte, Paris
1855. This author makes 288 species, ranked under 85 genera.) for a beak so
small and conical as that of the short-faced tumbler; for one so broad and
short as that of the barb; for one so long, straight, and narrow, with its
enormous wattles, as that of the English carrier; for an expanded upraised
tail like that of the fantail; or for an oesophagus like that of the
pouter. I do not for a moment pretend that the domestic races differ from
each other in their whole organisation as much as the more distinct natural
genera. I refer only to external characters, on which, however, it must be
confessed that most genera of birds have been founded. When, in a future
chapter, we discuss the principle of selection as followed by man, we shall
clearly see why the differences between the domestic races are almost
always confined to external, or at least to externally visible, characters.
Owing to the amount and gradations of difference between the several
breeds, I have found it indispensable in the following classification to
rank them under Groups, Races, and Sub-races; to which varieties and sub-
varieties, all strictly inheriting their proper characters, must often be
added. Even with the individuals of the same sub-variety, when long kept by
different fanciers, different strains can sometimes be recognised. There
can be no doubt that, if well-characterised forms of the several races had
been found wild, all would have been ranked as distinct species, and
several of them would certainly have been placed by ornithologists in
distinct genera. A good classification of the various domestic breeds is
extremely difficult, owing to the manner in which many of the forms
graduate into each other; but it is curious how exactly the same
difficulties are encountered, and the same rules have to be followed, as in
the classification of any natural but difficult group of organic beings. An
"artificial classification" might be followed which would present fewer
difficulties than a "natural classification;" but then it would interrupt
many plain affinities. Extreme forms can readily be defined; but
intermediate and troublesome forms often destroy our definitions. Forms
which may be called "aberrant" must sometimes be included within groups to
which they do not accurately belong. Characters of all kinds must be used;
but as with birds in a state of nature, those afforded by the beak are the
best and most readily appreciated. It is not possible to weigh the
importance of all the characters which have to be used so as to make the
groups and sub-groups of equal value. Lastly, a group may contain only one
race, and another and less distinctly defined group may contain several
races and sub-races, and in this case it is difficult, as in the
classification of natural species, to avoid placing too high a value on the
number of forms which a group may contain.
In my measurements I have never trusted to the eye; and when speaking of a
part being large or small, I always refer to the wild rock-pigeon (Columba
livia) as the standard of comparison. The measurements are given in
decimals of an inch.
(5/5. As I so often refer to the size of the C. livia, or rock-pigeon, it
may be convenient to give the mean between the measurements of two wild
birds, kindly sent me by Dr. Edmondstone from the Shetland Islands.
LENGTH IN INCHES:
From feathered base of beak to end of tail: 14.25
From feathered base of beak 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
Beak.--Thickness, measured vertically at distal end of nostrils: .23
Beak.--Breadth, measured at same place: .16
Feet.--From end of middle toe (without claw) to distal end of tibia: 2.77
Feet.--From end of middle toe to end of hind toe (without claws): 2.02
WEIGHT: 14 1/4 ounces.)
(FIGURE 17. THE ROCK PIGEON, or Columba livia. The parent-form of all
domesticated Pigeons. (5/6. 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. Tegetmeier. It may be confidently asserted that the
characters of the six breeds which have been figured are not in the least
exaggerated.))
DIAGRAM 1. DOMESTIC PIGEONS.
Columba livia or ROCK-PIGEON--
--GROUP I.--(SUB-GROUP (RACE) 1.)--German P.
--Lille P.--
--Dutch P.
--ENGLISH POUTER.
--GROUP II.--(SUB-GROUPS (RACES) 2, 3, 4.)--Kali-Par--Bussora--
--Murassa.
--Dragon--ENGLISH CARRIER.
--Bagadotten--Scanderoon--Pigeon Cygne--RUNT.
--TRONFO.
--BARB.
--GROUP III.--(SUB-GROUPS (RACES) 5, 6, 7, 8, 9.)--
--Java Fantail--FANTAIL
--Turbit--AFRICAN OWL.
--Persian Tumbler--Lotan Tumbler--Common Tumbler--SHORT-FACED TUMBLER.
--INDIAN FRILL-BACK.
--JACOBIN.
--GROUP IV.--(SUB-GROUPS (RACES) 10, 11.)--
--TRUMPETER.
--LAUGHER.
--ENGLISH FRILL-BACK.
--NUN.
--SPOT.
--SWALLOW.
--DOVECOTE PIGEON.
I will now give a brief description of all the principal breeds. Diagram 1.
may aid the reader in learning their names and seeing their affinities. The
rock-pigeon, or Columba livia (including under this name two or three
closely-allied sub-species or geographical races, hereafter to be
described), may be confidently viewed, as we shall see in the next chapter,
as the common parent-form. The names in italics on the right-hand side of
the page show us the most distinct breeds, or those which have undergone
the greatest amount of modification. The lengths of the dotted lines rudely
represent the degree of distinctness of each breed from the parent-stock,
and the names placed under each other in the columns show the more or less
closely connecting links. The distances of the dotted lines from each other
approximately represent the amount of difference between the several
breeds.
(FIGURE 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.
RACE I. POUTER PIGEONS. (KROPFTAUBEN, GERMAN. GROSSES-GORGES, OR BOULANS,
FRENCH.)
Oesophagus of great size, barely separated from the crop, often inflated.
Body and legs elongated. Beak of moderate dimensions.
[SUB-RACE 1/I.
The improved English Pouter, when its crop is fully inflated, presents a
truly astonishing appearance. The habit of slightly inflating the crop is
common to all domestic pigeons, but is carried to an extreme in the Pouter.
The crop does not differ, except in size, from that of other pigeons; but
is less plainly separated by an oblique constriction from the oesophagus.
The diameter of the upper part of the oesophagus is immense, even close up
to the head. The beak in one bird which I possessed was almost completely
buried when the oesophagus was fully expanded. The males, especially when
excited, pout more than the females, and they glory in exercising this
power. If a bird will not, to use the technical expression, "play," the
fancier, as I have witnessed, by taking the beak into his mouth, blows him
up like a balloon; and the bird, then puffed up with wind and pride, struts
about, retaining his magnificent size as long as he can. Pouters often take
flight with their crops inflated. After one of my birds had swallowed a
good meal of peas and water, as he flew up in order to disgorge them and
feed his nearly fledged young, I heard the peas rattling in his inflated
crop as if in a bladder. When flying, they often strike the backs of their
wings together, and thus make a clapping noise.
Pouters stand remarkably upright, and their bodies are thin and elongated.
In connexion with this form of body, the ribs are generally broader and the
vertebrae 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 .O3 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 34 1/2 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 28 1/4 and
14 3/4. There are many sub-varieties of the Pouter of different colours,
but these I pass over.
SUB-RACE 1/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 (5/7. 'Das Ganze der Taubenzucht' Weimar 1837 pl. 11 and 12.)
says that the wings are crossed over the tail, and do not reach to its
extremity.
SUB-RACE 1/III. THE LILLE POUTER.
I know this breed only from description. (5/8. Boitard and Corbie 'Les
Pigeons' etc. page 177 pl. 6.) 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 Corbie; 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 1/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 always be trusted. This sub-race seems considerably different.
The upper part of the oesophagus is much less distended. The bird stands
less upright. The feet are not feathered, and the legs and beak are
shorter. In these respects there is an approach in form to the common rock-
pigeon. The tail-feathers are very long, yet the tips of the closed wings
extend beyond the end of the tail; and the length of the wings, from tip to
tip, and of the body, is greater than in the English Pouter.]
(FIGURE 19. ENGLISH CARRIER.)
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
figures 19 and 20) had existed as wild species, no ornithologist would have
placed them in the same genus with each other or with the rock-pigeon. This
group may, as a general rule, be recognised by the beak being long, with
the skin over the nostrils swollen and often carunculated or wattled, and
with that round the eyes bare and likewise carunculated. The mouth is very
wide, and the feet are large. Nevertheless the Barb, which must be classed
in this same group, has a very short beak, and some runts have very little
bare skin round their eyes.
RACE II. CARRIERS. (TURKISCHE TAUBEN; PIGEONS TURCS, DRAGONS.)
Beak elongated, narrow, pointed; eyes surrounded by much naked, generally
carunculated, skin; neck and body elongated.
[SUB-RACE 2/I. THE ENGLISH CARRIER.
[This is a fine bird, of large size, close feathered, generally dark-
coloured, with an elongated neck. The beak is attenuated and of wonderful
length: in one specimen it was 1.4 inch in length from the feathered base
to the tip; therefore nearly twice as long as that of the rock-pigeon,
which measured only .77. Whenever I compare proportionally any part in the
carrier and rock-pigeon, I take the length of the body from the base of the
beak to the end of the tail as the standard of comparison; and according to
this standard, the beak in one Carrier was nearly half an inch longer than
in the rock-pigeon. The upper mandible is often slightly arched. The tongue
is very long. The development of the carunculated skin or wattle round the
eyes, over the nostrils, and on the lower mandible, is prodigious. The
eyelids, measured longitudinally, were in some specimens exactly twice as
long as in the rock-pigeon. The external orifice or furrow of the nostrils
was also twice as long. The open mouth in its widest part was in one case
.75 of an inch in width, whereas in the rock-pigeon it is only about .4 of
an inch. This great width of mouth is shown in the skeleton by the reflexed
edges of the ramus of the lower jaw. The head is flat on the summit and
narrow between the orbits. The feet are large and coarse; the length, as
measured from end of hind toe to end of middle toe (without the claws), was
in two specimens 2.6 inches; and this, proportionally with the rock-pigeon,
is an excess of nearly a quarter of an inch. One very fine Carrier measured
31 1/2 inches from tip to tip of wing. Birds of this sub-race are too
valuable to be flown as carriers.]
SUB-RACE 2/II. DRAGONS; PERSIAN CARRIERS.
[The English Dragon differs from the improved English Carrier in being
smaller in all its dimensions, and in having less wattle round the eyes and
over the nostrils, and none on the lower mandible. Sir W. Elliot sent me
from Madras a Bagdad Carrier (sometimes called khandesi), the name of which
shows its Persian origin: it would be considered here a very poor Dragon;
the body was of the size of the rock-pigeon, with the beak a little longer,
namely, 1 inch from the tip to the feathered base. The skin round the eyes
was only slightly wattled, whilst that over the nostrils was fairly
wattled. The Hon. C. Murray, also, sent me two Carriers direct from Persia;
these had nearly the same character as the Madras bird, being about as
large as the rock-pigeon, but the beak in one specimen was as much as 1.15
in length; the skin over the nostrils was only moderately, and that round
the eyes scarcely at all wattled.]
SUB-RACE 2/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 figure 24.D. when I treat of
the skeleton. The eyes are surrounded by a wide space of bright red skin,
which, as well as that over the nostrils, is moderately wattled. The
breast-bone is remarkably protuberant, being abruptly bowed outwards. The
feet and tarsi are of great length, larger than in first-rate English
Carriers. The whole bird is of large size, but in proportion to the size of
the body the feathers of the wing and tail are short; a wild rock-pigeon,
of considerably less size, had tail-feathers 4.6 inches in length, whereas
in the large Bagadotten these feathers were scarcely over 4.1 inches in
length. Riedel (5/9. 'Die Taubenzucht' Ulm 1824 s. 42.) remarks that it is
a very silent bird.]
SUB-RACE 2/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 (5/10. 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.), 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 as the wild rock-pigeon. The
shape of the beak, with some little carunculated skin over the nostrils,--
the much elongated eyelids,--the broad mouth measured internally,--the
narrow head,--the feet proportionally a little longer than in the rock-
pigeon,--and the general appearance, all show that this bird is an
undoubted Carrier; yet in one specimen the beak was of exactly the same
length as in the rock-pigeon. In the other specimen the beak (as well as
the opening of the nostrils) was only a very little longer, viz., by .08 of
an inch. Although there was a considerable space of bare and slightly
carunculated skin round the eyes, that over the nostrils was only in a
slight degree rugose. Sir W. Elliot informs me that in the living bird the
eye seems remarkably large and prominent, and the same fact is noticed in
the Persian treatise; but the bony orbit is barely larger than that in the
rock-pigeon.
Amongst the several breeds sent to me from Madras by Sir W. Elliot there is
a pair of the Kali 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 Kali 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 TAUBEN AND HINKELTAUBEN OF
NEUMEISTER; PIGEON BAGADAIS, PIGEON ROMAIN.)
Beak long, massive; body of great size.
[Inextricable confusion reigns in the classification, affinities, and
naming of Runts. Several characters which are generally pretty constant in
other pigeons, such as the length of the wings, tail, legs, and neck, and
the amount of naked skin round the eyes, are excessively variable in Runts.
When the naked skin over the nostrils and round the eyes is considerably
developed and wattled, and when the size of body is not very great, Runts
graduate in so insensible a manner into Carriers, that the distinction is
quite arbitrary. This fact is likewise shown by the names given to them in
different parts of Europe. Nevertheless, taking the most distinct forms, at
least five sub-races (some of them including well-marked varieties) can be
distinguished, which differ in such important points of structure, that
they would be considered as good species in a state of nature.]
SUB-RACE 3/I. SCANDEROON OF ENGLISH WRITERS (DIE FLORENTINER AND
HINKELTAUBEN OF NEUMEISTER).
[Birds of this sub-race, of which I kept one alive and have since seen two
others, differ from the Bagadotten of Neumeister only in not 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 distal end of the radius of the wing. My
bird, as measured from the base of the beak to the root of the tail, was
fully 2 inches longer than the rock-pigeon; yet the tail itself was only 4
inches in length, whereas in the rock-pigeon, which is a much smaller bird,
the tail is 4 5/8 inches in length.
The Hinkel- or Florentiner Taube of Neumeister (Table 13 figure 1) agrees
with the above description in all the specified characters (for the beak is
not mentioned), except that Neumeister expressly says that the neck is
short, whereas in my Scanderoon it was remarkably long and bowed; so that
the Hinkel forms a well-marked variety.]
SUB-RACE 3/II. PIGEON CYGNE AND PIGEON BAGADAIS OF BOITARD AND CORBIE
(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 38 1/2 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 6 1/4 inches in length, and therefore 2
1/4 inches longer than that of the Scanderoon,--a bird of nearly the same
size. The beak is longer, thicker, and broader than in the rock-pigeon,
proportionally with the size of body. The eyelids, nostrils, and internal
gape of mouth are all proportionally very large, as in Carriers. The foot,
from the end of the middle to end of hind toe, was actually 2.85 inches in
length, which is an excess of .32 of an inch over the foot of the rock-
pigeon, proportionally to the relative size of the two birds.]
SUB-RACE 3/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-characterised birds, there can be no doubt of
the propriety of the separation. They are heavy, massive birds, with
shorter necks, legs, and beaks than in the foregoing races. The skin over
the nostrils is swollen, but not carunculated; the naked skin round the
eyes is not very wide, and only slightly carunculated; and I have seen a
fine so-called Spanish Runt with hardly any naked skin round the eyes. Of
the two varieties to be seen in England, one, which is the rarer, has very
long wings and tail, and agrees pretty closely with the last sub-race; the
other, with shorter wings and tail, is apparently the Pigeon romain
ordinaire of Boitard and Corbie. These Runts are apt to tremble like
Fantails. They are bad flyers. A few years ago Mr. Gulliver (5/11. 'Poultry
Chronicle' volume 2 page 573.) exhibited a Runt which weighed 1 pound 14
ounces; 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 pounds 2 1/2 ounces. A very fine rock-pigeon from the Shetland
Islands weighed only 14 1/2 ounces.]
SUB-RACE 3/IV. TRONFO OF ALDROVANDI (LEGHORN RUNT?).
[In Aldrovandi's work published in 1600 there is a coarse woodcut of a
great Italian pigeon, with an elevated tail, short legs, massive body, and
with the beak short and thick. I had imagined that this latter character so
abnormal in the group, was merely a false representation from bad drawing;
but Moore, in his work published in 1735, says that he possessed a Leghorn
Runt of which "the beak was very short for so large a bird." In other
respects Moore's bird resembled the first sub-race or Scanderoon, for it
had a long bowed neck, long legs, short beak, and elevated tail, and not
much wattle about the head. So that Aldrovandi's and Moore's birds must
have formed distinct varieties, both of which seem to be now extinct in
Europe. Sir W. Elliot, however, informs me that he has seen in Madras a
short-beaked Runt imported from Cairo.]
SUB-RACE 3/V. MURASSA (ADORNED PIGEON) OF MADRAS.
[Skins of these handsome chequered birds were sent me from Madras by Sir W.
Elliot. They are rather larger than the largest rock-pigeon, with longer
and more massive beaks. The skin over the nostrils is rather full and very
slightly carunculated, and they have some naked skin round the eyes; feet
large. This breed is intermediate between the rock-pigeon and a very poor
variety of Runt or Carrier.
From these several descriptions we see that with Runts, as with Carriers,
we have a fine gradation from the rock-pigeon (with the Tronfo diverging as
a distinct branch) to our largest and most massive Runts. But the chain of
affinities, and many points of resemblance, between Runts and carriers,
make me believe that these two races have not descended by independent
lines from the rock-pigeon, but from some common parent, as represented in
the Table, which had already acquired a moderately long beak with slightly
swollen skin over the nostrils, and with some slightly carunculated naked
skin round the eyes.]
(FIGURE 20. ENGLISH BARB.)
RACE IV. BARBS. (INDISCHE TAUBEN; 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 much
more closely than do young pigeons of other and equally distinct breeds. At
this early age, the length of beak, the swollen skin over the rather open
nostrils, the gape of the mouth, and the size of the feet, are the same in
both; although these parts afterwards become widely different. We thus see
that embryology (as the comparison of very young animals may perhaps be
called) comes into play in the classification of domestic varieties, as
with species in a state of nature.
Fanciers, with some truth, compare the head and beak of the Barb to that of
a bullfinch. The Barb, if found in a state of nature would certainly have
been placed in a new genus formed for its reception. The body is a little
larger than that of the rock-pigeon, but the beak is more than .2 of an
inch shorter; although shorter, it is both vertically and horizontally
thicker. From the outward flexure of the rami of the lower jaw, the mouth
internally is very broad, in the proportion of .6 to .4 to that of the
rock-pigeon. The whole head is broad. The skin over the nostril is swollen,
but not carunculated, except slightly in first-rate birds when old; whilst
the naked skin round the eye is broad and much carunculated. It is
sometimes so much developed, that a bird belonging to Mr. Harrison Weir
could hardly see to pick up food from the ground. The eyelids in one
specimen were nearly twice as long as those of the rock-pigeon. The feet
are coarse and strong, but proportionally rather shorter than in the rock-
pigeon. The plumage is generally dark and uniform. Barbs, in short, may be
called short-beaked Carriers, bearing the same relation to Carriers that
the Tronfo of Aldrovandi does to the common Runt.]
GROUP III.
This group is artificial, and includes a heterogeneous collection of
distinct forms. It may be defined by the beak, in well-characterised
specimens of the several races, being shorter than in the rock-pigeon, and
by the skin round the eyes not being much developed.
(FIGURE 21. ENGLISH FANTAIL.)
RACE V.-FANTAILS.
SUB-RACE 5/I. EUROPEAN FANTAILS (PFAUENTAUBEN; 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, according to MM.
Boitard and Corbie, 42. I have counted in one of my own birds 33, and at
Calcutta Mr. Blyth (5/12. 'Annals and Mag. of Nat. History' volume 19 1847
page 105.) has counted in an IMPERFECT tail 34 feathers. In Madras, as I am
informed by Sir W. Elliot, 32 is the standard number; but in England number
is much less valued than the position and expansion of the tail. The
feathers are arranged in an irregular double row; their permanent fanlike
expansion 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 vertebrae. I have examined many
specimens of various colours from different countries, and there was no
trace of the oil-gland; this is a curious case of abortion. (5/13. This
gland occurs in most birds; but Nitzsch (in his 'Pterylographie' 1840 page
55) states that it is absent in two species of Columba, in several species
of Psittacus, in some species of Otis, and in most or all birds of the
Ostrich family. It can hardly be an accidental coincidence 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.)
The neck is thin and bowed backwards. The breast is broad and protuberant.
The feet are small. The carriage of the bird is very different from that of
other pigeons; in good birds the head touches the tail-feathers, which
consequently often become crumpled. They habitually tremble much: and their
necks have an extraordinary, apparently convulsive, backward and forward
movement. Good birds walk in a singular manner, as if their small feet were
stiff. Owing to their large tails, they fly badly on a windy day. The dark-
coloured varieties are generally larger than white Fantails.
Although between the best and common Fantails, now existing in England,
there is a vast difference in the position and size of the tail, in the
carriage of the head and neck, in the convulsive movements of the neck, in
the manner of walking, and in the breadth of the breast, the differences so
graduate away, that it is impossible to make more than one sub-race. Moore,
however, an excellent old authority (5/14. See the two excellent editions
published by Mr. J.M. Eaton in 1852 and 1858 entitled 'A Treatise on Fancy
Pigeons.') 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 5/II. JAVA FANTAIL.
[Mr. Swinhoe sent me from Amoy, in China, the skin of a Fantail belonging
to a breed known to have been imported from Java. It was coloured in a
peculiar manner, unlike any European Fantail; and, for a Fantail, had a
remarkably short beak. Although a good bird of the kind, it had only 14
tail-feathers; but Mr. Swinhoe has counted in other birds of this breed
from 18 to 24 tail-feathers. From a rough sketch sent to me, it is evident
that the tail is not so much expanded or so much upraised as in even
second-rate European Fantails. The bird shakes its neck like our Fantails.
It had a well-developed oil-gland. Fantails were known in India, as We
shall hereafter see, before the year 1600; and we may suspect that in the
Java Fantail we see the breed in its earlier and less improved condition.]
(FIGURE 22. AFRICAN OWL.)
RACE VI. TURBIT AND OWL. (MOVENTAUBEN; PIGEONS A CRAVATE.)
Feathers divergent along the front of the neck and breast; beak very short,
vertically rather thick; oesophagus somewhat enlarged.
Turbits and Owls differ from each other slightly in the shape of the head;
the former have a crest, and the beak is differently curved; 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. They
have 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 is inflated, it is 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. (TUMMLER, OR BURZELTAUBEN; CULBUTANTS.)
During flight, tumble backwards; body generally small; beak generally
short, sometimes excessively short and conical.
This race may be divided into four sub-races, namely, Persian, Lotan,
Common, and short-faced Tumblers. These sub-races include many varieties
which breed true. I have examined eight skeletons of various kinds of
Tumblers: excepting in one imperfect and doubtful specimen, the ribs are
only seven in number, whereas the rock-pigeon has eight ribs.
SUB-RACE 7/I. PERSIAN TUMBLERS.
I received a pair direct from Persia, from the Hon. C. Murray. They are
rather smaller birds than the wild rock-pigeon, about the size of the
common dovecote pigeon, white and mottled, slightly feathered on the feet,
with the beak just perceptibly shorter than in the rock-pigeon. H.M.
Consul, Mr. Keith Abbott, informs me that the difference in the length of
beak is so slight, that only practised Persian fanciers can distinguish
these Tumblers from the common pigeon of the country. He informs me that
they fly in flocks high up in the air and tumble well. Some of them
occasionally appear to become giddy and tumble to the ground, in which
respect they resemble some of our Tumblers.
SUB-RACE 7/II. LOTAN, OR LOWTUN: INDIAN GROUND TUMBLERS.
These birds present one of the most remarkable inherited habits or
instincts ever 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 dovecote
pigeon. The beak is proportionally only slightly shorter and rather thinner
than in the rock-pigeon. These birds when gently shaken and placed on the
ground immediately begin tumbling head over heels, and they continue thus
to tumble until taken up and soothed,--the ceremony being generally to blow
in their faces, as in recovering a person from a state of hypnotism or
mesmerism. It is asserted that they will continue to roll over till they
die, if not taken up. There is abundant evidence with respect to these
remarkable peculiarities; but what makes the case the more worthy of
attention is, that the habit has been inherited since before the year 1600,
for the breed is distinctly described in the 'Ayeen Akbery.' (5/15. English
translation by F. Gladwin 4th edition volume 1. The habit of the Lotan is
also described in the Persian treatise 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.' volume 14 1847 page 104; he says that they "may be seen at any of
the Calcutta bird-dealers.") 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 birds of another sub-variety, called the Kalmi Lotan, which begin
to roll over if only touched on the neck with a rod or wand.
SUB-RACE 7/III. COMMON ENGLISH TUMBLERS.
These birds have exactly the same habits as the Persian Tumbler, but tumble
better. The English bird is rather smaller than the Persian, and the beak
is plainly shorter. Compared with the rock-pigeon, and proportionally with
the size of body, the beak is from .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, longer necks, and are feather-footed.
They tumble to an extraordinary degree; as Mr. Brent remarks (5/16.
'Journal of Horticulture' October 22, 1861 page 76.), "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" (5/17. See the account of the House-
tumblers kept at Glasgow, in the 'Cottage Gardener' 1858 page 285. Also Mr.
Brent's paper 'Journal of Horticulture' 1861 page 76.), which is a Scotch
variety, not differing in general appearance and form of beak from the
common Tumbler. Mr. Brent states that these birds generally begin to tumble
"almost as soon as they can well fly; at three months old they tumble well,
but still fly strong; at five or six months they tumble excessively; and in
the second year they mostly give up flying, on account of their tumbling so
much and so close to the ground. Some fly round with the flock, throwing a
clean summersault every few yards, till they are obliged to settle from
giddiness and exhaustion. These are called Air Tumblers, and they commonly
throw from twenty to thirty summersaults in a minute, each clear and clean.
I have one red cock that I have on two or three occasions timed by my
watch, and counted forty summersaults in the minute. Others tumble
differently. At first they throw a single summersault, then it is double,
till it becomes a continuous roll, which puts an end to flying, for if they
fly a few yards over they go, and roll till they reach the ground. Thus I
had one kill herself, and another broke his leg. Many of them turn over
only a few inches from the ground, and will tumble two or three times in
flying across their loft. These are called House-tumblers, from tumbling in
the house. The act of tumbling seems to be one over which they have no
control, an involuntary movement which they seem to try to prevent. I have
seen a bird sometimes in his struggles fly a yard or two straight upwards,
the impulse forcing him backwards while he struggles to go forwards. If
suddenly startled, or in a strange place, they seem less able to fly than
if quiet in their accustomed loft." These House-tumblers differ from the
Lotan or Ground Tumbler of India, in not requiring to be shaken in order to
begin tumbling. The breed has probably been formed merely by selecting the
best common Tumblers, though it is possible that they may have been crossed
at some former period with Lotans.
(FIGURE 23. SHORT-FACED ENGLISH TUMBLER.)
SUB-RACE 7/1V. 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
Columbidae. Their heads are nearly globular and upright in front, so that
some fanciers say (5/18. J.M. Eaton 'Treatise on Pigeons' 1852 page 9.)
"the head should resemble a cherry with a barleycorn stuck in it." These
are the smallest kind of pigeons. Mr. Esquilant possessed a blue Baldhead,
two years old, which when alive weighed, before feeding-time, only 6 ounces
5 drs.; two others, each weighed 7 ounces. We have seen that a wild rock-
pigeon weighed 14 ounces 2 drs., and a Runt 34 ounces 4 drs. Short-faced
Tumblers have a remarkably erect carriage, with prominent breasts, drooping
wings, and very small feet. The length of the beak from the tip to the
feathered base was in one good bird only .4 of an inch; in a wild rock-
pigeon it was exactly double this length. As these Tumblers have shorter
bodies than the wild rock-pigeon, they ought of course to have shorter
beaks; but proportionally with the size of 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 scutellae. The primary wing-feathers are not rarely nine instead of
ten in number. The improved short-faced Tumblers have almost lost the power
of tumbling; but there are several authentic accounts of their occasionally
tumbling. There are several sub-varieties, such as Bald-heads, Beards,
Mottles, and Almonds; the latter are remarkable from not acquiring their
perfectly-coloured plumage until they have moulted three or four times.
There is good reason to believe that most of these sub-varieties, some of
which breed truly, have arisen since the publication of Moore's treatise in
1735. (5/19. J.M. Eaton 'Treatise' edition 1858 page 76.)
Finally, in regard to the whole group of Tumblers, it is impossible to
conceive a more perfect gradation than I have now lying before me, from the
rock-pigeon, through Persian, Lotan, and common Tumblers, up to the
marvellous short-faced birds; which latter, no ornithologist, judging from
mere external structure, would place in the same genus with the rock-
pigeon. The differences between the successive steps in this series are not
greater than those which may be observed between common dovecote-pigeons
(C. livia) brought from different countries.]
RACE VIII. INDIAN FRILL-BACK.
Beak very short; feathers reversed.
[A specimen of this bird, in spirits, was sent to me from Madras by Sir W.
Elliot. It is wholly different from the Frill-back often exhibited in
England. It is a smallish bird, about the size of the common Tumbler, but
has a beak in all its proportions like our short-faced Tumblers. The beak,
measured from the tip to the feathered base, was only .46 of an inch in
length. The feathers over the whole body are reversed or curl backwards.
Had this bird occurred in Europe, I should have thought it only a monstrous
variety of our improved Tumbler: but as short-faced Tumblers are not known
in India, I think it must rank as a distinct breed. Probably this is the
breed seen by Hasselquist in 1757 at Cairo, and said to have been imported
from India.]
RACE IX. JACOBIN. (ZOPF- OR PERRFICKENTAUBE; NONNAIN.)
Feathers of the neck forming a hood; wings and tail long; beak moderately
short.
[This pigeon can at once be recognised by its hood, almost enclosing the
head and meeting in front of the neck. The hood seems to be merely an
exaggeration of the crest of reversed feathers on the back of the head,
which is common to many sub-varieties, and which in the Latztaube (5/20.
Neumeister 'Taubenzucht' tab. 4. figure 1.) 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 1 1/4 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 2 1/4 inches too long, and the two wings, from tip
to tip, 5 1/4 inches too long. In disposition this bird is singularly
quiet, seldom flying or moving about, as Bechstein and Riedel have likewise
remarked in Germany. (5/21. Riedel 'Die Taubenzucht' 1824 s. 26. Bechstein
'Naturgeschichte Deutschlands' b. 4 s. 36 1795) The latter author also
notices the length of the wings and tail. The beak is nearly .2 of an inch
shorter in proportion to the size of the body than in the rock-pigeon; but
the internal gape of the mouth is considerably wider.]
GROUP IV.
The birds of this group may be characterised by their resemblance in all
important points of structure, especially in the beak, to the rock-pigeon.
The Trumpeter forms the only well-marked race. Of the numerous other sub-
races and varieties I shall specify only a few of the most distinct, which
I have myself seen and kept alive.
RACE X. TRUMPETER. (TROMMELTAUBE; PIGEON TAMBOUR, GLOUGLOU.)
A tuft of feathers at the base of the beak curling forward; feet much
feathered; voice very peculiar; size exceeding that of the rock-pigeon.
[This is a well-marked breed, with a peculiar voice, wholly unlike that of
any other pigeon. The coo is rapidly repeated, and is continued for several
minutes; hence their name of Trumpeters. They are also characterised by a
tuft of elongated feathers, which curls forward over the base of the beak,
and which is possessed by no other breed. Their feet are so heavily
feathered, that they almost appear like little wings. They are larger birds
than the rock-pigeon, but their beak is of very nearly the same
proportional size. Their feet are rather small. This breed was perfectly
characterised in Moore's time, in 1735. Mr. Brent says that two varieties
exist, which differ in size.]
RACE XI. SCARCELY DIFFERING IN STRUCTURE FROM THE WILD COLUMBA LIVIA.
SUB-RACE 11/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 character supposed seldom to vary
with birds. Although the voice of the Laugher is very different from that
of the Trumpeter, yet one of my Trumpeters used to utter a single note like
that of the Laugher. I have kept two varieties of Laughers, which differed
only in one variety being turn-crowned; the smooth-headed kind, for which I
am indebted to the kindness of Mr. Brent, besides its peculiar note, used
to coo in a singular and pleasing manner, which, independently, struck both
Mr. Brent and myself as resembling that of the turtle-dove. Both varieties
come from Arabia. This breed was known by Moore in 1735. A pigeon which
seems to say Yak-roo is mentioned in 1600 in the 'Ayeen Akbery' and is
probably the same breed. Sir W. Elliot has also sent me from Madras a
pigeon called Yahui, said to have come from Mecca, which does not differ in
appearance from the Laugher; it has "a deep melancholy voice, like Yahu,
often repeated." Yahu, yahu, means Oh God, oh God; and Sayzid Mohammed
Musari, in the treatise written about 100 years ago, says that these birds
"are not flown, because they repeat the name of the most high God." Mr.
Keith Abbott, however, informs me that the common pigeon is called Yahoo in
Persia.]
SUB-RACE 11/II. COMMON FRILL-BACK (DIE STRUPPTAUBE).
Beak rather longer than in the rock-pigeon; feathers reversed .
[This is a considerably larger bird than the rock-pigeon, and with the
beak, proportionally with the size of body, a little (viz. by .04 of an
inch) longer. The feathers, especially on the wing-coverts, have their
points curled upwards or back-wards.]
SUB-RACE 11/III. NUNS (PIGEONS COQUILLES).
[These elegant birds are smaller than the rock-pigeon. The beak is actually
1.7, and proportionally with the size of the body .1 of an inch shorter
than in the rock-pigeons, although of the same thickness. In young birds
the scutellae on the tarsi and toes are generally of a leaden-black colour;
and this is a remarkable character (though observed in a lesser degree in
some other breeds), as the colour of the legs in the adult state is subject
to very little variation in any breed. I have on two or three occasions
counted thirteen or fourteen feathers in the tail; this likewise occurs in
the barely distinct breed called Helmets. Nuns are symmetrically coloured,
with the head, primary wing-feathers, tail, and tail-coverts of the same
colour, namely, black or red, and with the rest of the body white. This
breed has retained the same character since Aldrovandi wrote in 1600. I
have received from Madras almost similarly coloured birds.]
SUB-RACE 11/IV. SPOTS (DIE BLASSTAUBEN; PIGEONS HEURTES).
[These birds are a very little larger than the rock-pigeon, with the beak a
trace smaller in all its dimensions, and with the feet decidedly smaller.
They are symmetrically coloured, with a spot on the forehead, with the tail
and tail-coverts of the same colour, the rest of the body being white. This
breed existed in 1676 (5/22. Willughby 'Ornithology' edited by Ray.); and
in 1735 Moore remarks that they breed truly, as is the case at the present
day.]
SUB-RACE 11/V. SWALLOWS.
[These birds, as measured from tip to tip of wing, or from the end of the
beak to the end of the tail, exceed in size the rock-pigeon; but their
bodies are much less bulky; their feet and legs are likewise smaller. The
beak is of about the same length, but rather slighter. Altogether their
general appearance is considerably different from that of the rock-pigeon.
Their heads and wings are of the same colour, the rest of the body being
white. Their flight is said to be peculiar. This seems to be a modern
breed, which, however, originated before the year 1795 in Germany, for it
is described by Bechstein.
Besides the several breeds now described, three or four other very distinct
kinds existed lately, or perhaps still exist, in Germany and France.
Firstly, the Karmeliten, or carme pigeon, which I have not seen; it is
described as of small size, with very short legs, and with an extremely
short beak. Secondly, the Finnikin, which is now extinct in England. It
had, according to Moore's (5/23. J.M. Eaton's edition (1858) of Moore page
98.) treatise, published in 1735, a tuft of feathers on the hinder part of
the head, which ran down its back not unlike a horse's mane. "When it is
salacious it rises over the hen and turns round three or four times,
flapping its wings, then reverses and turns as many times the other way."
The Turner, on the other hand, when it "plays to the female, turns only one
way." Whether these extraordinary statements may be trusted I know not; but
the inheritance of any habit may be believed, after what we have seen with
respect to the Ground-tumbler of India. MM. Boitard and Corbie describe a
pigeon (5/24. Pigeon pattu plongeur. 'Les Pigeons' etc. page 165.) 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, etc., which are all remarkable from their manner of flight. Mr.
Brent informs me that he has seen one of these breeds in Germany with its
wing-feathers injured from having been so often struck together but he did
not see it flying. An old stuffed specimen of a Finnikin in the British
Museum presents no well-marked character. Thirdly, a singular pigeon with a
forked tail is mentioned in some treatises; and as Bechstein (5/25.
'Naturgeschichte Deutschlands' b. 4 s. 47.) briefly describes and figures
this bird, with a tail "having completely the structure of that of the
house-swallow," it must once have existed, for Bechstein was far too good a
naturalist to have confounded any distinct species with the domestic
pigeon. Lastly, an extraordinary pigeon imported from Belgium has lately
been exhibited at the Philoperisteron Society in London (5/26. Mr. W.B.
Tegetmeier 'Journal of Horticulture' January 20, 1863 page 58.), 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 15 1/2 inches from tip to beak to end of tail, and 32 1/2 inches from
tip to tip of wing; now the wild rock-pigeon weighs 14 1/2 ounces, and
measures from tip to beak to end of tail 15 inches, and from tip to tip of
wing only 26 3/4 inches.]
I have now described all the domestic pigeons known to me, and have added a
few others on reliable authority. I have classed them under four Groups, in
order to mark their affinities and degrees of difference; but the third
group is artificial. The kinds examined by me form eleven races, which
include several sub-races; and even these latter present differences that
would certainly have been thought of specific value if observed in a state
of nature. The sub-races likewise include many strictly inherited
varieties; so that altogether there must exist, as previously remarked,
above 150 kinds which can be distinguished, though generally by characters
of extremely slight importance. Many of the genera of the Columbidae,
admitted by ornithologists, do not differ in any great degree from each
other; taking this into consideration, there can be no doubt that several
of the most strongly characterised domestic forms, if found wild, would
have been placed in at least five new genera. Thus a new genus would have
been formed for the reception of the improved English Pouter: a second
genus for Carriers and Runts; and this would have been a wide or
comprehensive genus, for it would have admitted common Spanish Runts
without any wattle, short-beaked Runts like the Tronfo, and the improved
English Carrier: a third genus would have been formed for the Barb: a
fourth for the Fantail: and lastly, a fifth for the short beaked, not-
wattled pigeons, such as Turbits and short-faced Tumblers. The remaining
domestic forms might have been included, in the same genus with the wild
rock-pigeon.
INDIVIDUAL VARIABILITY; VARIATIONS OF A REMARKABLE NATURE.
The differences which we have as yet considered are characteristic of
distinct breeds; but there are other differences, either confined to
individual birds, or often observed in certain breeds but not
characteristic of them. These individual differences are of importance, as
they might in most cases be secured and accumulated by man's power of
selection and thus an existing breed might be greatly modified or a new one
formed. Fanciers notice and select only those slight differences which are
externally visible; but the whole organisation is so tied together by
correlation of growth, that a change in one part is frequently accompanied
by other changes. For our purpose, modifications of all kinds are equally
important, and if affecting a part which does not commonly vary, are of
more importance than a modification in some conspicuous part. At the
present day any visible deviation of character in a well-established breed
is rejected as a blemish; but it by no means follows that at an early
period, before well-marked breeds had been formed, such deviations would
have been rejected; on the contrary, they would have been eagerly preserved
as presenting a novelty, and would then have been slowly augmented, as we
shall hereafter more clearly see, by the process of unconscious selection.
[I have made numerous measurements of the various parts of the body in the
several breeds, and have hardly ever found them quite the same in birds of
the same breed,--the differences being greater than we commonly meet with
in wild species within the same district. To begin with the primary
feathers of the wing and tail; but I must first mention, as some readers
may not be aware of the fact, that the number of the primary wing and tail-
feathers in wild birds is generally constant, and characterises, not only
whole genera, but even whole families. When the tail-feathers are unusually
numerous, as for instance in the swan, they are apt to be variable in
number; but this does not apply to the several species and genera of the
Columbidae, which never (as far as I can hear) have less than twelve or
more than sixteen tail-feathers; and these numbers characterise, with rare
exception, whole sub-families. (5/27. 'Coup-d'oeil sur L'Ordre des Pigeons'
par C.L. Bonaparte 'Comptes Rendus' 1854-55. Mr. Blyth in 'Annals of Nat.
Hist.' volume 19 1847 page 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.") The wild
rock-pigeon has twelve tail-feathers. With Fantails, as we have seen, the
number varies from fourteen to forty-two. In two young birds in the same
nest I counted twenty-two and twenty-seven feathers. Pouters are very
liable to have additional tail-feathers, and I have seen on several
occasions fourteen or fifteen in my own birds. Mr. Bult had a specimen,
examined by Mr. Yarrell, with seventeen tail-feathers. I had a Nun with
thirteen, and another with fourteen tail-feathers; and in a Helmet, a breed
barely distinguishable from the Nun, I have counted fifteen, and have heard
of other such instances. On the other hand, Mr. Brent possessed a Dragon,
which during its whole life never had more than ten tail-feathers; and one
of my Dragons, descended from Mr. Brent's, had only eleven. I have seen a
Bald-head 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 Columbidae, 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 primaries of a white colour being one of the points in Short-
faced Bald-head-Tumblers. Mr. Brent, however, had an Air-Tumbler (not
short-faced) which had in both wings eleven primaries. Mr. Corker, the
eminent breeder of prize Carriers, assures me that some of his birds had
eleven primaries in both wings. I have seen eleven in one wing in two
Pouters. I have been assured by three fanciers that they have seen twelve
in Scanderoons; but as Neumeister asserts that in the allied Florence Runt
the middle flight-feather is often double, the number twelve may have been
caused by two of the ten primaries having each two shafts to a single
feather. The secondary wing-feathers are difficult to count, but the number
seems to vary from twelve to fifteen. The length of the wing and tail
relatively to the body, and of the wings to the tail, certainly varies; I
have especially noticed this in Jacobins. In Mr. Bult's magnificent
collection of Pouters, the wings and tail varied greatly in length; and
were sometimes so much elongated that the birds could hardly play upright.
In the relative length of the few first primaries I have observed only a
slight degree of variability. Mr. Brent informs me that he has observed the
shape of the first feather to vary very slightly. But the variation in
these latter points is extremely slight compared with the differences which
may be observed in the natural species of the Columbidae.
In the beak I have seen very considerable differences in birds of the same
breed, as in carefully bred Jacobins and Trumpeters. In Carriers there is
often a conspicuous difference in the degree of attenuation and curvature
of the beak. So it is indeed in many breeds: thus I had two strains of
black Barbs, which evidently differed in the curvature of the upper
mandible. In width of mouth I have found a great difference in two
Swallows. In Fantails of first-rate merit I have seen some birds with much
longer and thinner necks than in others. Other analogous facts could be
given. We have seen that the oil-gland is aborted in all Fantails (with the
exception of the sub-race from Java), and, I may add, so hereditary is this
tendency to abortion, that some, although not all, of the mongrels which I
reared 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 scutellae on the toes often varies in the same breed, and
sometimes even differs on the two feet of the same individual; the Shetland
rock-pigeon has fifteen on the middle, and six on the hinder toe; whereas I
have seen a Runt with sixteen on the middle and eight on the hind toe; and
a short-faced Tumbler with only twelve and five on these same toes. The
rock-pigeon has no sensible amount of skin between its toes; but I
possessed a Spot and a Nun with the skin extending for a space of a quarter
of an inch from the fork, between the two INNER toes. On the other hand, as
will hereafter be more fully shown, pigeons with feathered feet very
generally have the bases of their OUTER toes connected by skin. I had a red
Tumbler, which had a coo unlike that of its fellows, approaching in tone to
that of the Laugher: this bird had the habit, to a degree which I never saw
equalled in any other pigeon, of often walking with its wings raised and
arched in an elegant-manner. I need say nothing on the great variability,
in almost every breed, in size of body, in colour, in the feathering of the
feet, and in the feathers on the back of the head being reversed. But I may
mention a remarkable Tumbler (5/28. Described and figured in the 'Poultry
Chronicle' volume 3 1855 page 82.) 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 a hen Jacobin with the
feathers on the thigh so long as to reach the ground, and a cock having,
but in a lesser degree, the same peculiarity: from these two birds he bred
others similarly characterised, which were exhibited at the Philoperisteron
Soc. I bred a mongrel pigeon which had fibrous feathers, and the wing and
tail-feathers so short and imperfect that the bird could not fly even a
foot in height.]
There are many singular and inherited peculiarities in the plumage of
pigeons: thus Almond-Tumblers do not acquire their perfect mottled feathers
until they have moulted three or four times: the Kite Tumbler is at first
brindled black and red with a barred appearance, but when "it throws its
nest feathers it becomes almost black, generally with a bluish tail, and a
reddish colour on the inner webs of the primary wing-feathers." (5/29. 'The
Pigeon Book' by Mr. B.P. Brent 1859 page 41.) Neumeister describes a breed
of a black colour with white bars on the wing and a white crescent-shaped
mark on the breast; these marks are generally rusty-red before the first
moult, but after the third or fourth moult they undergo a change; the wing-
feathers and the crown of the head likewise then become white or grey.
(5/30. 'Die staarhalsige Taube. Das Ganze, etc.' s. 21 tab. 1. figure 4.)
It is an important fact, and I believe there is hardly an exception to the
rule, that the especial characters for which each breed is valued are
eminently variable: thus, in the Fantail, the number and direction of the
tail-feathers, the carriage of the body, and the degree of trembling are
all highly variable points; in Pouters, the degree to which they pout, and
the shape of their inflated crops; in the Carrier, the length, narrowness,
and curvature of the beak, and the amount of wattle; in Short-faced
Tumblers, the shortness of the beak, the prominence of the forehead, and
general carriage (5/31. 'A Treatise on the Almond-Tumbler' by J.M. Eaton
1852 page 8 et passim.), and in the Almond-Tumbler the colour of the
plumage; in common Tumblers, the manner of tumbling; in the Barb, the
breadth and shortness of the beak and the amount of eye-wattle; in Runts,
the size of body; in Turbits the frill; and lastly in Trumpeters, the
cooing, as well as the size of the tuft of feathers over the nostrils.
These, which are the distinctive and selected characters of the several
breeds, are all eminently variable.
There is another interesting fact with respect to the characters of the
several breeds, namely, that they are often most strongly displayed in the
male bird. In Carriers, when the males and females are exhibited in
separate pens, the wattle is plainly seen to be much more developed in the
males, though I have seen a hen Carrier belonging to Mr. Haynes heavily
wattled. Mr. Tegetmeier informs me that, in twenty Barbs in Mr. P.H.
Jones's possession, the males had generally the largest eye-wattles; Mr.
Esquilant also believes in this rule, but Mr. H. Weir, a first-rate judge,
entertains some doubt on the subject. 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 excellently; but this is an unusual
circumstance. Mr. Harrison Weir, a successful breeder of prize Fantails,
informs me that his male birds often have a greater number of tail-feathers
than the females. Mr. Eaton asserts (5/32. 'A Treatise, etc.' page 10.)
that if a cock and hen Tumbler were of equal merit, the hen would be worth
double the money; and as pigeons always pair, so that an equal number of
both sexes is necessary for reproduction, this seems to show that high
merit is rarer in the female than in the male. In the development of the
frill in Turbits, of the hood in Jacobins, of the tuft in Trumpeters, of
tumbling in Tumblers, there is no difference between the males and females.
I may here add a rather different case, namely, the existence in France
(5/33. Boitard and Corbie 'Les Pigeons' etc. 1824 page 173.) of a wine-
coloured variety of the Pouter, in which the male is generally chequered
with black, whilst the female is never so chequered. Dr. Chapuis also
remarks (5/34. 'Le Pigeon Voyageur Belge' 1865 page 87. I have given in my
'Descent of Man' 6th edition page 466 some curious cases, on the authority
of Mr. Tegetmeier, of silver-coloured (i.e. very pale blue) birds being
generally females, and of the ease with which a race thus characterised
could be produced. Bonizzi (see 'Variazioni dei Columbi domestici' Padova
1873) states that certain coloured spots are often different in the two
sexes, and the certain tints are commoner in females than in male pigeons.)
that in certain light-coloured pigeons the males have their feathers
striated with black, and these striae 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 remarkable facts, for
there is no sensible difference at any age between the two sexes in the
aboriginal rock-pigeon; and not often any strongly marked difference
throughout the family of the Columbidae. (5/35. Prof. A. Newton 'Proc.
Zoolog. Soc.' 1865 page 716 remarks that he knows no species which present
any remarkable sexual distinction; but Mr. Wallace informs me, that in the
sub-family of the Treronidae the sexes often differ considerably in colour.
See also on sexual differences in the Columbidae, Gould 'Handbook to the
Birds of Australia' volume 2 pages 109-149.)
OSTEOLOGICAL CHARACTERS.
In the skeletons of the various breeds there is much variability; and
though certain differences occur frequently, and others rarely, in certain
breeds, yet none can be said to be absolutely characteristic of any breed.
Considering that strongly-marked domestic races have been formed chiefly by
man's selection, we ought not to expect to find great and constant
differences in the skeleton; for fanciers neither see, nor do they care
for, modifications of structure in the internal framework. Nor ought we to
expect changes in the skeletons from changed habits of life; as every
facility is given to the most distinct breeds to follow the same habits,
and the much modified races are never allowed to wander abroad and procure
their own food in various ways. Moreover, I find, on comparing the
skeletons of Columba livia, oenas, palumbus, and turtur, which are ranked
by all systematists in two or three distinct though allied genera, that the
differences are extremely slight, certainly less than between the skeletons
of some of the most distinct domestic breeds. How far the skeleton of the
wild rock-pigeon is constant I have had no means of judging, as I have
examined only two.
(FIGURE 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.)
[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 proportionally broader. In the Short-faced Tumbler
the skull is more globular: all the bones of the face are much shortened,
and the front of the skull and descending nasal bones are almost
perpendicular: the maxillo-jugal arch and premaxillary bones form an almost
straight line; the space between the prominent edges of the eye-orbits is
depressed. In the Barb the premaxillary bones are much shortened, and their
anterior portion is thicker than in the rock-pigeon, as is the lower part
of the nasal bone. In two Nuns the ascending branches of the
premaxillaries, near their tips, were somewhat attenuated, and in these
birds, as well as in some others, for instance in the Spot, the occipital
crest over the foramen was considerably more prominent than in the rock-
pigeon.
(FIGURE 25. LOWER JAWS, seen from above, of natural size. A. Rock-pigeon.
B. Runt. C. Barb.
FIGURE 26. SKULL OF RUNT, seen from above, of natural size, showing the
reflexed margin of the distal portion of the lower jaw.
FIGURE 27. LATERAL VIEW OF JAWS, of natural size. A. Rock-pigeon. B.
Shortfaced Tumbler. C. Bagadotten Carrier.)
In the lower jaw, the articular surface is proportionably smaller in many
breeds than in the rock-pigeon; and the vertical diameter, more especially
of the outer part of the articular surface, is considerably shorter. May
not this be accounted for by the lessened use of the jaws, owing to
nutritious food having been given during a long period to all highly
improved pigeons? In Runts, Carriers, and Barbs (and in a lesser degree in
several breeds), the whole side of the jaw near the articular end is bent
inwards in a highly remarkable manner; and the superior margin of the
ramus, beyond the middle, is reflexed in an equally remarkable manner, as
may be seen in figure 25, in comparison with the jaw of the rock-pigeon.
This reflection of the upper margin of the lower jaw is plainly connected
with the singularly wide gape of the mouth, as has been described in Runts,
Carriers, and Barbs. The reflection is well shown in figure 26 of the head
of a Runt seen from above; here a wide open space may be observed on each
side, between the edges of the lower jaw and of the premaxillary bones. In
the rock-pigeon, and in several domestic breeds, the edges of the lower jaw
on each side come close up to the premaxillary bones, so that no open space
is left. The degree of downward curvature of the distal half of the lower
jaw also differs to an extraordinary degree in some breeds, as may be seen
in the drawings (figure 27 A) of the rock-pigeon, (B) of the Short-faced
Tumbler, and (C) of the Bagadotten Carrier of Neumeister. In some Runts the
symphysis of the lower jaw is remarkably solid. No one would readily have
believed that jaws differing in the several above-specified points so
greatly could have belonged to the same species.
VERTEBRAE.
All the breeds have twelve cervical vertebrae. (5/36. I am not sure that I
have designated the different kinds of vertebra correctly: but I observe
that different anatomists 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.) But in a Bussorah Carrier from India the twelfth
vertebra carried a small rib, a quarter of an inch in length, with a
perfect double articulation.
The DORSAL VERTEBRAE 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, eight bear ribs; the eighth rib
being very thin and the seventh having no process. In Pouters all the ribs
are extremely broad, and, in three out of four skeletons examined by me,
the eighth rib was twice or even thrice as broad as in the rock-pigeon; and
the seventh pair had distinct processes. In many breeds there are only
seven ribs, as in seven out of eight skeletons of various Tumblers, and in
several skeletons of Fantails, Turbits and Nuns.
In all these breeds the seventh pair was very small, and was destitute of
processes, in which respect it differed from the same rib in the rock-
pigeon. In one Tumbler, and in the Bussorah Carrier, even the sixth pair
had no process. The hypapophysis of the second dorsal vertebra varies much
in development; being sometimes (as in several, but not all Tumblers)
nearly as prominent as that of the third dorsal vertebra; and the two
hypapophyses together tend to form an ossified arch. The development of the
arch, formed by the hypapophyses of the third and fourth dorsal vertebrae,
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 fourteen, and, as we
shall immediately see, an additional number of caudal vertebrae. In Runts
and Carriers there is generally the proper number, namely twelve; but in
one Runt, and in the Bussorah Carrier, there were only eleven. In Tumblers
there are either eleven, or twelve, or thirteen sacral vertebrae.
The CAUDAL VERTEBRAE are seven in number in the rock-pigeon. In Fantails,
which have their tails so largely developed, there are 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 vertebrae. I have seen eight in a Nun and Jacobin. Tumblers,
though such small birds, always have the normal number seven; as have
Carriers, with one exception, in which there were only six.
The following table will serve as a summary, and will show the most
remarkable deviations in the number of the vertebra and ribs which I have
observed:--
TABLE 4. NUMBER OF VERTEBRAE AND RIBS IN:
1. THE ROCK PIGEON.
Cervical Vertebrae: 12.
Dorsal Vertebrae: 8.
Dorsal Ribs: 8.
The 6th pair with processes, the 7th pair without a process.
Sacral Vertebrae: 12.
Caudal Vertebrae: 7.
Total: 39.
2. POUTER, FROM MR. BULT.
Cervical Vertebrae: 12.
Dorsal Vertebrae: 8.
Dorsal Ribs: 8.
The 6th and 7th pair with processes.
Sacral Vertebrae: 14.
Caudal Vertebrae: 8 or 9.
Total: 42 or 43.
3. TUMBLER, DUTCH ROLLER.
Cervical Vertebrae: 12.
Dorsal Vertebrae: 8.
Dorsal Ribs: 7.
The 6th and 7th pair without processes.
Sacral Vertebrae: 11.
Caudal Vertebrae: 7.
Total: 38.
4. BUSSORAH CARRIER.
Cervical Vertebrae: 12.
The twelfth bore a small rib.
Dorsal Vertebrae: 8.
Dorsal Ribs: 7.
The 6th and 7th pair without processes.
Sacral Vertebrae: 11.
Caudal Vertebrae: 7.
Total: 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.
(FIGURE 28. SCAPULAE, of natural size. A. Rock-pigeon. B. Short-faced
Tumbler.)
In the bones of the extremities I could detect no difference, except in
their proportional lengths; for instance, the metatarsus in a Pouter was
1.65 inch, and in a Short-faced Tumbler only .95 in length; and this is a
greater difference than would naturally follow from their differently-sized
bodies; but long legs in the Pouter, and small feet in the Tumbler, are
selected points. In some Pouters the SCAPULA is rather straighter, and in
some Tumblers it is straighter, with the apex less elongated, than in the
rock-pigeon: in figure 28, the scapula of the rock-pigeon (A), and of a
short-faced Tumbler (B), are given. The processes at the summit of the
CORACOID, which receive the extremities of the furculum, form a more
perfect cavity in some Tumblers than in the rock-pigeon: in Pouters these
processes are larger and differently shaped, and the exterior angle of the
extremity of the coracoid, which is articulated to the sternum, is squarer.
(FIGURE 29. FURCULA, of natural size. A. Short-faced Tumbler. B and C
Fantail. D. Pouter.)
The two arms of the FURCULUM in Pouters diverge less, proportionally to
their length, than in the rock-pigeon; and the symphysis is more solid and
pointed. In Fantails the degree of divergence of the two arms varies in a
remarkable manner. In figure 29, B and C represent the furcula of two
Fantails; and it will be seen that the divergence in B is rather less even
than in the furculum of the short-faced, small-sized Tumbler (A), whereas
the divergence in C equals that in a rock-pigeon, or in the Pouter (D),
though the latter is a much larger bird. The extremities of the furculum,
where articulated to the coracoids, vary considerably in outline.
In the STERNUM the differences in form are slight, except in the size and
outline of the perforations, which, both in the larger and lesser sized
breeds, are sometimes small. These perforations, also, are sometimes either
nearly circular, or elongated as is often the case with Carriers. The
posterior perforations occasionally are not complete, being left open
posteriorly. The marginal apophyses forming the anterior perforations vary
greatly in development. The degree of convexity of the posterior part of
the sternum differs much, being sometimes almost perfectly flat. The
manubrium is rather more prominent in some individuals than in others, and
the pore immediately under it varies greatly in size.]
CORRELATION OF GROWTH.
By this term I mean that the whole organisation is so connected, that when
one part varies, other parts vary; but which of two correlated variations
ought to be looked at as the cause and which as the effect, or whether both
result from some common cause, we can seldom or never tell. The point of
interest for us is that, when fanciers, by the continued selection of
slight variations, have largely modified one part, they often
unintentionally produce other modifications. For instance, the beak is
readily acted on by selection, and, with its increased or diminished
length, the tongue increases or diminishes, but not in due proportion; for,
in a Barb and Short-faced Tumbler, both of which have very short beaks, the
tongue, taking the rock-pigeon as the standard of comparison, was
proportionally not shortened enough, whilst in two Carriers and in a Runt
the tongue, proportionally with the beak, was not lengthened enough, thus,
in a first-rate English Carrier, in which the beak from the tip to the
feathered base was exactly thrice as long as in a first-rate Short-faced
Tumbler, the tongue was only a little more than twice as long. But the
tongue varies in length independently of the beak: thus in a Carrier with a
beak 1.2 inch in length, the tongue was .67 in length: whilst in a Runt
which equalled the Carrier in length of body and in stretch of wings from
tip to tip, the beak was .92 whilst the tongue was .73 of an inch in
length, so that the tongue was actually longer than in the carrier with its
long beak. The tongue of the Runt was also very broad at the root. Of two
Runts, one had its beak longer by .23 of an inch, whilst its tongue was
shorter by .14 than in the other.
With the increased or diminished length of the beak the length of the slit
forming the external orifice of the nostrils varies, but not in due
proportion, for, taking the rock-pigeon as the standard, the orifice in a
Short-faced Tumbler was not shortened in due proportion with its very short
beak. On the other hand (and this could not have been anticipated), the
orifice in three English Carriers, in the Bagadotten Carrier, and in a Runt
(pigeon cygne), was longer by above the tenth of an inch than would follow
from the length of the beak proportionally with that of the rock-pigeon. In
one Carrier the orifice of the nostrils was thrice as long as in the rock-
pigeon, though in body and length of beak this bird was not nearly double
the size of the rock-pigeon. This greatly increased length of the orifice
of the nostrils seems to stand partly in correlation with the enlargement
of the wattled skin on the upper mandible and over the nostrils; and this
is a character which is selected by fanciers. So again, the broad, naked,
and wattled skin round the eyes of Carriers and Barbs is a selected
character; and in obvious correlation with this, the eyelids, measured
longitudinally, are proportionally more than double the length of those of
the rock-pigeon.
The great difference (see figure 27) in the curvature of the lower jaw in
the rock-pigeon, the Tumbler, and Bagadotten Carrier, stands in obvious
relation to the curvature of the upper jaw, and more especially to the
angle formed by the maxillo-jugal arch with the premaxillary bones. But in
Carriers, Runts, and Barbs the singular reflexion of the upper margin of
the middle part of the lower jaw (see figure 25) is not strictly correlated
with the width or divergence (as may be clearly seen in figure 26) of the
premaxillary bones, but with the breadth of the horny and soft parts of the
upper mandible, which are always overlapped by the edges of the lower
mandible.
In Pouters, the elongation of the body is a selected character, and the
ribs, as we have seen, have generally become very broad, with the seventh
pair furnished with processes; the sacral and caudal vertebrae have been
augmented in number; the sternum has likewise increased in length (but not
in the depth of the crest) by .4 of an inch more than would follow from the
greater bulk of the body in comparison with that of the rock-pigeon. In
Fantails, the length and number of the caudal vertebrae have increased.
Hence, during the gradual progress of variation and selection, the internal
bony framework and the external shape of the body have been, to a certain
extent, modified in a correlated manner.
Although the wings and tail often vary in length independently of each
other, it is scarcely possible to doubt that they generally tend to become
elongated or shortened in correlation. This is well seen in Jacobins, and
still more plainly in Runts, some varieties of which have their wings and
tail of great length, whilst others have both very short. With Jacobins,
the remarkable length of the tail and wing-feathers is not a character
which is intentionally selected by fanciers; but fanciers have been trying
for centuries, at least since the year 1600, to increase the length of the
reversed feathers on the neck, so that the hood may more completely enclose
the head; and it may be suspected that the increased length of the wing and
tail-feathers stand in correlation with the increased length of the neck-
feathers. Short-faced Tumblers have short wings in nearly due proportion
with the reduced size of their bodies; but it is remarkable, seeing that
the number of the primary wing-feathers is a constant character in most
birds, that these Tumblers generally have only nine instead of ten
primaries. I have myself observed this in eight birds; and the Original
Columbarian Society (5/37. J.M. Eaton 'Treatise' edition 1858 page 78.)
reduced the standard for Bald-head Tumblers from ten to nine white flight-
feathers, thinking it unfair that a bird which had only nine feathers
should be disqualified for a prize because it had not ten WHITE flight-
feathers. On the other hand, in Carriers and Runts, which have large bodies
and long wings, eleven primary feathers have occasionally been observed.
Mr. Tegetmeier has informed me of a curious and inexplicable case of
correlation, namely, that young pigeons of all breeds which when mature
become white, yellow, silver (i.e., extremely pale blue), or dun-coloured,
are born almost naked; whereas pigeons of other colours are born well-
clothed with down. Mr. Esquilant, however, has observed that young dun
Carriers are not so bare as young dun Barbs and Tumblers. Mr. Tegetmeier
has seen two young birds in the same nest, produced from differently
coloured parents, which differed greatly in the degree to which they were
at first clothed with down.
I have observed another case of correlation which at first sight appears
quite inexplicable, but on which, as we shall see in a future chapter, some
light can be thrown by the law of homologous parts varying in the same
manner. The case is, that, when the feet are much feathered, the roots of
the feathers are connected by a web of skin, and apparently in correlation
with this the two outer toes become connected for a considerable space by
skin. I have observed this in very many specimens of Pouters, Trumpeters,
Swallows, Roller-tumblers (likewise observed in this breed by Mr. Brent),
and in a lesser degree in other feather-footed pigeons.
The feet of the smaller and larger breeds are of course much smaller or
larger than those of the rock-pigeon; but the scutellae 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
scutellae 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 scutellae on the
feet is usually a constant character. The length of the feet and the length
of the beak apparently stand in correlation; but as disuse apparently has
affected the size of the feet, this case may come under the following
discussion.
ON THE EFFECTS OF DISUSE.
In the following discussion on the relative proportions of the feet,
sternum, furculum, scapulae, and wings, I may premise, in order to give
some confidence to the reader, that all my measurements were made in the
same manner, and that they were made without the least intention of
applying them to the following purpose.
[TABLE 5.I.
PIGEONS WITH THEIR BEAKS GENERALLY SHORTER THAN THAT OF THE ROCK-PIGEON,
PROPORTIONALLY TO THE SIZE OF THEIR BODIES.
Column 1. Name of Breed.
Column 2. Actual length of Feet (inches).
Column 3. Difference between actual and calculated length of feet, in
proportion to length of feet and size of body in the Rock-pigeon.
Column 3a. Too short by (inches).
Column 3b. Too long by (inches).
1. 2. 3a. 3b.
Wild rock-pigeon (mean measurement). 2.02
Short-faced Tumbler, bald-head. 1.57 0.11 ..
Short-faced Tumbler, almond. 1.60 0.16 ..
Tumbler, red magpie. 1.75 0.19 ..
Tumbler, red common (by standard to end of tail). 1.85 0.07 ..
Tumbler, common bald-head. 1.85 0.18 ..
Tumbler, roller. 1.80 0.06 ..
Turbit. 1.75 0.17 ..
Turbit. 1.80 0.01 ..
Turbit. 1.84 0.15 ..
Jacobin. 1.90 0.02 ..
Trumpeter, white. 2.02 0.06 ..
Trumpeter, mottled. 1.95 0.18 ..
Fantail (by standard to end of tail). 1.85 0.15 ..
Fantail (by standard to end of tail). 1.95 0.15 ..
Fantail crested var. Ditto. 1.95 0.0 0.0
Indian Frill-back Ditto. 1.8O 0.19 ..
English Frill-back. 2.10 0.03 ..
Nun. 1.82 0.02 ..
Laugher. 1.65 0.16 ..
Barb. 2.00 0.03 ..
Barb. 2.00 ..
0.03
Spot. 1.90 0.02 ..
Spot. 1.90 0.07 ..
Swallow, red. 1.85 0.18 ..
Swallow, blue. 2.00 ..
0.03
Pouter. 2.42 ..
0.11
Pouter, German. 2.30 ..
0.09
Bussorah Carrier. 2.17 ..
0.09
Number of specimens. 28 22 5
I measured most of the birds which came into my possession, from the
feathered BASE of the beak (the length of beak itself being so variable) to
the end of the tail, and to the oil-gland, but unfortunately (except in a
few cases) not to the root of the tail; I measured each bird from the
extreme tip to tip of wing; and the length of the terminal folded part of
the wing, from the extremity of the primaries to the joint of the radius. I
measured the feet without the claws, from the end of the middle toe to the
end of the hind toe; and the tarsus and middle toe together. I have taken
in every case the mean measurement of two wild rock-pigeons from the
Shetland Islands, as the standard of comparison. The following table shows
the actual length of the feet in each bird; and the difference between the
length which the feet ought to have had according to the size of body of
each, in comparison with the size of body and length of feet of the rock-
pigeon, calculated (with a few specified exceptions) by the standard of the
length of the body from the base of the beak to the oil-gland. I have
preferred this standard, owing to the variability of the length of tail.
But I have made similar calculations, taking as the standard the length
from tip to tip of wing, and likewise in most cases from the base of the
beak to the end of the tail; and the result has always been closely
similar. To give an example: the first bird in the table, being a Short-
faced Tumbler, is much smaller than the rock-pigeon, and would naturally
have shorter feet; but it is found on calculation to have feet too short by
.11 of an inch, in comparison with the feet of the rock-pigeon, relatively
to the size of the body in these two birds, as measured from the base of
beak to the oil-gland. So again, when this same Tumbler and the rock-pigeon
were compared by the length of their wings, or by the extreme length of
their bodies, the feet of the Tumbler were likewise found to be too short
in very nearly the same proportion. I am well aware that the measurements
pretend to greater accuracy than is possible, but it was less trouble to
write down the actual measurements given by the compasses in each case than
an approximation.
TABLE 5.II.
PIGEONS WITH THEIR BEAKS LONGER THAN THAT OF THE ROCK-PIGEON,
PROPORTIONALLY TO THE SIZE OF THEIR BODIES.
Column 1. Name of Breed.
Column 2. Actual length of Feet (inches).
Column 3. Difference between actual and calculated length of feet, in
proportion to length of feet and size of body in the Rock-pigeon.
Column 3a. Too short by (inches).
Column 3b. Too long by (inches).
1. 2. 3a. 3b.
Wild rock-pigeon (mean measurement). 2.02
Carrier. 2.60 ..
0.31
Carrier. 2.60 ..
0.25
Carrier. 2.40 ..
0.21
Carrier, Dragon. 2.25 ..
0.06
Bagadotten Carrier. 2.80 ..
0.56
Scanderoon, white. 2.80 ..
0.37
Scanderoon, Pigeon cygne. 2.85 ..
0.29
Runt. 2.75 ..
0.27
Number of specimens. 8 .. 8
In these two tables (Tables 5.I.and 5.II.) we see in the first column the
actual length of the feet in thirty-six birds belonging to various breeds,
and in the two other columns we see by how much the feet are too short or
too long, according to the size of bird, in comparison with the rock-
pigeon. In the first table twenty-two specimens have their feet too short,
on an average by a little above the tenth of an inch (viz. .107); and five
specimens have their feet on an average a very little too long, namely, by
.07 of an inch. But some of these latter 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 besides the one 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 5.I. there are a few partial exceptions to the beak being
proportionally shorter than in the rock-pigeon: thus the beak of the
English Frill-back is just perceptibly longer, and that of the Bussorah
Carrier of the same length or slightly longer, than in the rock-pigeon. The
beaks of Spots, Swallows, and Laughers are only a very little shorter, or
of the same proportional length, but slenderer. Nevertheless, these two
tables, taken conjointly, indicate pretty plainly some kind of correlation
between the length of the beak and the size of the feet. Breeders of cattle
and horses believe that there is an analogous connection between the length
of the limbs and head; they assert that a race-horse with the head of a
dray-horse, or a grey-hound with the head of a bulldog, would be a
monstrous production. As fancy pigeons are generally kept in small
aviaries, and are abundantly supplied with food, they must walk about much
less than the wild rock-pigeon; and it may be admitted as highly probable
that the reduction in the size of the feet in the twenty-two birds in the
first table has been caused by disuse (5/38. In an analogous, but converse,
manner, certain natural groups of the Columbidae, from being more
terrestrial in their habits than other allied groups, have larger feet. See
Prince Bonaparte 'Coup d'oeil sur l'Ordre des Pigeons.'), and that this
reduction has acted by correlation on the beaks of the great majority of
the birds in Table 5.I. When, on the other hand, the beak has been much
elongated by the continued selection of successive slight increments of
length, the feet by correlation have likewise become much elongated in
comparison with those of the wild rock-pigeon, notwithstanding their
lessened use.
As I had taken measures from the end of the middle toe to the heel of the
tarsus in the rock-pigeon and in the above thirty-six birds, I have made
calculations analogous with those above given, and the result is the same--
namely, that in the short-beaked breeds, with equally few exceptions as in
the former case, the middle toe conjointly with the tarsus has decreased in
length; whereas in the long-beaked breeds it has increased in length,
though not quite so uniformly as in the former case, for the leg, in some
varieties of the Runt varies much in length.
LENGTH OF STERNUM.
As fancy pigeons are generally confined in aviaries of moderate size, and
as even when not confined they do not search for their own food, they must
during many generations have used their wings incomparably less than the
wild rock-pigeon. Hence it seemed to me probable that all the parts of the
skeleton subservient to flight would be found to be reduced in size. With
respect to the sternum, I have carefully measured its extreme length in
twelve birds of different breeds, and in two wild rock-pigeons from the
Shetland Islands. For the proportional comparison I have tried three
standards of measurement, with all twelve birds namely, the length from the
base of the beak to the oil-gland, to the end of the tail, and from the
extreme tip to tip of wings. The result has been in each case nearly the
same, the sternum being invariably found to be shorter than in the wild
rock-pigeon. I will give only a single table, as calculated by the standard
from the base of the beak to the oil-gland; for the result in this case is
nearly the mean between the results obtained by the two other standards.
TABLE 5.III.
LENGTH OF STERNUM.
Column 1. Name of Breed.
Column 2. Actual Length in Inches.
Column 3. Too short by (inches).
1. 2. 3.
Wild Rock-pigeon. 2.55
Pied Scanderoon. 2.80 0.60
Bagadotten Carrier. 2.80 0.17
Dragon. 2.45 0.41
Carrier. 2.75 0.35
Short-faced Tumbler. 2.05 0.28
Barb. 2.35 0.34
Nun. 2.27 0.15
German Pouter. 2.36 0.54
Jacobin. 2.33 0.22
English Frill-back. 2.40 0.43
Swallow. 2.45 0.17
This table (Table 5.III.) shows that in these twelve breeds the sternum is
of an average one-third of an inch (exactly .332) shorter than in the rock-
pigeon, proportionally with the size of their bodies; so that the sternum
has been reduced by between one-seventh and one-eighth of its entire
length; and this is a considerable reduction.
I have also measured in twenty-one birds, including the above dozen, the
prominence of the crest of the sternum relatively to its length,
independently of the size of the body. In two of the twenty-one birds the
crest was prominent in the same relative degree as in the rock-pigeon; in
seven it was more prominent; but in five out of these seven, namely, in a
Fantail, two Scanderoons, and two English Carriers, this greater prominence
may to a certain extent be explained, as a prominent breast is admired and
selected by fanciers; in the remaining twelve birds the prominence was
less. Hence it follows that the crest exhibits a slight, though uncertain,
tendency to be reduced in prominence in a greater degree than does the
length of the sternum relatively to the size of body, in comparison with
the rock-pigeon.
I have measured the length of the scapula in nine different large and
small-sized breeds, and in all the scapula is proportionally shorter
(taking the same standard as before) than in the wild rock-pigeon. The
reduction in length on an average is very nearly one-fifth of an inch, or
about one-ninth of the length of the scapula in the rock-pigeon.
The arms of the furcula in all the specimens which I compared, diverged
less, proportionally with the size of body, than in the rock-pigeon; and
the whole furculum was proportionally shorter. Thus in a Runt, which
measured from tip to tip of wings 38 1/2 inches, the furculum was only a
very little longer (with the arms hardly more divergent) than in a rock-
pigeon which measured from tip to tip 26 1/2 inches. In a Barb, which in
all its measurements was a little larger than the same rock-pigeon, the
furculum was a quarter of an inch shorter. In a Pouter, the furculum had
not been lengthened proportionally with the increased length of the body.
In a Short-faced Tumbler, which measured from tip to tip of wings 24
inches, therefore only 2 1/2 inches less than the rock-pigeon, the furculum
was barely two-thirds of the length of that of the rock-pigeon.]
We thus clearly see that the sternum, scapula, and furculum are all reduced
in proportional length; but when we turn to the wings we find what at first
appears a wholly different and unexpected result. I may here remark that I
have not picked out specimens, but have used every measurement made by me.
Taking the length from the base of beak to the end of the tail as the
standard of comparison, I find that, out of thirty-five birds of various
breeds, twenty-five have wings of greater, and ten have them of less
proportional length, than in the rock-pigeon. But from the frequently
correlated length of the tail and wing-feathers, it is better to take as
the standard of comparison the length from the base of the beak to the oil-
gland; and by this standard, out of twenty-six of the same birds which had
been thus measured, twenty-one had wings too long, and only five had them
too short. In the twenty-one birds the wings exceeded in length those of
the rock-pigeon, on an average, by 1 1/3 inch; whilst in the five birds
they were less in length by only .8 of an inch. As I was much surprised
that the wings of closely confined birds should thus so frequently have
been increased in length, it occurred to me that it might be solely due to
the greater length of the wing-feathers; for this certainly is the case
with the Jacobin, which has wings of unusual length. As in almost every
case I had measured the folded wings, I subtracted the length of this
terminal part from that of the expanded wings, and thus I obtained, with a
moderate degree of accuracy, the length of the wings from the ends of the
two radii, answering from wrist to wrist in our arms. The wings, thus
measured in the same twenty-five birds, now gave a widely different result;
for they were proportionally with those of the rock-pigeon too short in
seventeen birds, and in only eight too long. Of these eight birds, five
were long-beaked (5/39. It perhaps deserves notice that besides 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.), and this fact perhaps indicates that there is some
correlation of the length of the beak with the length of the bones of the
wings, in the same manner as with that of the feet and tarsi. The
shortening of the humerus and radius in the seventeen birds may probably be
attributed to disuse, as in the case of the scapula and furculum 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 scapula and
furculum, have all been reduced in size in comparison with the same parts
in the rock-pigeon. And I presume that this may be attributed to disuse or
lessened exercise. The wings, as measured from the ends of the radii, have
likewise been generally reduced in length; but, owing to the increased
growth of the wing-feathers, the wings, from tip to tip, are commonly
longer than in the rock-pigeon. The feet, as well as the tarsi conjointly
with the middle toe, have likewise in most cases become reduced; and this
it is probable has been caused by their lessened use; but the existence of
some sort of correlation between the feet and beak is shown more plainly
than the effects of disuse. We have also some faint indication of a similar
correlation between the main bones of the wing and the beak.
SUMMARY ON THE POINTS OF DIFFERENCE BETWEEN THE SEVERAL DOMESTIC RACES, AND
BETWEEN THE INDIVIDUAL BIRDS.
The beak, together with the bones of the face, differ remarkably in length,
breadth, shape, and curvature. The skull differs in shape, and greatly in
the angle formed by the union of the pre-maxillary, nasal, and maxillo-
jugal bones. The curvature of the lower jaw and the reflection of its upper
margin, as well as the gape of the mouth, differ in a highly remarkable
manner. The tongue varies much in length, both independently and in
correlation with the length of the beak. The development of the naked,
wattled skin over the nostrils and round the eyes varies in an extreme
degree. The eyelids and the external orifices of the nostrils vary in
length, and are to a certain extent correlated with the degree of
development of the wattle. The size and form of the oesophagus and crop,
and their capacity for inflation, differ immensely. The length of the neck
varies. With the varying shape of the body, the breadth and number of the
ribs, the presence of processes, the number of the sacral vertebrae, and
the length of the sternum, all vary. The number and size of the coccygeal
vertebrae 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 furculum, differ. The oil-gland varies in
development, and is sometimes quite aborted. The direction and length of
certain feathers have been much modified, as in the hood of the Jacobin and
the frill of the Turbit. The wing and tail-feathers generally vary in
length together, but sometimes independently of each other and of the size
of the body. The number and position of the tail-feather vary to an
unparalleled degree. The primary and secondary wing-feathers occasionally
vary in number, apparently in correlation with the length of the wing. The
length of the leg and the size of the feet, and, in connection with the
latter, the number of the scutellae, all vary. A web of skin sometimes
connects the bases of the two inner toes, and almost invariably the two
outer toes when the feet are feathered.
The size of the body differs greatly: a Runt has been known to weigh more
than five times as much as a Short-faced Tumbler. The eggs differ in size
and shape. According to Parmentier (5/40. Temminck 'Hist. Nat. Gen. des
Pigeons et des Gallinaces' tome 1 1813 page 170.), some races use much
straw in building their nests, and others use little; but I cannot hear of
any recent corroboration of this statement. The length of time required for
hatching the eggs is uniform in all the breeds. The period at which the
characteristic plumage of some breeds is acquired, and at which certain
changes of colour supervene, differs. The degree to which the young birds
are clothed with down when first hatched is different, and is correlated in
a singular manner with the colour of the plumage. The manner of flight, and
certain inherited movements, such as clapping the wings, tumbling either in
the air or on the ground, and the manner of courting the female, present
the most singular differences. In disposition the several races differ.
Some races are very silent; others coo in a highly peculiar manner.
Although many different races have kept true in character during several
centuries, as we shall hereafter more fully see, yet there is far more
individual variability in the most constant breeds than in birds in a state
of nature. There is hardly any exception to the rule that those characters
vary most which are now most valued and attended to by fanciers, and which
consequently are now being improved by continued selection. This is
indirectly admitted by fanciers when they complain that it is much more
difficult to breed high fancy pigeons up to the proper standard of
excellence than the so-called toy pigeons, which differ from each other
merely in colour; for particular colours when once acquired are not liable
to continued improvement or augmentation. Some characters become attached,
from quite unknown causes, more strongly to the male than to the female
sex; so that we have in certain races, a tendency towards the appearance of
secondary sexual characters (5/41. This term was used by John Hunter for
such differences in structure between the males and females, as are not
directly connected with the act of reproduction, as the tail of the
peacock, the horns of deer, etc.) of which the aboriginal rock-pigeon
displays not a trace.
CHAPTER 1.VI.
PIGEONS--continued.
ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES.
HABITS OF LIFE.
WILD RACES OF THE ROCK-PIGEON.
DOVECOTE-PIGEONS.
PROOFS OF THE DESCENT OF THE SEVERAL RACES FROM COLUMBA LIVIA.
FERTILITY OF THE RACES WHEN CROSSED.
REVERSION TO THE PLUMAGE OF THE WILD ROCK-PIGEON.
CIRCUMSTANCES FAVOURABLE TO THE FORMATION OF THE RACES.
ANTIQUITY AND HISTORY OF THE PRINCIPAL RACES.
MANNER OF THEIR FORMATION.
SELECTION.
UNCONSCIOUS SELECTION.
CARE TAKEN BY FANCIERS IN SELECTING THEIR BIRDS.
SLIGHTLY DIFFERENT STRAINS GRADUALLY CHANGE INTO WELL-MARKED BREEDS.
EXTINCTION OF INTERMEDIATE FORMS.
CERTAIN BREEDS REMAIN PERMANENT, WHILST OTHERS CHANGE.
SUMMARY.
The differences described in the last chapter between the eleven chief
domestic races and between individual birds of the same race, would be of
little significance, if they had not all descended from a single wild
stock. The question of their origin is therefore of fundamental importance,
and must be discussed at considerable length. No one will think this
superfluous who considers the great amount of difference between the races,
who knows how ancient many of them are, and how truly they breed at the
present day. Fanciers almost unanimously believe that the different races
are descended from several wild stocks, whereas most naturalists believe
that all are descended from the Columba livia or rock-pigeon.
Temminck (6/1. Temminck 'Hist. Nat. Gen. des Pigeons' etc. tome 1 page
191.) has well observed, and Mr. Gould has made the same remark to me, that
the aboriginal parent must have been a species which roosted and built its
nest on rocks; and I may add that it must have been a social bird. For all
the domestic races are highly social, and none are known to build or
habitually to roost on trees. The awkward manner in which some pigeons,
kept by me in a summer-house near an old walnut-tree, occasionally alighted
on the barer branches, was evident. (6/2. 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
disturbed in their loft by their young being taken, roosted on them at
night.) Nevertheless, Mr. R. Scot Skirving informs me that he often saw
crowds of pigeons in Upper Egypt settling on low trees, but not on palms,
in preference to alighting on the mud hovels of the natives. In India Mr.
Blyth (6/3. 'Annals and Mag. of Nat. Hist.' 2nd series volume 20 1857 page
509; and in a late volume of the 'Journal of the Asiatic Society.') has
been assured that the wild C. livia, var. intermedia, sometimes roosts in
trees. I may here give a curious instance of compulsion leading to changed
habits: the banks of the Nile above lat. 28 deg 30' are perpendicular for a
long distance, so that when the river is full the pigeons cannot alight on
the shore to drink, and Mr. Skirving repeatedly saw whole flocks settle on
the water, and drink whilst they floated down the stream. These flocks seen
from a distance resembled flocks of gulls on the surface of the sea.
If any domestic race had descended from a species which was not social, or
which built its nest and roosted in trees (6/4. In works written on the
pigeon by fanciers I have sometimes observed the mistaken belief expressed
that the species which naturalists called ground-pigeons (in
contradistinction to arboreal pigeons) do not perch and build on trees. In
these same works by fanciers wild species resembling the chief domestic
races are often said to exist in various parts of the world; but such
species are quite unknown to naturalists.) the sharp eyes of fanciers would
assuredly have detected some vestige of so different an aboriginal habit.
For we have reason to believe that aboriginal habits are long retained
under domestication. Thus with the common ass we see signs of its original
desert life in its strong dislike to cross the smallest stream of water,
and in its pleasure in rolling in the dust. The same strong dislike to
cross a stream is common to the camel, which has been domesticated from a
very ancient period. Young pigs, though so tame, sometimes squat when
frightened, and thus try to conceal themselves even on an open and bare
place. Young turkeys, and occasionally even young fowls, when the hen gives
the danger-cry, run away and try to hide themselves, like young partridges
or pheasants, in order that their mother may take flight, of which she has
lost the power. The musk-duck (Cairina moschata) in its native country
often perches and roosts on trees (6/5. Sir R. Schomburgk in 'Journal R.
Geograph. Soc.' volume 13 1844 page 32.), and our domesticated musk-ducks,
though such sluggish birds, "are fond of perching on the tops of barns,
walls, etc., 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." (6/6. Rev. E.S. Dixon 'Ornamental
Poultry' 1848 pages 63, 66.) We know that the dog, however well and
regularly fed, often buries, like the fox, any superfluous food; and we see
him turning round and round on a carpet, as if to trample down grass to
form a bed; we see him on bare pavements scratching backwards as if to
throw earth over his excrement, although, as I believe, this is never
effected even where there is earth. In the delight with which lambs and
kids crowd together and frisk on the smallest hillock, we see a vestige of
their former alpine habits.
We have therefore good reason to believe that all the domestic races of the
pigeon are descended either from some one or from several species which
both roosted and built their nests on rocks, and were social in
disposition. As only five or six wild species have these habits, and make
any near approach in structure to the domesticated pigeon, I will enumerate
them.
[Firstly, the Columba leuconota resembles certain domestic varieties in its
plumage, with the one marked and never-failing difference of a white band
which crosses the tail at some distance from the extremity. This species,
moreover, inhabits the Himalaya, close to the limit of perpetual snow; and
therefore, as Mr. Blyth has remarked, is not likely to have been the parent
of our domestic breeds, which thrive in the hottest countries. Secondly,
the C. rupestris, of Central Asia, which is intermediate (6/7. 'Proc.
Zoolog. Soc.' 1859 page 400.) between the C. leuconota and livia; but has
nearly the same coloured tail as the former species. Thirdly, the Columba
littoralis builds and roosts, according to Temminck, on rocks in the
Malayan archipelago; it is white, excepting parts of the wing and the tip
of the tail, which are black; its legs are livid-coloured, and this is a
character not observed in any adult domestic pigeon; but I need not have
mentioned this species or the closely-allied C. luctuosa, as they in fact
belong to the genus Carpophaga. Fourthly, Columba guinea, which ranges from
Guinea (6/8. Temminck 'Hist. Nat. Gen. des Pigeons' tome 1; also 'Les
Pigeons' par Mme. Knip and Temminck. Bonaparte, however, in his 'Coup-
d'oeil' believes that two closely allied species are confounded together
under this name. The C. leucocephala 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.) 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
Strictoenas of Reichenbach, but is closely allied to 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 characterised by the feathers of the
neck having peculiar notched tips,--a character not observed in any
domestic race. Fifthly, the Columba oenas of Europe, which roosts on trees,
and builds its nest in holes, either in trees or the ground; this species,
as far as external characters go, might be the parent of several domestic
races; but, though it crosses readily with the true rock-pigeon, the
offspring, as we shall presently see, are sterile hybrids, and of such
sterility there is not a trace when the domestic races are intercrossed. It
should also be observed that if we were to admit, against all probability,
that any of the foregoing five or six species were the parents of some of
our domestic pigeons, not the least light would be thrown on the chief
differences between the eleven most strongly-marked races.
We now come to the best known rock-pigeon, the Columba livia, which is
often designated in Europe pre-eminently as the Rock-pigeon, and which
naturalists believe to be the parent of all the domesticated breeds. This
bird agrees in every essential character with the breeds which have been
only slightly modified. It differs from all other species in being of a
slaty-blue colour, with two black bars on the wings, and with the croup (or
loins) white. Occasionally birds are seen in Faroe and the Hebrides with
the black bars replaced by two or three black spots; this form has been
named by Brehm (6/9. 'Handbuch der Naturgesch. Vogel Deutschlands.') C.
amaliae, but this species has not been admitted as distinct by other
ornithologists. Graba (6/10. 'Tagebuch, Reise nach Faro' 1830 s. 62.) even
found a difference in the bars on the right and left wings of the same bird
in Faroe. Another and rather more distinct form is either truly wild or has
become feral on the cliffs of England and was doubtfully named by Mr. Blyth
(6/11. 'Annals and Mag. of Nat. Hist.' volume 19 1847 page 102. This
excellent paper on pigeons is well worth consulting.) as C. affinis, but is
now no longer considered by him as a distinct species. C. affinis is rather
smaller than the rock-pigeon of the Scottish islands, and has a very
different appearance owing to the wing-coverts being chequered with black,
with similar marks often extending over the back. The chequering consists
of a large black spot on the two sides, but chiefly on the outer side, of
each feather. The wing-bars in the true rock-pigeon and in the chequered
variety are, in fact, due to similar though larger spots symmetrically
crossing the secondary wing-feather and the larger coverts. Hence the
chequering arises merely from an extension of these marks to other parts of
the plumage. Chequered birds are not confined to the coasts of England; for
they were found by Graba at Faroe; and W. Thompson (6/12. 'Natural History
of Ireland' Birds volume 2 1850 page 11. For Graba see previous reference.)
says that at Islay fully half the wild rock-pigeons were chequered. Colonel
King, of Hythe, stocked his dovecote with young wild birds which he himself
procured from nests at the Orkney Islands; and several specimens, kindly
sent to me by him, were all plainly chequered. As we thus see that
chequered birds occur mingled with the true rock-pigeon at three distinct
sites, namely, Faroe, the Orkney Islands, and Islay, no importance can be
attached to this natural variation in the plumage.
Prince C.L. Bonaparte (6/13. 'Coup-d'oeil sur l'Ordre des Pigeons' 'Comptes
Rendus' 1854-55.), a great divider of species, enumerates, with a mark of
interrogation, as distinct from C. livia, the C. turricola of Italy, the C.
rupestris of Daouria, and the C. schimperi of Abyssinia; but these birds
differ from C. livia in characters of the most trifling value. In the
British Museum there is a chequered pigeon, probably the C. schimperi of
Bonaparte, from Abyssinia. To these may be added the C. gymnocyclus of G.R.
Gray from W. Africa, which is slightly more distinct, and has rather more
naked skin round the eyes than the rock-pigeon; but from information given
me by Dr. Daniell, it is doubtful whether this is a wild bird, for
dovecote-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 differs chiefly in the croup
being blue instead of snow-white; but as Mr. Blyth informs me, the tint
varies, being sometimes albescent. When this form is domesticated chequered
birds appear, just as occurs in Europe with the truly wild C. livia.
Moreover we shall immediately have proof that the blue and white croup is a
highly variable character; and Bechstein (6/14. 'Naturgeschichte
Deutschlands' b. 4 1795 s. 14.) asserts that with dovecote-pigeons in
Germany this is the most variable of all the characters of the plumage.
Hence it may be concluded that C. intermedia cannot be ranked as
specifically distinct from C. livia.
In Madeira there is a rock-pigeon which a few ornithologists have suspected
to be distinct from C. livia. I have examined numerous specimens collected
by Mr. E.V. Harcourt and Mr. Mason. They are rather smaller than the rock-
pigeon from the Shetland Islands, and their beaks are plainly thinner, but
the thickness of the beak varied in the several specimens. In plumage there
is remarkable diversity; some specimens are identical in every feather (I
speak after actual comparison) with the rock-pigeon of the Shetland
Islands; others are chequered, like C. affinis from the cliffs of England,
but generally to a greater degree, being almost black over the whole back;
others are identical with the so-called C. intermedia of India in the
degree of blueness of the croup; whilst others have this part very pale or
very dark blue, and are likewise chequered. So much variability raises a
strong suspicion that these birds are domestic pigeons which have become
feral.
From these facts it can hardly be doubted that C. livia, affinis,
intermedia, and the forms marked with an interrogation by Bonaparte ought
all to be included under a single species. But it is quite immaterial
whether or not they are thus ranked, and whether some one of these forms or
all are the progenitors of the various domestic kinds, as far as any light
can thus be thrown on the differences between the more strongly-marked
races. That common dovecote-pigeons, which are kept in various parts of the
world, are descended from one or from several of the above-mentioned wild
varieties of C. livia, no one who compares them will doubt. But before
making a few remarks on dovecote-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 dovecote more than twenty years ago
with young wild birds taken at the Orkney Islands, and since then they have
greatly multiplied. The accurate Macgillivray (6/15. 'History of British
Birds' volume 1 pages 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-pigeon can be easily tamed; and the former gentleman
asserts that the tamed birds breed four times a year. Dr. Lawrence
Edmondstone informs me that a wild rock-pigeon came and settled in his
dovecote 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.) asserts that he completely tamed a
wild rock-pigeon in the Hebrides; and several accounts are on records of
these pigeons having bred in dovecotes 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 (6/16. 'Annals and
Mag. of Nat. History' volume 19 1847 page 103 and volume for 1857 page
512.) asserts that wild birds come frequently to the dovecotes and mingle
freely with their inhabitants. In the ancient 'Ayeen Akbery' it is written
that, if a few wild pigeons be taken, "they are speedily joined by a
thousand others of their kind."
Dovecote-pigeons are those which are kept in dovecotes 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 dovecote-
pigeon exactly resembles the chequered variety of C. livia; but I have seen
dovecotes brought from Yorkshire without any trace of chequering, like the
wild rock-pigeon of the Shetland Islands. The chequered dovecotes from the
Orkney Islands, after having been domesticated by Colonel King for more
than twenty years, differed slightly from each other in the darkness of
their plumage and in the thickness of their beaks; the thinnest beak being
rather thicker than the thickest one in the Madeira birds. In Germany,
according to Bechstein, the common dovecote-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 dovecote-pigeons, which originally came from the
S. Natunas Islands in the Malay Archipelago, and which had been crossed
with the Singapore dovecotes: they were small and the darkest variety was
extremely like the dark chequered variety with a blue croup from Madeira;
but the beak was not so thin, though decidedly thinner than in the rock-
pigeon from the Shetland Islands. A dovecote-pigeon sent to me by Mr.
Swinhoe from Foochow, in China, was likewise rather small, but differed in
no other respect. I have also received through the kindness of Dr. Daniell,
four living dovecote-pigeons from Sierra Leone (6/17. Domestic pigeons of
the common kind are mentioned as being pretty numerous in John Barbut's
'Description of the Coast of Guinea' page 215 published in 1746; they are
said, in accordance with the name which they bear, to have been imported.)
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 dovecote. When the beaks of
these African pigeons were compared with the thinnest beaks of the wild
Madeira specimens, the contrast was great; the former being fully one-third
thicker in a vertical direction than the latter; so that any one at first
would have felt inclined to rank these birds as specifically distinct; yet
so perfectly graduated a series could be formed between the above-mentioned
varieties, that it was obviously impossible to separate them.]
To sum up: the wild Columba livia, including under this name C. affinis,
intermedia, and the other still more closely-affined geographical races,
has a vast range from the southern coast of Norway and the Faroe Islands to
the shores of the Mediterranean, to Madeira and the Canary Islands, to
Abyssinia, India, and Japan. It varies greatly in plumage, being in many
places chequered with black, and having either a white or blue croup or
loins; it varies also slightly in the size of the beak and body. Dovecote-
pigeons, which no one disputes are descended from one or more of the above
wild forms, present a similar but greater range of variation in plumage, in
the size of body, and in the length and thickness of the beak. There seems
to be some relation between the croup being blue or white, and the
temperature of the country inhabited by both wild and dovecote pigeons; for
nearly all the dovecote-pigeons in the northern parts of Europe have a
white croup, like that of the wild European rock-pigeon; and nearly all the
dovecote-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 dovecote-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, etc., are
descended from distinct aboriginal stocks, yet admit that the so-called
toy-pigeons, which differ from the rock-pigeon in little except colour, are
descended from this bird. By toy-pigeons are meant such birds as Spots,
Nuns, Helmets, Swallows, Priests, Monks, Porcelains, Swabians, Archangels,
Breasts, Shields, and others in Europe, and many others in India. It would
indeed be as puerile to suppose that all these birds are descended from so
many distinct wild stocks as to suppose this to be the case with the many
varieties of the gooseberry, heartsease, or dahlia. Yet these kinds all
breed true, and many of them include sub-varieties which likewise transmit
their character truly. 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 dovecote-pigeons. Although we may
safely admit that dovecote-pigeons, which vary slightly, and that toy-
pigeons, which vary in a greater degree in accordance with their more
highly-domesticated condition, are descended from C. livia, including under
this name the above-enumerated wild geographical races; yet the question
becomes far more difficult when we consider the eleven principal races,
most of which have been profoundly modified. It can, however, be shown, by
indirect evidence of a perfectly conclusive nature, that these principal
races are not descended from so many wild stocks; and if this be once
admitted, few will dispute that they are the descendants of C. livia, which
agrees with them so closely in habits and in most characters, which varies
in a state of nature, and which has certainly undergone a considerable
amount of variation, as in the toy-pigeons. We shall moreover presently see
how eminently favourable circumstances have been for a great amount of
modification in the more carefully tended breeds.
The reasons for concluding that the several principal races are not
descended from so many aboriginal and unknown stocks may be grouped under
the following six heads:--
FIRSTLY.
If the eleven chief races have not arisen from the variation of some one
species, together with its geographical races, they must be descended from
several extremely distinct aboriginal species; for no amount of crossing
between only six or seven wild forms could produce races so distinct as
Pouters, Carriers, Runts, Fantails, Turbits, Short-faced Tumblers,
Jacobins, and Trumpeters. How could crossing produce, for instance, a
Pouter or a Fantail, unless the two supposed aboriginal parents possessed
the remarkable characters of these breeds? I am aware that some
naturalists, following Pallas, believe that crossing gives a strong
tendency to variation, independently of the characters inherited from
either parent. They believe that it would be easier to raise a Pouter or
Fantail pigeon from crossing two distinct species, neither of which
possessed the characters of these races, than from any single species. I
can find few facts in support of this doctrine, and believe in it only to a
limited degree; but in a future chapter I shall have to recur to this
subject. For our present purpose the point is not material. The question
which concerns us is, whether or not many new and important characters have
arisen since man first domesticated the pigeon. On the ordinary view,
variability is due to changed conditions of life; on the Pallasian
doctrine, variability, or the appearance of new characters, is due to some
mysterious effect from the crossing of two species, neither of which
possesses the characters in question. In some few instances it is possible
that well-marked races may have been formed by crossing; for instance, a
Barb might perhaps be formed by a cross between a long-beaked Carrier,
having large eye-wattles, and some short-beaked pigeon. That many races
have been in some degree modified by crossing, and that certain varieties
which are distinguished only by peculiar tints have arisen from crosses
between differently-coloured varieties, is almost certain. On the doctrine,
therefore, that the chief races owe their differences to their descent from
distinct species, we must admit that at least eight or nine, or more
probably a dozen species, all having the same habit of breeding and
roosting on rocks and living in society, either now exist somewhere, or
formerly existed, but have become extinct as wild birds. Considering how
carefully wild pigeons have been collected throughout the world, and what
conspicuous birds they are, especially when frequenting rocks, it is
extremely improbable that eight or nine species, which were long ago
domesticated and therefore must have inhabited some anciently known
country, should still exist in the wild state and be unknown to
ornithologists.
The hypothesis that such species formerly existed, but have become extinct,
is in some slight degree more probable. But the extinction of so many
species within the historical period is a bold hypothesis, seeing how
little influence man has had in exterminating the common rock-pigeon, which
agrees in all its habits of life with the domestic races. The C. livia now
exists and flourishes on the small northern islands of Faroe, on many
islands off the coast of Scotland, on Sardinia, and the shores of the
Mediterranean, and in the centre of India. Fanciers have sometimes imagined
that the several supposed parent-species were originally confined to small
islands, and thus might readily have been exterminated; but the facts just
given do not favour the probability of their extinction, even on small
islands. Nor is it probable, from what is known of the distribution of
birds, that the islands near Europe should have been inhabited by peculiar
species of pigeons; and if we assume that distant oceanic islands were the
homes of the supposed parent-species, we must remember that ancient voyages
were tediously slow, and that ships were then ill-provided with fresh food,
so that it would not have been easy to bring home living birds. I have said
ancient voyages, for nearly all the races of the pigeon were known before
the year 1600, so that the supposed wild species must have been captured
and domesticated before that date.
SECONDLY.
The doctrine that the chief domestic races are descended from several
aboriginal species, implies that several species were formerly so
thoroughly domesticated as to breed readily when confined. Although it is
easy to tame most wild birds, experience shows us that it is difficult to
get them to breed freely under confinement; although it must be owned that
this is less difficult with pigeons than with most other birds. During the
last two or three hundred years, many birds have been kept in aviaries, but
hardly one has been added to our list of thoroughly reclaimed species: yet
on the above doctrine we must admit that in ancient times nearly a dozen
kinds of pigeons, now unknown in the wild state, were thoroughly
domesticated.
THIRDLY.
Most of our domesticated animals have run wild in various parts of the
world; but birds, owing apparently to their partial loss of the power of
flight, less often than quadrupeds. Nevertheless I have met with accounts
showing that the common fowl has become feral in South America and perhaps
in West Africa, and on several islands: the turkey was at one time almost
feral on the banks of the Parana; and the Guinea-fowl has become perfectly
wild at Ascension and in Jamaica. In this latter island the peacock, also,
"has become a maroon bird." The common duck wanders from its home and
becomes almost wild in Norfolk. Hybrids between the common and musk-duck
which have become wild have been shot in North America, Belgium, and near
the Caspian Sea. The goose is said to have run wild in La Plata. The common
dovecote-pigeon has become wild at Juan Fernandez, Norfolk Island,
Ascension, probably at Madeira, on the shores of Scotland, and, as is
asserted, on the banks of the Hudson in North America. (6/18. With respect
to feral pigeons--for Juan Fernandez see Bertero in 'Annal. des Sc. Nat.'
tome 21 page 351. For Norfolk Islands see Rev. E.S. Dixon in the 'Dovecote'
1851 page 14 on the authority of Mr. Gould. For Ascension I rely on MS.
information given me by Mr. Layard. For the banks of the Hudson, see Blyth
in 'Annals of Nat. Hist.' volume 20 1857 page 511. For Scotland see
Macgillivray 'British Birds' volume 1 page 275; also Thompson 'Nat. Hist.
of Ireland, Birds' volume 2 page 11. For ducks see Rev. E.S. Dixon
'Ornamental Poultry' 1847 page 122. For the feral hybrids of the common and
musk-ducks see Audubon 'American Ornithology' and Selys-Longchamp 'Hybrides
dans la Famille des Anatides.' For the goose Isidore Geoffroy St.-Hilaire
'Hist. Nat. Gen.' tome 3 page 498. For guinea-fowls see Gosse 'Naturalist's
Sojourn in Jamaica' page 124; and his 'Birds of Jamaica' for fuller
particulars. I saw the wild guinea-fowl in Ascension. For the peacock see
'A Week at Port Royal' by a competent authority, Mr. R. Hill, page 42. For
the turkey I rely on oral information; I ascertained that they were not
Curassows. With respect to fowls I will give the references in the next
chapter.) But how different is the case, when we turn to the eleven chief
domestic races of the pigeon, which are supposed by some authors to be
descended from so many distinct species! no one has ever pretended that any
one of these races has been found wild in any quarter of the world; yet
they have been transported to all countries, and some of them must have
been carried back to their native homes. On the view that all the races are
the product of variation, we can understand why they have not become feral,
for the great amount of modification which they have undergone shows how
long and how thoroughly they have been domesticated; and this would unfit
them for a wild life.
FOURTHLY.
If it be assumed that the characteristic differences between the various
domestic races are due to descent from several aboriginal species, we must
conclude that man chose for domestication in ancient times, either
intentionally or by chance, a most abnormal set of pigeons; for that
species resembling such birds as Pouters, Fantails, Carriers, Barbs, Short-
faced Tumblers, Turbits, etc., would be in the highest degree abnormal, as
compared with all the existing members of the great pigeon family, cannot
be doubted. Thus we should have to believe that man not only formerly
succeeded in thoroughly domesticating several highly abnormal species, but
that these same species have since all become extinct, or are at least now
unknown. This double accident is so extremely improbable that the assumed
existence of so many abnormal species would require to be supported by the
strongest evidence. On the other hand, if all the races are descended from
C. livia, we can understand, as will hereafter be more fully explained, how
any slight deviation in structure which first appeared would continually be
augmented by the preservation of the most strongly marked individuals; and
as the power of selection would be applied according to man's fancy, and
not for the bird's own good, the accumulated amount of deviation would
certainly be of an abnormal nature in comparison with the structure of
pigeons living in a state of nature.
I have already alluded to the remarkable fact that the characteristic
differences between the chief domestic races are eminently variable; we see
this plainly in the great difference in the number of the tail-feathers in
the Fantail, in the development of the crop in Pouters, in the length of
the beak in Tumblers, in the state of the wattle in Carriers, etc. If these
characters are the result of successive variations added together by
selection, we can understand why they should be so variable: for these are
the very parts which have varied since the domestication of the pigeon, and
therefore would be likely still to vary; these variations moreover have
been recently, and are still being accumulated by man's selection;
therefore they have not as yet become firmly fixed.
FIFTHLY.
All the domestic races pair readily together, and, what is equally
important, their mongrel offspring are perfectly fertile. To ascertain this
fact I made many experiments, which are given in the note below; and
recently Mr. Tegetmeier has made similar experiments with the same result.
(6/19. 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
while publishing. 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 undoubtedly have thus united all.
The case of five distinct breeds being blended together with unimpaired
fertility is important, because Gartner has shown that it is a very
general, though not, as he thought, universal rule, that complex 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. Pistor ('Das Ganze der Feldtaubenzucht' 1831 s. 15) asserts that
the mongrels from Barbs and Fantails are sterile: I have proved this to be
erroneous, not only by crossing those 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. Gen. des Pigeons' tome 1 page 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 Dovecote' page 107) between Turbits and
Dovecotes and Nuns. I have crossed Turbits with Barbs, as has M. Boitard
(page 34), 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. 4 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 occasionally sterile.) The
accurate Neumeister asserts that when dovecotes are crossed with pigeons of
any other breed, the mongrels are extremely fertile and hardy. (6/20. 'Das
Ganze der Taubenzucht' s. 18.) MM. Boitard and Corbie (6/21. 'Les Pigeons'
etc. page 35.) affirm, after their great experience, that the more distinct
the breeds are which are crossed, the more productive are their mongrel
offspring. I admit that the doctrine first broached by Pallas is highly
probable, if not actually proved, namely, that closely allied species,
which in a state of nature or when first captured would have been in some
degree sterile if crossed, lose this sterility after a long course of
domestication; yet when we consider the great difference between such races
as Pouters, Carriers, Runts, Fantails, Turbits, Tumblers etc., 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 (6/22. Domestic pigeons pair readily with
the allied C. oenas (Bechstein 'Naturgesch. Deutschlands' b. 4 s. 3); and
Mr. Brent has made the same cross several 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) paired with a Dragon, but never
laid eggs. Bechstein further states (s. 26) that the domestic pigeon will
cross with C. palumbus, Turtur risoria and T. vulgaris, 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.' volume 7 1834 page 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 female T. risoria, and the latter
laid many eggs, but all were sterile. MM. Boitard and Corbie ('Les Pigeons'
page 235) state that the hybrids from these two turtle-doves are invariably
sterile both inter se and with either pure parent. The experiment was tried
by M. Corbie "avec une espece d'obstination;" and likewise by M. Mauduyt,
and by M. Vieillot. Temminck also found the hybrids from these two species
quite barren. Therefore, when Bechstein ('Naturgesch. Deutschlands Vogel'
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 November 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, and from T. vulgaris and
Ectopistes migratorius, were sterile. Two of the latter male hybrids paired
with their pure parents, 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.
Geoffrey Saint-Hilaire 'Hist. Nat. Generale' tome 3 page 180) from Turtur
auritus with T. cambayensis and with T. suratensis; but nothing is said of
their fertility. At the Zoological Gardens of London the Goura coronata and
victoriae produced a hybrid which paired with the pure G. coronata, and
laid several eggs, but these proved barren. In 1860 Columba gymnophthalmos
and maculosa produced hybrids in these same gardens.) all the cases which I
have collected) that hardly a single well-ascertained instance is known of
hybrids between two true species of pigeons being fertile, inter se, or
even when crossed with one of their pure parents.
SIXTHLY.
Excluding certain important characteristic differences, the chief races
agree most closely both with each other and with C. livia in all other
respects. As previously observed, all are eminently sociable; all dislike
to perch or roost, and refuse to build in trees; all lay two eggs, and this
is not a universal rule with the Columbidae; all, as far as I can hear,
require the same time for hatching their eggs; all can endure the same
great range of climate; all prefer the same food, and are passionately fond
of salt; all exhibit (with the asserted exception of the Finnikin and
Turner which do not differ much in any other character) the same peculiar
gestures when courting the females; and all (with the exception of
Trumpeters and Laughers, which likewise do not differ much in any other
character) coo in the same peculiar manner, unlike the voice of any other
wild pigeon. All the coloured breeds display the same peculiar metallic
tints on the breast, a character far from general with pigeons. Each race
presents nearly the same range of variation in colour; and in most of the
races we have the same singular correlation between the development of down
in the young and the future colour of plumage. All have the proportional
length of their toes, and of their primary wing-feathers, nearly the same,-
-characters which are apt to differ in the several members of the
Columbidae. In those races which present some remarkable deviation of
structure, such as in the tail of Fantails, crop of Pouters, beak of
Carriers and Tumblers, etc., the other parts remain nearly unaltered. Now
every naturalist will admit that it would be scarcely possible to pick out
a dozen natural species in any family which should agree closely in habits
and in general structure, and yet should differ greatly in a few characters
alone. This fact is explicable through the doctrine of natural selection;
for each successive modification of structure in each natural species is
preserved, solely because it is of service; and such modifications when
largely accumulated imply a great change in the habits of life, and this
will almost certainly lead to other changes of structure throughout the
whole organisation. On the other hand, if the several races of the pigeon
have been produced by man through selection and variation, we can readily
understand how it is that they should still all resemble each other in
habits and in those many characters which man has not cared to modify,
whilst they differ to so prodigious a degree in those parts which have
struck his eye or pleased his fancy.
Besides the points above enumerated, in which all the domestic races
resemble C. livia and each other, there is one which deserves special
notice. The wild rock-pigeon is of a slaty-blue colour; the wings are
crossed by two bars; the croup varies in colour, being generally white in
the pigeon of Europe, and blue in that of India; the tail has a black bar
close to the end, and the outer webs of the outer tail-feathers are edged
with white, except near the tips. These combined characters are not found
in any wild pigeon besides C. livia. I have looked carefully through the
great collections of pigeons in the British Museum, and I find that a dark
bar at the end of the tail is common; that the white edging to the outer
tail-feathers is not rare; but that the white croup is extremely rare, and
the two black bars on the wings occur in no other pigeon, excepting the
alpine C. leuconota and C. rupestris of Asia. Now if we turn to the
domestic races, it is highly remarkable, as an eminent fancier, Mr.
Wicking, observed to me, that, whenever a blue bird appears in any race,
the wings almost invariably show the double black bars. (6/23. 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
signifies 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.) The primary wing-
feathers may be white or black, and the whole body may be of any colour,
but if the wing-coverts are blue, the two black bars are sure to appear. I
have myself seen, or acquired trustworthy evidence, as given below (6/24. I
have observed blue birds with all the above-mentioned marks in the
following races, which seemed to be perfectly 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
croup in some was bluish or pure blue. Mr. Wicking bred blue Fantails from
two black birds. Carriers (including the Bagadotten of Neumeister) 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, assures me that, if black carriers are matched for
many successive generations, 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 instances 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 excessively 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.), of blue
birds with black bars on the wing, with the croup either white or very pale
or dark blue, with the tail having a terminal black bar, and with the outer
feathers externally edged with white or very pale coloured, in the
following races, which, as I carefully observed in each case, appeared to
be perfectly true: namely, in Pouters, Fantails, Tumblers, Jacobins,
Turbits, Barbs, Carriers, Runts of three distinct varieties, Trumpeters,
Swallows, and in many other toy-pigeons, which as being closely allied to
C. livia, are not worth enumerating. Thus we see that, in purely-bred races
of every kind known in Europe, blue birds occasionally appear having all
the marks which characterise C. livia, and which concur in no other wild
species. Mr. Blyth, also, has made the same observation with respect to the
various domestic races known in India.
Certain variations in the plumage are equally common in the wild C. livia,
in dovecote-pigeons, and in all the most highly modified races. Thus, in
all, the croup varies from white to blue, being most frequently white in
Europe, and very generally blue in India. (6/25. 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 white, 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.) We
have seen that the wild C. livia in Europe, and dovecotes in all parts of
the world, often have the upper wing-coverts chequered with black; and all
the most distinct races, when blue, are occasionally chequered in precisely
the same manner. Thus I have seen Pouters, Fantails, Carriers, Turbits,
Tumblers (Indian and English), Swallows, Bald-pates, and other toy-pigeons
blue and chequered; and Mr. Esquilant has seen a chequered Runt. I bred
from two pure blue Tumblers a chequered bird.
The facts hitherto given refer to the occasional appearance in pure races
of blue birds with black wing-bars, and likewise of blue and chequered
birds; but it will now be seen that when two birds belonging to distinct
races are crossed, neither of which have, nor probably have had during many
generations, a trace of blue in their plumage, or a trace of wing-bars and
the other characteristic marks, they very frequently produce mongrel
offspring of a blue colour, sometimes chequered, with black wing-bars,
etc.; or if not of a blue colour, yet with the several characteristic marks
more or less plainly developed. I was led to investigate this subject from
MM. Boitard and Corbie (6/26. 'Les Pigeons' etc. page 37.) having asserted
that from crosses between certain breeds it is rare to get anything but
bisets or dovecote pigeons, which, as we know, are blue birds with the
usual characteristic marks. We shall hereafter see that this subject
possesses, independently of our present object, considerable interest, so
that I will give the results of my own trials in full. I selected for
experiment races which, when pure, very seldom produce birds of a blue
colour, or have bars on their wings and tail.
The Nun is white, with the head, tail, and primary wing-feathers black; it
is a breed which was established as long ago as the year 1600. I crossed a
male Nun with a female red common Tumbler, which latter variety generally
breeds true. Thus neither parent had a trace of blue in the plumage, or of
bars on the wing and tail. I should premise that common Tumblers are rarely
blue in England. From the above cross I reared several young: one was red
over the whole back, but with the tail as blue as that of the rock-pigeon;
the terminal bar, however, was absent, but the outer feathers were edged
with white: a second and third nearly resembled the first, but the tail in
both presented a trace of the bar at the end: a fourth was brownish, and
the wings showed a trace of the double bar: a fifth was pale blue over the
whole breast, back, croup, and tail, but the neck and primary wing-feathers
were reddish; the wings presented two distinct bars of a red colour; the
tail was not barred, but the outer feathers were edged with white. I
crossed this last curiously coloured bird with a black mongrel of
complicated descent, namely, from a black Barb, a Spot, and Almond-tumbler,
so that the two young birds produced from this cross included the blood of
five varieties, none of which had a trace of blue or of wing and tail-bars:
one of the two young birds was brownish-black, with black wing-bars; the
other was reddish-dun, with reddish wing-bars, paler than the rest of the
body, with the croup pale blue, the tail bluish with a trace of the
terminal bar.
Mr. Eaton (6/27. 'Treatise on Pigeons' 1858 page 145.) matched two Short-
faced Tumblers, namely, a splash cock and kite hen (neither of which are
blue or barred), and from the first nest he got a perfect blue bird, and
from the second a silver or pale blue bird, both of which, in accordance
with all analogy, no doubt presented the usual characteristic marks.
I crossed two male black Barbs with two female red Spots. These latter have
the whole body and wings white, with a spot on the forehead, the tail and
tail-coverts red; the race existed at least as long ago as 1676, and now
breeds perfectly true, as was known to be the case in the year 1735. (6/28.
J. Moore 'Columbarium' 1735; in J.M. Eaton's edition 1852 page 71.) Barbs
are uniformly-coloured birds, with rarely even a trace of bars on the wing
or tail; they are known to breed very true. The mongrels thus raised were
black or nearly black, or dark or pale brown, sometimes slightly piebald
with white: of these birds no less than six presented double wing-bars; in
two the bars were conspicuous and quite black; in seven some white feathers
appeared on the croup; and in two or three there was a trace of the
terminal bar to the tail, but in none were the outer tail-feathers edged
with white.
I crossed black Barbs (of two excellent strains) with purely-bred, snow-
white Fantails. The mongrels were generally quite black, with a few of the
primary wing and tail feathers white: others were dark reddish-brown, and
others snow-white: none had a trace of wing-bars or of the white croup. I
then paired together two of these mongrels, namely, a brown and black bird,
and their offspring displayed wing-bars, faint, but of a darker brown than
the rest of body. In a second brood from the same parents a brown bird was
produced, with several white feathers confined to the croup.
I crossed a male dun Dragon belonging to a family which had been dun-
coloured without wing-bars during several generations, with a uniform red
Barb (bred from two black Barbs); and the offspring presented decided but
faint traces of wing-bars. I crossed a uniform red male Runt with a White
trumpeter; and the offspring had a slaty-blue tail with a bar at the end,
and with the outer feathers edged with white. I also crossed a female black
and white chequered Trumpeter (of a different strain from the last) with a
male Almond-tumbler, neither of which exhibited a trace of blue, or of the
white croup, or of the bar at end of tail: nor is it probable that the
progenitors of these two birds had for many generations exhibited any of
these characters, for I have never even heard of a blue Trumpeter in this
country, and my Almond-tumbler was purely bred; yet the tail of this
mongrel was bluish, with a broad black bar at the end, and the croup was
perfectly white. It may be observed in several of these cases, that the
tail first shows a tendency to become by reversion blue; and this fact of
the persistency of colour in the tail and tail-coverts (6/29. 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 mongrel
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 aill-coverts, which were pale fawn, and except the faintest trace of
double wing-bars of the same pale fawn tint.) will surprise no one who has
attended to the crossing of pigeons.
The last case which I will give is the most curious. I paired a mongrel
female Barb-fantail with a mongrel male Barb-spot; neither of which
mongrels had the least blue about them. Let it be remembered that blue
Barbs are excessively rare; that Spots, as has been already stated, were
perfectly characterised in the year 1676, and breed perfectly true; this
likewise is the case with white Fantails, so much so that I have never
heard of white Fantails throwing any other colour. Nevertheless the
offspring from the above two mongrels was of exactly the same blue tint as
that of the wild rock-pigeon from the Shetland Islands over the whole back
and wings; the double black wing-bars were equally conspicuous; the tail
was exactly alike in all its characters, and the croup was pure white; the
head, however, was tinted with a shade of red, evidently derived from the
Spot, and was of a paler blue than in the rock-pigeon, as was the stomach.
So that two black Barbs, a red Spot, and a white Fantail, as the four
purely-bred grandparents, produced a bird exhibiting the general blue
colour, together with every characteristic mark, the wild Columba livia.
With respect to crossed breeds frequently producing blue birds chequered
with black, and resembling in all respects both the dovecote-pigeon and the
chequered wild variety of the rock-pigeon, the statement before referred to
by MM. Boitard and Corbie would almost suffice; but I will give three
instances of the appearance of such birds from crosses in which one alone
of the parents or great-grandparents was blue, but not chequered. I crossed
a male blue Turbit with a snow-white Trumpeter, and the following year with
a dark, leaden-brown, Short-faced Tumbler; the offspring from the first
cross were as perfectly chequered as any dovecote-pigeon; and from the
second, so much so as to be nearly as black as the most darkly chequered
rock-pigeon from Madeira. Another bird, whose great-grandparents were a
white Trumpeter, a white Fantail, a white Red-spot, a red Runt, and a blue
Pouter, was slaty-blue and chequered exactly like a dovecote-pigeon. I may
here add a remark made to me by Mr. Wicking, who has had more experience
than any other person in England in breeding pigeons of various colours:
namely, that when a blue, or a blue and chequered bird, having black wing-
bars, once appears in any race and is allowed to breed, these characters
are so strongly transmitted that it is extremely difficult to eradicate
them.
What, then, are we to conclude from this tendency in all the chief domestic
races, both when purely bred and more especially when intercrossed, to
produce offspring of a blue colour, with the same characteristic marks,
varying in the same manner, as in Columbia livia? If we admit that these
races are 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, etc., without
obtaining a single blue or barred bird; yet by crossing these breeds I
reared in the first and second generation, during the course of only three
or four years, a considerable number of young birds, more or less plainly
coloured blue, and with most of the characteristic marks. When black and
white, or black and red birds, are crossed, it would appear that a slight
tendency exists in both parents to produce blue offspring, and that this,
when combined, overpowers the separate tendency in either parent to produce
black, or white, or red offspring.
If we reject the belief that all the races of the pigeon are the modified
descendants of C. livia, and suppose that they are descended from several
aboriginal stocks, then we must choose between the three following
assumptions: firstly, that at least eight or nine species formerly existed
which were aboriginally coloured in various ways, but have since varied in
exactly the same manner so as to assume the colouring of C. livia; but this
assumption throws not the least light on the appearance of such colours and
marks when the races are crossed. Or secondly, we may assume that the
aboriginal species were all coloured blue, and had the wing-bars and other
characteristic marks of C. livia,--a supposition which is highly
improbable, as besides this one species no existing member of the
Columbidae presents these combined characters; and it would not be possible
to find any other instance of several species identical in plumage, yet as
different in important points of structure as are Pouters, Fantails,
Carriers, Tumblers, etc. Or lastly, we may assume that all the races,
whether descended from C. livia or from several aboriginal species,
although they have been bred with so much care and are so highly valued by
fanciers, have all been crossed within a dozen or score of generations with
C. livia, and have thus acquired their tendency to produce blue birds with
the several characteristic marks. I have said that it must be assumed that
each race has been crossed with C. livia within a dozen, or, at the utmost,
within a score of generations; for there is no reason to believe that
crossed offspring ever revert to one of their ancestors when removed by a
greater number of generations. In a breed which has been crossed only once,
the tendency to reversion will naturally become less and less in the
succeeding generations, as in each there will be less and less of the blood
of the foreign breed; but when there has been no cross with a distinct
breed, and there is a tendency in both parents to revert to some long-lost
character, this tendency, for all that we can see to the contrary, may be
transmitted undiminished for an indefinite number of generations. These two
distinct cases of reversion are often confounded together by those who have
written on inheritance.
Considering, on the one hand, the improbability of the three assumptions
which have just been discussed, and, on the other hand, how simply the
facts are explained on the principle of reversion, we may conclude that the
occasional appearance in all the races, both when purely bred and more
especially when crossed, of blue birds, sometimes chequered, with double
wing-bars, with white or blue croups, with a bar at the end of the tail,
and with the outer tail-feathers edged with white, affords an argument of
the greatest weight in favour of the view that all are descended from
Columba livia, including under this name the three or four wild varieties
or sub-species before enumerated.
To sum up the six foregoing arguments, which are opposed to the belief that
the chief domestic races are the descendants of at least eight or nine or
perhaps a dozen species; for the crossing of any less number would not
yield the characteristic differences between the several races. FIRSTLY,
the improbability that so many species should still exist somewhere, but be
unknown to ornithologists, or that they should have become within the
historical period extinct, although man has had so little influence in
exterminating the wild C. livia. SECONDLY, the improbability of man in
former times having thoroughly domesticated and rendered fertile under
confinement so many species. THIRDLY, these supposed species having nowhere
become feral. FOURTHLY, the extraordinary fact that man should,
intentionally or by chance, have chosen for domestication several species,
extremely abnormal in character; and furthermore, the points of structure
which render these supposed species so abnormal being now highly variable.
FIFTHLY, the fact of all the races, though differing in many important
points of structure, producing perfectly fertile mongrels; whilst all the
hybrids which have been produced between even closely allied species in the
pigeon-family are sterile. SIXTHLY, the remarkable statements just given on
the tendency in all the races, both when purely bred and when crossed, to
revert in numerous minute details of colouring to the character of the wild
rock-pigeon, and to vary in a similar manner. To these arguments may be
added the extreme improbability that a number of species formerly existed,
which differed greatly from each other in some few points, but which
resembled each other as closely as do the domestic races in other points of
structure, in voice, and in all their habits of life. When these several
facts and arguments are fairly taken into consideration, it would require
an overwhelming amount of evidence to make us admit that the chief domestic
races are descended from several aboriginal stocks; and of such evidence
there is absolutely none.
The belief that the chief domestic races are descended from several wild
stocks no doubt has arisen from the apparent improbability of such great
modifications of structure having been effected since man first
domesticated the rock-pigeon. Nor am I surprised at any degree of
hesitation in admitting their common parentage: formerly, when I went into
my aviaries and watched such birds as Pouters, Carriers, Barbs, Fantails,
and Short-faced Tumblers, etc., I could not persuade myself that all had
descended from the same wild stock, and that man had consequently in one
sense created these remarkable modifications. Therefore I have argued the
question of their origin at great, and, as some will think, superfluous
length.
Finally, in favour of the belief that all the races are descended from a
single stock, we have in Columba livia a still existing and widely
distributed species, which can be and has been domesticated in various
countries. This species agrees in most points of structure and in all its
habits of life, as well as occasionally in every detail of plumage, with
the several domestic races. It breeds freely with them, and produces
fertile offspring. It varies in a state of nature (6/30. It deserves
notice, as bearing on the general subject of variation, that not only C.
livia presents several wild forms, regarded 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 predicament. This is the case, as Mr.
Blyth has remarked to me, with Treron, Palumbus, and Turtur.), and still
more so when semi-domesticated, as shown by comparing the Sierra Leone
pigeons with those of India, or with those which apparently have run wild
in Madeira. It has undergone a still greater amount of variation in the
case of the numerous toy-pigeons, which no one supposes to be descended
from distinct species; yet some of these toy-pigeons have transmitted their
character truly for centuries. Why, then, should we hesitate to believe in
that greater amount of variation which is necessary for the production of
the eleven chief races? It should be borne in mind that in two of the most
strongly-marked races, namely, Carriers and Short-faced Tumblers, the
extreme forms can be connected with the parent-species by graduated
differences not greater than those which may be observed between the
dovecote-pigeons inhabiting different countries, or between the various
kinds of toy-pigeons,--gradations which must certainly be attributed to
variation.
That circumstances have been eminently favourable for the modification of
the pigeon through variation and selection will now be shown. The earliest
record, as has been pointed out to me by Professor Lepsius, of pigeons in a
domesticated condition, occurs in the fifth Egyptian dynasty, about 3000
B.C. (6/31. 'Denkmaler' abth. 2 bl. 70.); 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. (6/32. 'The 'Dovecote' by the Rev. E.S. Dixon 1851 pages 11-13.
Adolphe Pictet (in his 'Les Origines Indo-Europeennes' 1859 page 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 of the pigeon in the East.) In the time of the Romans, as we
hear from Pliny (6/33. English translation 1601 book 10 ch. 37.), 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
monarch of Iran and Turan sent him some very rare breeds. His Majesty,"
says the courtly historian, "by crossing the breeds, which method was never
practised before, has improved them astonishingly." (6/34. 'Ayeen Akbery'
translated by F. Gladwin 4to edition volume 1 page 270.) Akber Khan
possessed seventeen distinct kinds, eight of which were valuable for beauty
alone. At about this same period of 1600 the Dutch, according to
Aldrovandi, were as eager about pigeons as the Romans had formerly been.
The breeds which were kept during the fifteenth century in Europe and in
India apparently differed from each other. Tavernier, in his Travels in
1677, speaks, as does Chardin in 1735, of the vast number of pigeon-houses
in Persia; and the former remarks that, as Christians were not permitted to
keep pigeons, some of the vulgar actually turned Mahometans for this sole
purpose. The Emperor of Morocco had his favourite keeper of pigeons, as is
mentioned in Moore's treatise, published 1737. In England, from the time of
Willughby in 1678 to the present day, as well as in Germany and in France,
numerous treatises have been published on the pigeon. In India, about a
hundred years ago, a Persian treatise was written; and the writer thought
it no light affair, for he begins with a solemn invocation, "in the name of
God, the gracious and merciful." Many large towns, in Europe and the United
States, now have their societies of devoted pigeon-fanciers: at present
there are three such societies in London. In India, as I hear from Mr.
Blyth, the inhabitants of Delhi and of some other great cities are eager
fanciers. Mr. Layard informs me that most of the known breeds are kept in
Ceylon. In China, according to Mr. Swinhoe of Amoy, and Dr. Lockhart of
Shangai, Carriers, Fantails, Tumblers, and other varieties are reared with
care, especially by the bonzes or priests. The Chinese fasten a kind of
whistle to the tail-feathers of their pigeons, and as the flock wheels
through the air they produce a sweet sound. In Egypt the late Abbas Pacha
was a great fancier of Fantails. Many pigeons are kept at Cairo and
Constantinople, and these have lately been imported by native merchants, as
I hear from Sir W. Elliot, into Southern India, and sold at high prices.
The foregoing statements show in how many countries, and during how long a
period, many men have been passionately devoted to the breeding of pigeons.
Hear how an enthusiastic fancier at the present day writes: "If it were
possible for noblemen and gentlemen to know the amazing amount of solace
and pleasure derived from Almond Tumblers, when they begin to understand
their properties, I should think that scarce any nobleman or gentleman
would be without their aviaries of Almond Tumblers." (6/35. J.M. Eaton
'Treatise on the Almond Tumbler' 1851; Preface page 6.) The pleasure thus
taken is of paramount importance, as it leads amateurs carefully to note
and preserve each slight deviation of structure which strikes their fancy.
Pigeons are often closely confined during their whole lives; they do not
partake of their naturally varied diet; they have often been transported
from one climate to another; and all these changes in their conditions of
life would be likely to cause variability. Pigeons have been domesticated
for nearly 5000 years, and have been kept in many places, so that the
numbers reared under domestication must have been enormous: and this is
another circumstance of high importance, for it obviously favours the
chance of rare modifications of structure occasionally appearing. Slight
variations of all kinds would almost certainly be observed, and, if valued,
would, owing to the following circumstances, be preserved and propagated
with unusual facility. Pigeons, differently from any other domesticated
animal, can easily be mated for life, and, though kept with other pigeons,
rarely prove unfaithful to each other. Even when the male does break his
marriage-vow, he does not permanently desert his mate. I have bred in the
same aviaries many pigeons of different kinds, and never reared a single
bird of an impure strain. Hence a fancier can with the greatest ease select
and match his birds. He will also 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.
[HISTORY OF THE PRINCIPAL RACES OF THE PIGEON. (6/36. As in the following
discussion I often speak of the present time, I should state that this
chapter was completed in the year 1858.)
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 is, than of any
other domesticated animal. Some of the cases are interesting as proving how
long domestic varieties may be propagated with exactly the same or nearly
the same characters; and other cases are still more interesting as showing
how slowly but steadily races have been greatly modified during successive
generations. In the last chapter I stated that Trumpeters and Laughers,
both so remarkable for their voices, seem to have been perfectly
characterised in 1735; and Laughers were apparently known in India before
the year 1600. Spots in 1676, and Nuns in the time of Aldrovandi, before
1600, were coloured exactly as they now are. Common Tumblers and Ground
Tumblers displayed 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 characterised 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 characterised in Aldrovandi's time (6/37.
'Ornithologie' 1600 volume 2 page 360.), 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 1/2 or 6 3/4 long
"must be allowed to be a very good one." Mr. Bult, the most successful
breeder of Pouters in the world, informs me that at present (1858) the
standard length of the body is not less than 18 inches; but he has measured
one bird 19 inches in length, and has heard of 20 and 22 inches, but doubts
the truth of these latter statements. The standard length of the leg is now
7 inches, but Mr. Bult has recently measured two of his own birds with legs
7 1/2 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 7 1/2 inches in length. The extremely slight
improvement in Pouters, except in the length of the leg, during the last
123 years, may be partly accounted for by the neglect which they suffered,
as I am informed by Mr. Bult, until within the last 20 or 30 years. About
1765 (6/38. 'A Treatise on Domestic Pigeons' dedicated to Mr. Mayor 1765
Preface page 14.) there was a change of fashion, stouter and more feathered
legs being preferred to thin and nearly naked legs.
FANTAILS.
The first notice of the existence of this breed is in India, before the
year 1600, as given in the 'Ayeen Akbery' (6/39. Mr. Blyth has given a
translation of part of the 'Ayeen Akbery' in 'Annals and Mag. of Nat.
Hist.' volume 19 1847 page 104.); at this date, judging from Aldrovandi,
the breed was unknown in Europe. In 1677 Willughby speaks of a Fantail with
26 tail-feathers; in 1735 Moore saw one with 36 feathers; and in 1824 MM.
Boitard and Corbie 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 valued. The old
descriptions do not suffice to show whether in these latter respects there
has been much improvement: but if Fantails with their heads and tails
touching had formerly existed, as at the present time, the fact would
almost certainly have been noticed. The Fantails which are now found in
India probably show the state of the race, as far as carriage is concerned,
at the date of their introduction into Europe; and some, said to have been
brought from Calcutta, which I kept alive, were in a marked manner inferior
to our exhibition birds. The Java Fantail shows the same difference in
carriage; and although Mr. Swinhoe has counted 18 and 24 tail-feathers in
his birds, a first-rate specimen sent to me had only 14 tail-feathers.
JACOBINS.
This breed existed before 1600, but the hood, judging from the figure given
by Aldrovandi, did not enclose the head nearly so perfectly as at present:
nor was the head then white; nor were the wings and tail so long, but this
last character might have been overlooked by the rude artist. In Moore's
time, in 1735, the Jacobin was considered the smallest kind of pigeon, and
the bill is said to be very short. Hence either the Jacobin, or the other
kinds with which it was then compared, must since that time have been
considerably modified; for Moore's description (and it must be remembered
that he was a first-rate judge) is clearly not applicable, as far as size
of body and length of beak are concerned, to our present Jacobins. In 1795,
judging from Bechstein, the breed had assumed its present character.
TURBITS.
It has generally been supposed by the older writers on pigeons, that the
Turbit is the Cortbeck of Aldrovandi; but if this be the case, it is an
extraordinary fact that the characteristic frill should not have been
noticed. The beak, moreover, of the Cortbeck is described as closely
resembling that of the Jacobin, which shows a change in the one or the
other race. The Turbit, with its characteristic frill, and bearing its
present name, is described by Willughby in 1677; and the bill is said to be
like that of the bullfinch,--a good comparison, but now more strictly
applicable to the beak of the Barb. The sub-breed called the Owl was well
known in Moore's time, in 1735.
TUMBLERS.
Common Tumblers, as well as Ground Tumblers, perfect as far as tumbling is
concerned, existed in India before the year 1600; and at this period
diversified modes of flight, such as flying at night, the ascent to a great
height, and manner of descent, seem to have been much attended to in India,
as at the present time. Belon (6/40. 'L'Histoire de la Nature des Oiseaux'
page 314.) in 1555 saw in Paphlagonia what he describes as "a very new
thing, viz. pigeons which flew so high in the air that they were lost to
view, but returned to their pigeon-house without separating." This manner
of flight is characteristic of our present Tumblers, but it is clear that
Belon would have mentioned the act of tumbling if the pigeons described by
him had tumbled. Tumblers were not known in Europe in 1600, as they are not
mentioned by Aldrovandi, who discusses the flight of pigeons. They are
briefly alluded to by Willughby, in 1687, as small pigeons "which show like
footballs in the air." The short-faced race did not exist at this period,
as Willughby could not have overlooked birds so remarkable for their small
size and short beaks. We can even trace some of the steps by which this
race has been produced. Moore in 1735 enumerates correctly the chief points
of excellence, but does not give any description of the several sub-breeds;
and from this fact Mr. Eaton infers (6/41. 'Treatise on Pigeons' 1852 page
64.) that the Short-faced Tumbler had not then come to full perfection.
Moore even speaks of the Jacobin as being the smallest pigeon. Thirty years
afterwards, in 1765, in the Treatise dedicated to Mayor, short-faced Almond
Tumblers are fully described, but the author, an excellent fancier,
expressly states in his Preface (page 14) that, "from great care and
expense in breeding them, they have arrived to so great perfection and are
so different from what they were 20 or 30 years past, that an old fancier
would have condemned them for no other reason than because they are not
like what used to be thought good when he was in the fancy before." Hence
it would appear that there was a rather sudden change in the character of
the short-faced Tumbler at about this period; and there is reason to
suspect that a dwarfed and half-monstrous bird, the parent-form of the
several short-faced sub-breeds, then appeared. I suspect this because
short-faced Tumblers are born with their beaks (ascertained by careful
measurement) as short, proportionally with the size of their bodies, as in
the adult bird; and in this respect they differ greatly from all other
breeds, which slowly acquire during growth their various characteristic
qualities.
Since the year 1765 there has been some change in one of the chief
characters of the short-faced Tumbler, namely, in the length of the beak.
Fanciers measure the "head and beak" from the tip of the beak to the front
corner of the eyeball. About the year 1765 a "head and beak" was considered
good (6/42. J.M. Eaton 'Treatise on the Breeding and Managing of the Almond
Tumbler' 1851. Compare page 5 of Preface, page 9 and page 32), which,
measured in the usual manner, was 7/8 of an inch in length; now it ought
not to exceed 5/8 of an inch; "it is however possible," as Mr. Eaton
candidly confesses, "for a bird to be considered as pleasant or neat even
at 6/8 of an inch, but exceeding that length it must be looked upon as
unworthy of attention." Mr. Eaton states that he has never seen in the
course of his life more than two or three birds with the "head and beak"
not exceeding half an inch in length; "still I believe in the course of a
few years that the head and beak will be shortened, and that half-inch
birds will not be considered so great a curiosity as at the present time."
That Mr. Eaton's opinion deserves attention cannot be doubted, considering
his success in winning prizes at our exhibitions. Finally in regard to the
Tumbler it may be concluded from the facts above given that it was
originally introduced into Europe, probably first into England, from the
East; and that it then resembled our common English Tumbler, or more
probably the Persian or Indian Tumbler, with a beak only just perceptibly
shorter than that of the common dovecote-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 description and figures; four breeds,
however, existed in the year 1600 which evidently were allied both to Barbs
and Carriers. To show how difficult it is to recognise some of the breeds
described by Aldrovandi I will give the different opinions in regard to the
above four kinds, named by him C. indica, cretensis, gutturosa, and
persica. Willughby thought that the Columba indica was a Turbit, but the
eminent fancier Mr. Brent believes that it was an inferior Barb: C.
cretensis, with a short beak and a swelling on the upper mandible, cannot
be recognised: C. (falsely called) gutturosa, which from its rostrum,
breve, crassum, et tuberosum seems to me to come nearest to the Barb, Mr.
Brent believes to be a Carrier; and lastly, the C. persica et turcica, Mr.
Brent thinks, and I quite concur with him, was a short-beaked Carrier with
very little wattle. In 1687 the Barb was known in England, and Willughby
describes the beak as like that of the Turbit; but it is not credible that
his Barbs 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, yet he
adds, "the bill is not short, but of a moderate length;" a description
which no one would apply to our present Carriers, so conspicuous for the
extraordinary length of their beaks. The old names given in Europe to the
Carrier, and the several names now in use in India, indicate that Carriers
originally came from Persia; and Willughby's description would perfectly
apply to the Bussorah Carrier as it now exists in Madras. In later times we
can partially trace the progress of change in our English Carriers: Moore,
in 1735, says "an inch and a half is reckoned a long beak, though there are
very good Carriers that are found not to exceed an inch and a quarter."
These birds must have resembled or perhaps been a little superior to the
Carriers, previously described, now found in Persia. In England at the
present day "there are," as Mr. Eaton (6/43. 'Treatise on Pigeons' 1852
page 41.) states, "beaks that would measure (from edge of eye to tip of
beak) one inch and three-quarters, and some few even two inches in
length."]
From these historical details we see that nearly all the chief domestic
races existed before the year 1600. Some remarkable only for colour appear
to have been identical with our present breeds, some were nearly the same,
some considerably different, and some have since become extinct. Several
breeds, such as Finnikins and Turners, the swallow-tailed pigeon of
Bechstein and the Carmelite, seem to have originated and to have
disappeared within this same period. Any one now visiting a well-stocked
English aviary would certainly pick out as the most distinct kinds, the
massive Runt, the Carrier with its wonderfully elongated beak and great
wattles, the Barb with its short broad beak and eye-wattles, the short-
faced Tumbler with its small conical beak, the Pouter with its great crop,
long legs and body, the Fantail with its upraised, widely-expanded, well-
feathered tail, the Turbit with its frill and short blunt beak, and the
Jacobin with his hood. Now, if this same person could have viewed the
pigeons kept before 1600 by Akber Khan in India and by Aldrovandi in
Europe, he would have seen the Jacobin with a less perfect hood; the Turbit
apparently without its frill; the Pouter with shorter legs, and in every
way less remarkable--that is, if Aldrovandi's Pouter resembled the old
German kind; the Fantail would have been far less singular in appearance,
and would have had much fewer feathers in its tail; he would have seen
excellent flying Tumblers, but he would in vain have looked for the
marvellous short-faced breeds; he would have seen birds allied to Barbs,
but it is extremely doubtful whether he would have met with our actual
Barbs; and lastly, he would have found Carriers with beaks and wattle
incomparably less developed than in our English Carriers. He might have
classed most of the breeds in the same groups as at present; but the
differences between the groups were then far less strongly pronounced than
at present. In short, the several breeds had at this early period not
diverged in so great a degree as now 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
dovecotes in their native country, without any care in selecting and
matching them, they are liable to little more variation than the wild C.
livia, namely, in the wings becoming chequered with black, in the croup
being blue or white, and in the size of the body. When, however, dovecote-
pigeons are transported into diversified countries, such as Sierra Leone,
the Malay archipelago, and Madeira, they are exposed to new conditions of
life; and apparently in consequence 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 in their native climate, to
considerably different conditions; for they cannot obtain their natural
diversity of food; and, what is probably more important, they are
abundantly fed, whilst debarred from taking much exercise. Under these
circumstances we might expect to find, from the analogy of all other
domesticated animals, a greater amount of individual variability than with
the wild pigeon; and this is the case. The want of exercise apparently
tends to reduce the size of the feet and organs of flight; and then, from
the law of correlation of growth, the beak apparently becomes affected.
From what we now see occasionally taking place in our aviaries, we may
conclude that sudden variations or sports, such as the appearance of a
crest of feathers on the head, of feathered feet, of a new shade of colour,
of an additional feather in the tail or wing, would occur at rare intervals
during the many centuries which have elapsed since the pigeon was first
domesticated. At the present day such "sports" are generally rejected as
blemishes; and there is so much mystery in the breeding of pigeons that, if
a valuable sport did occur, its history would often be concealed. Before
the last hundred and fifty years, there is hardly a chance of the history
of any such sport having been recorded. But it by no means follows from
this that such sports in former times, when the pigeon had undergone much
less variation, would have been rejected. We are profoundly ignorant of the
cause of each sudden and apparently spontaneous variation, as well as of
the infinitely numerous shades of difference between the birds of the same
family. But in a future chapter we shall see that all such variations
appear to be the indirect result of changes of some kind in the conditions
of life.
Hence, after a long course of domestication, we might expect to see in the
pigeon much individual variability, and occasional sudden variations, as
well as slight modifications from the lessened use of certain parts,
together with the effects of correlation of growth. But without selection
all this would produce only a trifling or no result; for without such aid
differences of all kinds would, from the two following causes, soon
disappear. In a healthy and vigorous lot of pigeons many more young birds
are killed for food or die than are reared to maturity; so that an
individual having any peculiar character, if not selected, would run a good
chance of being destroyed; and if not destroyed, the peculiarity in
question would generally be obliterated by free intercrossing. It might,
however, occasionally happen that the same variation repeatedly occurred,
owing to the action of peculiar and uniform conditions of life, and in this
case it would prevail independently of selection. But when selection is
brought into play all is changed; for this is the foundation-stone in the
formation of new races; and with the pigeon, circumstances, as we have
already seen, are eminently favourable for selection. When a bird
presenting some conspicuous variation has been preserved, and its offspring
have been selected, carefully matched, and again propagated, and so onwards
during successive generations, the principle is so obvious that nothing
more need be said about it. This may be called METHODICAL SELECTION, for
the breeder has a distinct object in view, namely, to preserve some
character which has actually appeared; or to create some improvement
already pictured in his mind.
Another form of selection has hardly been noticed by those authors who have
discussed this subject, but is even more important. This form may be called
UNCONSCIOUS SELECTION, for the breeder selects his birds unconsciously,
unintentionally, and without method, yet he surely though slowly produces a
great result. I refer to the effects which follow from each fancier at
first procuring and afterwards rearing as good birds as he can, according
to his skill, and according to the standard of excellence at each
successive period. He does not wish permanently to modify the breed; he
does not look to the distant future, or speculate on the final result of
the slow accumulation during many generations of successive slight changes;
he is content if he possesses a good stock, and more than content if he can
beat his rivals. The fancier in the time of Aldrovandi, when in the year
1600 he admired his own Jacobins, Pouters, or Carriers, never reflected
what their descendants in the year 1860 would become: he would have been
astonished could he have seen our Jacobins, our improved English Carriers,
and our Pouters; he would probably have denied that they were the
descendants of his own once-admired stock, and he would perhaps not have
valued them, for no other reason, as was written in 1765, "than because
they were not like what used to be thought good when he was in the fancy."
No one will attribute the lengthened beak of the Carrier, the shortened
beak of the Short-faced Tumbler, the lengthened leg of the Pouter, the more
perfectly enclosed hood of the Jacobin, etc.--changes effected since the
time of Aldrovandi, or even since a much later period,--to the direct and
immediate action of the conditions of life. For these several races have
been modified in various and even in directly opposite ways, though kept
under the same climate and treated in all respects in as nearly uniform a
manner as possible. Each slight change in the length or shortness of the
beak, in the length of leg, etc., has no doubt been indirectly and remotely
caused by some change in the conditions to which the bird has been
subjected, but we must attribute the final result, as is manifest in those
cases of which we have any historical record, to the continued selection
and accumulation of many slight successive variations.
The action of unconscious selection, as far as pigeons are concerned,
depends on a universal principle in human nature, namely, on our rivalry,
and desire to outdo our neighbours. We see this in every fleeting fashion,
even in our dress, and it leads the fancier to endeavour to exaggerate
every peculiarity in his breeds. A great authority on pigeons (6/44. Eaton
'Treatise on Pigeons' 1858 page 86.), says, "Fanciers do not and will not
admire a medium standard, that is, half and half, which is neither here nor
there, but admire extremes." After remarking that the fancier of Short-
faced Beard Tumblers wishes for a very short beak, and that the fancier of
Long-faced Beard Tumblers wishes for a very long beak, he says, with
respect to one of intermediate length, "Don't deceive yourself. Do you
suppose for a moment the short or the long-faced fancier would accept such
a bird as a gift? Certainly not; the short-faced fancier could see no
beauty in it; the long-faced fancier would swear there was no use in it,
etc." In these comical passages, written seriously, we see the principle
which has ever guided fanciers, and has led to such great modifications in
all the domestic races which are valued solely for their beauty or
curiosity.
Fashions in pigeon-breeding endure for long periods; we cannot change the
structure of a bird as quickly as we can the fashion of our dress. In the
time of Aldrovandi, no doubt the more the pouter inflated his crop, the
more he was valued. Nevertheless, fashions do to a certain extent change;
first one point of structure and then another is attended to; or different
breeds are admired at different times and in different countries. As the
author just quoted remarks, "the fancy ebbs and flows; a thorough fancier
now-a-days never stoops to breed toy-birds;" yet these very "toys" are now
most carefully bred in Germany. Breeds which at the present time are highly
valued in India are considered worthless in England. No doubt, when breeds
are neglected, they degenerate; still we may believe that, as long as they
are kept under the same conditions of life, characters once gained will be
partially retained for a long time, and may form the starting-point for a
future course of selection.
Let it not be objected to this view of the action of unconscious selection
that fanciers would not observe or care for extremely slight differences.
Those alone who have associated with fanciers can be thoroughly aware of
their accurate powers of discrimination acquired by long practice, and of
the care and labour which they bestow on their birds. I have known a
fancier deliberately study his birds day after day to settle which to match
together and which to reject. Observe how difficult the subject appears to
one of the most eminent and experienced fanciers. Mr. Eaton, the winner of
many prizes, says, "I would here particularly guard you against keeping too
great a variety of pigeons, otherwise you will know a little about all the
kinds, but nothing about one as it ought to be known." "It is possible
there may be a few fanciers that have a good general knowledge of the
several fancy pigeons, but there are many who labour under the delusion of
supposing they know what they do not." Speaking exclusively of one sub-
variety of one race, namely, the short-faced almond tumbler, and after
saying that some fanciers sacrifice every property to obtain a good head
and beak, and that other fanciers sacrifice everything for plumage, he
remarks: "Some young fanciers who are over covetous go in for all the five
properties at once, and they have their reward by getting nothing." In
India, as I hear from Mr. Blyth, pigeons are likewise selected and matched
with the greatest care. We must not judge of the slight divergences from
existing varieties which would have been valued in ancient days, by those
which are now valued after the formation of so many races, each with its
own standard of perfection, kept uniform by our numerous Exhibitions. The
ambition of the most energetic fancier may be fully satisfied by the
difficulty of excelling other fanciers in the breeds already established,
without trying to form a new one.
A difficulty with respect to the power of selection will perhaps already
have occurred to the reader, namely, what could have led fanciers first to
attempt to make such singular breeds as Pouters, Fantails, Carriers, etc.?
But it is this very difficulty which the principle of unconscious selection
removes. Undoubtedly no fancier ever did intentionally make such an
attempt. All that we need suppose is that a variation occurred sufficiently
marked to catch the discriminating eye of some ancient fancier, and then
unconscious selection carried on for many generations, that is, the wish of
succeeding fanciers to excel their rivals, would do the rest. In the case
of the Fantail we may suppose that the first progenitor of the breed had a
tail only slightly erected, as may now be seen in certain Runts (6/45. See
Neumeister's figure of the Florence Runt, tab. 13 in 'Das Ganze der
Taubenzucht.') with some increase in the number of the tail-feathers, as
now occasionally occurs with Nuns. In the case of the Pouter we may suppose
that some bird inflated its crop a little more than other pigeons, as is
now the case in a slight degree with the oesophagus of the Turbit. We do
not 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 (6/46. Mr. W.J. Moore gives a full account of the Ground
Tumblers of India ('Indian Medical Gazette' January and February 1873), and
says the pricking the base of the brain, and giving hydrocyanic acid,
together with strychnine, to an ordinary pigeon, brings on convulsive
movements exactly like those of a Tumbler. One pigeon, the brain of which
had been pricked, completely recovered, and ever afterwards occasionally
made somersaults.) and before the year 1600 pigeons remarkable for their
diversified manner of flight were much valued in India, and by the order of
the Emperor Akber Khan were sedulously trained and carefully matched.
In the foregoing cases we have supposed that a sudden variation,
conspicuous enough to catch a fancier's eye, first appeared; but even this
degree of abruptness in the process of variation is not necessary for the
formation of a new breed. When the same kind of pigeon has been kept pure,
and has been bred during a long period by two or more fanciers, slight
differences in the strain can often be recognised. Thus I have seen first-
rate Jacobins in one man's possession which certainly differed slightly in
several characters from those kept by another. I possessed some excellent
Barbs descended from a pair which had won a prize, and another lot
descended from a stock formerly kept by that famous fancier Sir John
Sebright, and these plainly differed in the form of the beak; but the
differences were so slight that they could hardly be given by words. Again,
the common English and Dutch Tumbler differ in a somewhat greater degree,
both in length of beak and shape of head. What first caused these slight
differences cannot be explained any more than why one man has a long nose
and another a short one. In the strains long kept distinct by different
fanciers, such differences are so common that they cannot be accounted for
by the accident of the birds first chosen for breeding having been
originally as different as they now are. The explanation no doubt lies in
selection of a slightly different nature having been applied in each case;
for no two fanciers have exactly the same taste, and consequently no two,
in choosing and carefully matching their birds, prefer or select exactly
the same. As each man naturally admires his own birds, he goes on
continually exaggerating by selection whatever slight peculiarities they
may possess. This will more especially happen with fanciers living in
different countries, who do not compare their stocks or aim at a common
standard of perfection. Thus, when a mere strain has once been formed,
unconscious selection steadily tends to augment the amount of difference,
and thus converts the strain into a sub-breed and this ultimately into a
well-marked breed or race.
The principle of correlation of growth should never be lost sight of. Most
pigeons have small feet, apparently caused by their lessened use, and from
correlation, as it would appear, their beaks have likewise become reduced
in length. The beak is a conspicuous organ, and, as soon as it had thus
become perceptibly shortened, fanciers would almost certainly strive to
reduce it still more by the continued selection of birds with the shortest
beaks; whilst at the same time other fanciers, as we know has actually been
the case, would in other sub-breeds, strive to increase its length. With
the increased length of the beak, the tongue becomes greatly lengthened, as
do the eyelids with the increased development of the eye-wattles; with the
reduced or increased size of the feet, the number of the scutellae vary;
with the length of the wing, the number of the primary wing-feathers
differ; and with the increased length of the body in the pouter the number
of the sacral vertebrae is 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 vertebrae had been visible and had been attended to
by fanciers, assuredly an additional number might easily have been fixed in
the Pouter. If these latter characters had once been rendered constant, we
should never have suspected that they had at first been highly variable, or
that they had arisen from correlation, in the one case with the shortness
of the wings, and in the other case with the length of the body.
In order to understand how the chief domestic races have become distinctly
separated from each other, it is important to bear in mind, that fanciers
constantly try to breed from the best birds, and consequently that those
which are inferior in the requisite qualities are in each generation
neglected; so that after a time the less improved parent-stocks and many
subsequently formed intermediate grades become extinct. This has occurred
in the case of the Pouter, Turbit, and Trumpeter, for these highly improved
breeds are now left without any links closely connecting them either with
each other or with the aboriginal rock-pigeon. In other countries, indeed,
where the same care has not been applied, or where the same fashion has not
prevailed, the earlier forms may long remain unaltered, or altered only in
a slight degree, and we are thus sometimes enabled to recover the
connecting links. This is the case in Persia and India with the Tumbler and
Carrier, which there differ but slightly from the rock-pigeon in the
proportions of their beaks. So again in Java, the Fantail sometimes has
only fourteen caudal feathers, and the tail is much less elevated and
expanded than in our improved birds; so that the Java bird forms a link
between a first-rate Fantail and the rock-pigeon.
Occasionally a breed may be retained for some particular quality in a
nearly unaltered condition in the same country, together with highly
modified off-shoots or sub-breeds, which are valued for some distinct
property. We see this exemplified in England, where the common Tumbler,
which is valued only for its flight, does not differ much from its parent-
form, the Eastern Tumbler; whereas the Short-faced Tumbler has been
prodigiously modified, from being valued, not for its flight, but for other
qualities. But the common-flying Tumbler of Europe has already begun to
branch out into slightly different sub-breeds, such as the common English
Tumbler, the Dutch Roller, the Glasgow House-tumbler, and the Long-faced
Beard Tumbler, etc.; and in the course of centuries, unless fashions
greatly change, these sub-breeds will diverge through the slow and
insensible process of unconscious selection, and become modified, in a
greater and greater degree. After a time the perfectly graduated links
which now connect all these sub-breeds together, will be lost, for there
would be no object and much difficulty in retaining such a host of
intermediate sub-varieties.
The principle of divergence, together with the extinction of the many
previously existing intermediate forms, is so important for understanding
the origin of domestic races, as well as of species in a state of nature,
that I will enlarge a little more on this subject. Our third main group
includes Carriers, Barbs, and Runts, which are plainly related to one
another, yet wonderfully distinct in several important characters.
According to the view given in the last chapter, these three races have
probably descended from an unknown race having an intermediate character,
and this race from the rock-pigeon. Their characteristic differences are
believed to be due to different breeders having at an early period admired
different points of structure; and then, on the acknowledged principle of
admiring extremes, having gone on breeding, without any thought of the
future, as good birds as they could,--Carrier-fanciers preferring long
beaks with much wattle,--Barb-fanciers preferring short thick beaks with
much eye-wattle,--and Runt-fanciers not caring about the beak or wattle,
but only for the size and weight of the body. This process would have led
to the neglect and final extinction of the earlier, inferior, and
intermediate birds; and thus it has come to pass, that in Europe these
three races are now so extraordinarily distinct from each other. But in the
East, whence they were originally brought, the fashion has been different,
and we there see breeds which connect the highly modified English Carrier
with the rock-pigeon, and others which to a certain extent connect Carriers
and Runts. Looking back to the time of Aldrovandi, we find that there
existed in Europe, before the year 1600, four breeds which were closely
allied to Carriers and Barbs, but which competent authorities cannot now
identify with our present Barbs and Carriers; nor can Aldrovandi's Runts be
identified with our present Runts. These four breeds certainly did not
differ from each other nearly so much as do our existing English Carriers,
Barbs, and Runts. All this is exactly what might have been anticipated. If
we could collect all the pigeons which have ever lived, from before the
time of the Romans to the present day, we should be able to group them in
several lines, diverging from the parent rock-pigeon. Each line would
consist of almost insensible steps, occasionally broken by some slightly
greater variation or sport, and each would culminate in one of our present
highly modified forms. Of the many former connecting links, some would be
found to have become absolutely extinct without having left any issue,
whilst others, though extinct, would be recognised as the progenitors of
the existing races.
I have heard it remarked as a strange circumstance that we occasionally
hear of the local or complete extinction of domestic races, whilst we hear
nothing of their origin. How, it has been asked, can these losses be
compensated, and more than compensated, for we know that with almost all
domesticated animals the races have largely increased in number since the
time of the Romans? But on the view here given, we can understand this
apparent contradiction. The extinction of a race within historical times is
an event likely to be noticed; but its gradual and scarcely sensible
modification through unconscious selection, and its subsequent divergence,
either in the same or more commonly in distant countries, into two or more
strains, and their gradual conversion into sub-breeds, and these into well-
marked breeds are events which would rarely be noticed. The death of a
tree, that has attained gigantic dimensions, is recorded; the slow growth
of smaller trees and their increase in number excite no attention.
In accordance with the belief in the great power of selection, and of the
little direct power of changed conditions of life, except in causing
general variability or plasticity of organisation, it is not surprising
that dovecote-pigeons have remained unaltered from time immemorial; and
that some toy-pigeons, which differ in little else besides colour from the
dovecote-pigeon, have retained the same character for several centuries.
For when one of these toy-pigeons had once become beautifully and
symmetrically coloured,--when, for instance, a Spot had been produced with
the crown of its head, its tail, and tail-coverts of a uniform colour, the
rest of the body being snow-white,--no alteration or improvement would be
desired. On the other hand, it is not surprising that during this same
interval of time our highly-bred pigeons have undergone an astonishing
amount of change; for in regard to them there is no defined limit to the
wish of the fancier, and there is no known limit to the variability of
their characters. What is there to stop the fancier desiring to give to his
Carrier a longer and longer beak, or to his Tumbler a shorter and shorter
beak? nor has the extreme limit of variability in the beak, if there be any
such limit, as yet been reached. Notwithstanding the great improvement
effected within recent times in the Short-faced Almond Tumbler, Mr. Eaton
remarks, "the field is still as open for fresh competitors as it was one
hundred years ago;" but this is perhaps an exaggerated assertion, for the
young of all highly-improved fancy birds are extremely liable to disease
and death.
I have heard it objected that the formation of the several domestic races
of the pigeon throws no light on the origin of the wild species of the
Columbidae, 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 shape of the primary wing-feathers, in the relative
length of the hind toe, or in habits of life, as in roosting and building
in trees. But the above objection shows how completely the principle of
selection has been misunderstood. It is not likely that characters selected
by the caprice of man should resemble differences preserved under natural
conditions either from being of direct service to each species, or from
standing in correlation with other modified and serviceable structures.
Until man selects birds differing in the relative length of the wing-
feathers or toes, etc., 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 scutellae. With respect to the
domestic races not roosting or building in trees, it is obvious that
fanciers would never attend to or select such changes in habits; but we
have seen that the pigeons in Egypt, which do not for some reason like
settling on the low mud hovels of the natives, are led, apparently by
compulsion, to perch in crowds on the trees. We may even affirm that, if
our domestic races had become greatly modified in any of the above
specified respects, and it could be shown that fanciers had never attended
to such points, or that they did not stand in correlation with other
selected characters, the fact, on the principles advocated in this chapter,
would have offered a serious difficulty.
Let us briefly sum up the last two chapters on the pigeon. We may conclude
with confidence that all the domestic races, notwithstanding their great
amount of difference, are descended from the Columba livia, including under
this name certain wild races. But the differences between the latter throw
no light whatever on the characters which distinguish the domestic races.
In each breed or sub-breed the individual birds are more variable than
birds in a state of nature; and occasionally they vary in a sudden and
strongly-marked manner. This plasticity of organisation apparently results
from changed conditions of life. Disuse has reduced certain parts of the
body. Correlation of growth so ties the organisation together, that when
one part varies other parts vary at the same time. When several breeds have
once been formed, their intercrossing aids the progress of modification,
and has even produced new sub-breeds. But as, in the construction of a
building, mere stones or bricks are of little avail without the builder's
art, so, in the production of new races, selection has been the presiding
power. Fanciers can act by selection on excessively slight individual
differences, as well as on those greater differences which are called
sports. Selection is followed methodically when the fancier tries to
improve and modify a breed according to a prefixed standard of excellence;
or he acts unmethodically and unconsciously, by merely trying to rear as
good birds as he can, without any wish or intention to alter the breed. The
progress of selection almost inevitably leads to the neglect and ultimate
extinction of the earlier and less improved forms, as well as of many
intermediate links in each long line of descent. Thus it has come to pass
that most of our present races are so marvellously distinct from each
other, and from the aboriginal rock-pigeon.
CHAPTER 1.VII.
FOWLS.
BRIEF DESCRIPTIONS OF THE CHIEF BREEDS.
ARGUMENTS IN FAVOUR OF THEIR DESCENT FROM SEVERAL SPECIES.
ARGUMENTS IN FAVOUR OF ALL THE BREEDS HAVING DESCENDED FROM GALLUS BANKIVA.
REVERSION TO THE PARENT-STOCK IN COLOUR.
ANALOGOUS VARIATIONS.
ANCIENT HISTORY OF THE FOWL.
EXTERNAL DIFFERENCES BETWEEN THE SEVERAL BREEDS.
EGGS.
CHICKENS.
SECONDARY SEXUAL CHARACTERS.
WING-AND TAIL-FEATHERS, VOICE, DISPOSITION, ETC.
OSTEOLOGICAL DIFFERENCES IN THE SKULL, VERTEBRAE, ETC.
EFFECTS OF USE AND DISUSE ON CERTAIN PARTS.
CORRELATION OF GROWTH.
As some naturalists may not be familiar with the chief breeds of the fowl,
it will be advisable to give a condensed description of them. (7/1. 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 this chapter; and from his well-known knowledge, the statements
here given may be fully trusted. Mr. Tegetmeier has likewise assisted me in
every possible way in obtaining for me information and specimens. I must
not let this opportunity pass without expressing my cordial thanks to Mr.
B.P. Brent, a well-known writer on poultry, for continuous assistance and
the gift of many specimens.) From what I have read and seen of specimens
brought from several quarters of the world, I believe that most of the
chief kinds have been imported into England, but many sub-breeds are
probably still unknown here. The following discussion on the origin of the
various breeds and on their characteristic differences does not pretend to
completeness, but may be of some interest to the naturalist. The
classification of the breeds cannot, as far as I can see, be made natural.
They differ from each other in different degrees, and do not afford
characters in subordination to each other, by which they can be ranked in
group under group. They seem all to have diverged by independent and
different roads from a single type. Each chief breed includes differently
coloured sub-varieties, most of which can be truly propagated, but it would
be superfluous to describe them. I have classed the various crested fowls
as sub-breeds under the Polish fowl; but I have great doubts whether this
is a natural arrangement, showing true affinity or blood relationship. It
is scarcely possible to avoid laying stress on the commonness of a breed;
and if certain foreign sub-breeds had been largely kept in this country
they would perhaps have been raised to the rank of main-breeds. Several
breeds are abnormal in character; that is, they differ in certain points
from all wild Gallinaceous birds. At first I made a division of the breeds
into normal and abnormal, but the result was wholly unsatisfactory.
[1. GAME BREED.
This may be considered as the typical breed, as it deviates only slightly
from the wild Gallus bankiva, or, as perhaps more correctly named,
ferrugineus. Beak strong; comb single and upright. Spurs long and sharp.
Feathers closely appressed to the body. Tail with the normal number of 14
feathers. Eggs often pale buff. Disposition indomitably courageous,
exhibited even in the hens and chickens. An unusual number of differently
coloured varieties exist, such as black and brown-breasted reds, duckwings,
blacks, whites, piles, etc., 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
appressed to the body; neck-hackles short, narrow, and hard. Eggs often
pale buff. Chickens feather late. Disposition savage. Of Eastern origin.
3. COCHIN, OR SHANGAI BREED.
Size great; wing feathers short, arched, much hidden in the soft downy
plumage; barely capable of flight; tail short, generally formed of 16
feathers, developed at a late period in the young males; legs thick,
feathered; spurs short, thick; nail of middle toe flat and broad; an
additional toe not rarely developed; skin yellowish. Comb and wattle well
developed. Skull with deep medial furrow; occipital foramen, sub-
triangular, vertically elongated. Voice peculiar. Eggs rough, buff-
coloured. Disposition extremely quiet. Of Chinese origin.
4. DORKING BREED.
Size great; body square, compact; feet with an additional toe; comb well
developed, but varies much in form; wattles well developed; colour of
plumage various. Skull remarkably broad between the orbits. Of English
origin.
The white Dorking may be considered as a distinct sub-breed, being a less
massive bird.
(FIGURE 30. SPANISH FOWL.)
5. SPANISH BREED (figure 30).
Tall, with stately carriage; tarsi long; comb single, deeply serrated, of
immense size; wattles largely developed; the large ear-lobes and sides of
face white. Plumage black glossed with green. Do not incubate. Tender in
constitution, the comb being often injured by frost. Eggs white, smooth, of
large size. Chickens feather late but the young cocks show their masculine
characters, and crow at an early age. Of Mediterranean origin.
The ANDALUSIANS may be ranked as a sub-breed: they are of a slaty-blue
colour, and their chickens are well feathered. A smaller, short-legged
Dutch sub-breed has been described by some authors as distinct.
(FIGURE 31. HAMBURGH FOWL.)
6. HAMBURGH BREED (figure 31).
Size moderate; comb flat, produced backwards, covered with numerous small
points; wattle of moderate dimensions; ear lobe white; legs blueish, thin.
Do not incubate. Skull, with the tips of the ascending branches of the
premaxillary and with the nasal bones standing a little separate from each
other; anterior margin of the frontal bones less depressed than usual.
There are two sub-breeds; the SPANGLED Hamburgh, of English origin, with
the tips of the feathers marked with a dark spot; and the PENCILLED
Hamburgh, of Dutch origin, with dark transverse lines across each feather,
and with the body rather smaller. Both these sub-breeds include gold and
silver varieties, as well as some other sub-varieties. Black Hamburghs have
been produced by a cross with the Spanish breed.
(FIGURE 32. POLISH FOWL.)
7. CRESTED OR POLISH BREED (figure 32).
Head with a large, rounded crest of feathers, supported on a hemispherical
protuberance of the frontal bones, which includes the anterior part of the
brain. The ascending branches of premaxillary bones and the inner nasal
processes are much shortened. The orifice of the nostrils raised and
crescentic. Beak short. Comb absent, or small and of crescentic shape;
wattles either present or replaced by a beard-like tuft of feathers. Legs
leaden-blue. Sexual differences appear late in life. Do not incubate. There
are several beautiful varieties which differ in colour and slightly in
other respects.
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. (7.2. The best account of Sultans is by
Miss Watts in 'The Poultry Yard' 1856 page 79. I owe to Mr. Brent's
kindness the examination of some specimens of this breed.)
SUB-BREED (b) PTARMIGANS.
An inferior breed closely allied to the last, white, rather small, legs
much feathered, with the crest pointed; comb small, cupped; wattles small.
SUB-BREED (c) GHOONDOOKS.
Another Turkish breed having an extraordinary appearance; black and
tailless; crest and beard large; legs feathered. The inner processes of the
two nasal bones come into contact with each other, owing to the complete
abortion of the ascending branches of the premaxillaries. I have seen an
allied white, tailless breed from Turkey.
SUB-BREED (D) CREVE-COEUR.
A French breed of large size, barely capable of flight, with short black
legs, head crested, comb produced into two points or horns, sometimes a
little branched like the horns of a stag; both beard and wattles present.
Eggs large. Disposition quiet. (7/3. A good description, with figures, is
given of this sub-breed in the 'Journal of Horticulture' June 10, 1862 page
206.)
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, well
developed; plumage invariably mottled with black, white, and straw-yellow;
head furnished with a crest, on a triple comb placed transversely; both
wattles and beard present. (7/4. A description, with figures, is given of
this breed in 'Journal of Horticulture' June 3, 1862 page 186. Some writers
describe the comb as two-horned.)
SUB-BREED (g) GUELDERLANDS.
No comb, head said to be surmounted by a longitudinal crest of soft velvety
feathers; nostrils said to be crescentic; wattles well developed; legs
feathered; colour black. From North America. The Breda fowl seems to be
closely allied to the Guelderland.
8. BANTAM BREED.
Originally from Japan (7/5. Mr. Crawfurd 'Descript. Dict. of the Indian
Islands' page 113. Bantams are mentioned in an ancient native Japanese
Encyclopaedia, as I am informed by Mr. Birch of the British Museum.)
characterised 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. RUMPLESS FOWLS.
These are so variable in character (7/6. 'Ornamental and Domestic Poultry'
1848.) that they hardly deserve to be called a breed. Any one who will
examine the caudal vertebrae will see how monstrous the breed is.
10. CREEPERS OR JUMPERS.
These are characterised by an almost monstrous shortness of legs, so that
they move by jumping rather than by walking; they are said not to scratch
up the ground. I have examined a Burmese variety, which had a skull of
rather unusual shape.
11. FRIZZLED OR CAFFRE FOWLS.
Not uncommon in India, with the feathers curling backwards, and with the
primary feathers of the wing and tail imperfect; periosteum of bones black.
12. SILK FOWLS.
Feathers silky, with the primary wing and tail-feathers imperfect; skin and
periosteum of bones black; comb and wattles dark leaden-blue; ear-lappets
tinged with blue; legs thin, often furnished with an additional toe. Size
rather small.
13. SOOTY FOWLS.
An Indian breed, having the peculiar appearance of a white bird smeared
with soot, with black skin and periosteum. The hens alone are thus
characterised.]
From this synopsis we see that the several breeds differ considerably, and
they would have been nearly as interesting for us as pigeons, if there had
been equally good evidence that all had descended from one parent-species.
Most fanciers believe that they are descended from several primitive
stocks. The Rev. E.S. Dixon (7/7. 'Ornamental and Domestic Poultry' 1848.)
argues strongly on this side of the question; and one fancier even
denounces the opposite conclusion by asking, "Do we not perceive pervading
this spirit, the spirit of the DEIST?" Most naturalists, with the exception
of a few, such as Temminck, believe that all the breeds have proceeded from
a single species; but authority on such a point goes for little. Fanciers
look to all parts of the world as the possible sources of their unknown
stocks; thus ignoring the laws of geographical distribution. They know well
that the several kinds breed truly even in colour. They assert, but, as we
shall see, on very weak grounds, that most of the breeds are extremely
ancient. They are strongly impressed with the great difference between the
chief kinds, and they ask with force, can differences in climate, food, or
treatment have produced birds so different as the black stately Spanish,
the diminutive elegant Bantam, the heavy Cochin with its many
peculiarities, and the Polish fowl with its great top-knot and protuberant
skull? But fanciers, whilst admitting and even overrating the effects of
crossing the various breeds, do not sufficiently regard the probability of
the occasional birth, during the course of centuries, of birds with
abnormal and hereditary peculiarities; they overlook the effects of
correlation of growth--of the long-continued use and disuse of parts, and
of some direct result from changed food and climate, though on this latter
head I have found no sufficient evidence; and lastly, they all, as far as I
know, entirely overlook the all-important subject of unconscious or
unmethodical selection, though they are well aware that their birds differ
individually and that by selecting the best birds for a few generations
they can improve their stocks.
An amateur writes (7/8. Ferguson 'Illustrated Series of Rare and Prize
Poultry' 1834 page 6 Preface.) as follows: "The fact that poultry have
until lately received but little attention at the hands of the fancier, and
been entirely confined to the domains of the producer for the market, would
alone suggest the improbability of that constant and unremitting attention
having been observed in breeding, which is requisite to the consummating in
the offspring of any two birds transmittable forms not exhibited by the
parents." This at first sight appears true. But in a future chapter on
Selection, abundant facts will be given showing not only that careful
breeding, but that actual selection was practised during ancient periods,
and by barely civilised races of man. In the case of the fowl I can adduce
no direct facts showing that selection was anciently practised; but the
Romans at the commencement of the Christian era kept six or seven breeds,
and Columella "particularly recommends as the best, those sorts that have
five toes and white ears." (7/9. Rev. E.S. Dixon in his 'Ornamental
Poultry' page 203 gives an account of Columella's work.) 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. (7/10. Mr. Crawfurd 'On the Relation of the Domesticated
Animals to Civilization' separately printed page 6; first read before the
Brit. Assoc. at Oxford 1860.) Azara (7/11. 'Quadrupedes du Paraguay' tome 2
page 324.), who wrote towards the close of the last century, states that in
the interior parts of South America, where I should not have expected that
the least care would have been taken of poultry, a black-skinned and black-
boned breed is kept, from being considered fertile and its flesh good for
sick persons. Now every one who has kept poultry knows how impossible it is
to keep several breeds distinct unless the utmost care be taken in
separating the sexes. Will it then be pretended that those persons who, in
ancient times and in semi-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 ancient
times intended to form a new breed, or to modify an old breed according to
some ideal standard of excellence. He who cared for poultry would merely
wish to obtain, and afterwards to rear, the best birds which he could; but
this occasional preservation of the best birds would in the course of time
modify the breed, as surely, though by no means as rapidly, as does
methodical selection at the present day, If one person out of a hundred or
out of a thousand attended to the breeding of his birds, this would be
sufficient; for the birds thus tended would soon become superior to others,
and would form a new strain; and this strain would, as explained in the
last chapter, slowly have its characteristic differences augmented, and at
last be converted into a new sub-breed or breed. But breeds would often be
for a time neglected and would deteriorate; they would, however, partially
retain their character, and afterwards might again come into fashion and be
raised to a standard of perfection higher than their former standard; as
has actually occurred quite recently with Polish fowls. If, however, a
breed were utterly neglected, it would become extinct, as has recently
happened with one of the Polish sub-breeds. Whenever in the course of past
centuries a bird appeared with some slight abnormal structure, such as with
a lark-like crest on its head, it would probably often have been preserved
from that love of novelty which leads some persons in England to keep
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 characters, though at first
only slight in degree; from the effects of the use and the disuse of parts;
possibly from the direct effects of changed climate and food; from
correlation of growth; from occasional reversions to old and long-lost
characters; from the crossing of breeds, when more than one had been
formed; but, above all, from unconscious selection carried on during many
generations, there is no insuperable difficulty, to the best of my
judgment, in believing that all the breeds have descended from some one
parent-source. Can any single species be named from which we may reasonably
suppose that all are descended? The Gallus bankiva apparently fulfils every
requirement. I have already given as fair an account as I could of the
arguments in favour of the multiple origin of the several breeds; and now I
will give those in favour of their common descent from G. bankiva.
[But it will be convenient first briefly to describe all the known species
of Gallus. The G. sonneratii does not range into the northern parts of
India; according to Colonel Sykes (7/12. 'Proc. Zoolog. Soc.' 1832 page
151.), it presents at different heights of 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 laminae,
transversely banded with three colours; and I have met no authentic account
of any such character having been observed in any domestic breed. (7/13.
These feathers have been described by Dr. W. Marshall 'Der Zoolog. Garten'
April 1874 page 124. I examined the feathers of some hybrids raised in the
Zoological Gardens between the male G. sonneratii and a red game-hen, and
they exhibited the true character of those of G. sonneratii, except that
the horny laminae were much smaller.) This species also differs greatly
from the common fowl, in the comb being finely serrated, and in the loins
being destitute of true hackles. Its voice is utterly different. It crosses
readily in India with domestic hens; and Mr. Blyth (7/14. See also an
excellent letter on the Poultry of India by Mr. Blyth in 'Gardener's
Chronicle' 1851 page 619.) raised nearly 100 hybrid chickens; but they were
tender and mostly died whilst young. Those which were reared were
absolutely sterile when crossed inter se or with either parent. At the
Zoological Gardens, however, some 'hybrids of the same parentage were not
quite so sterile: Mr. Dixon, as he informed me, made, with Mr. Yarrell's
aid, particular inquiries on this subject, and was assured that out of 50
eggs only five or six chickens were reared. Some, however, of these half-
bred birds were crossed with one of their parents, namely, a Bantam, and
produced a few extremely feeble chickens. Mr. Dixon also procured some of
these same birds and crossed them in several ways, but all were more or
less infertile. Nearly similar experiments have recently been tried on a
great scale in the Zoological Gardens with almost the same result. (7/15.
Mr. S.J. Salter in 'Natural History Review' April 1863 page 276.) Out of
500 eggs, raised from various first crosses and hybrids, between G.
sonneratii, bankiva, and varius, only 12 chickens were reared, and of these
only three were the product of hybrids inter se. From these facts, and from
the above-mentioned strongly-marked differences in structure between the
domestic fowl and G. sonneratii, we may reject this latter species as the
parent of any domestic breed.
Ceylon possesses a fowl peculiar to the island, viz. G. stanleyii; this
species approaches so closely (except in the colouring of the comb) to the
domestic fowl, that Messrs. Layard and Kellaert (7/16. See also Mr.
Layard's paper in 'Annals and Mag. of Nat. History' 2nd series volume 14
page 62.) would have considered it, as they inform me, as one of the
parent-stocks, had it not been for its singularly different voice. This
bird, like the last, crosses readily with tame hens, and even visits
solitary farms and ravishes them. Two hybrids, a male and female, thus
produced, were found by Mr. Mitford to be quite sterile: both inherited the
peculiar voice of G. stanleyii. This species, then, may in all probability
be rejected as one of the primitive stocks of the domestic fowl.
Java and the islands eastward as far as Flores are inhabited by G. varius
(or furcatus), which differs in so many characters--green plumage,
unserrated comb, and single median wattle--that no one supposes it to have
been the parent of any one of our breeds; yet, as I am informed by Mr.
Crawfurd (7/17. See also Mr. Crawfurd 'Descriptive Dict. of the Indian
Islands' 1856 page 113.), hybrids are commonly raised between the male G.
varius and the common hen, and are kept for their great beauty, but are
invariably sterile: this, however, was not the case with some bred in the
Zoological Gardens. These hybrids were at one time thought to be
specifically distinct, and were named G. aeneus. Mr. Blyth and others
believe that the G. temminckii (7/18. Described by Mr. G.R. Gray 'Proc.
Zoolog. Soc' 1849 page 62.) (of which the history is not known) is a
similar hybrid. Sir J. Brooke sent me some skins of domestic fowls from
Borneo, and across the tail of one of these, as Mr. Tegetmeier observed,
there were transverse blue bands like those which he had seen on the tail-
feathers of hybrids from G. varius, reared in the Zoological Gardens. This
fact apparently indicates that some of the fowls of Borneo have been
slightly affected by crosses with G. varius, but the case may possibly be
one of analogous variation. I may just allude to the G. giganteus, so often
referred to in works on poultry as a wild species; but Marsden (7/19. The
passage from Marsden is given by Mr. Dixon in his 'Poultry Book' page 176.
No ornithologist now ranks this bird as a distinct species.) the first
describer, speaks of it as a tame breed; and the specimen in the British
Museum evidently has the aspect of a domestic variety.
The last species to be mentioned, namely, Gallus bankiva, has a much wider
geographical range than the three previous species; it inhabits Northern
India as far west as Sinde, and ascends the Himalaya to a height of 4000
ft.; it inhabits Burmah, the Malay peninsula, the Indo-Chinese countries,
the Philippine Islands, and the Malayan archipelago as far eastward as
Timor. This species varies considerably in the wild state. Mr. Blyth
informs me that the specimens, both male and female, brought from near the
Himalaya, are rather paler coloured than those from other parts of India;
whilst those from the Malay peninsula and Java are brighter coloured than
the Indian birds. I have seen specimens from these countries, and the
difference of tint in the hackles was conspicuous. The Malayan hens were a
shade redder on the breast and neck than the Indian hens. The Malayan males
generally had a red ear-lappet, instead of a white one as in India; but Mr.
Blyth has seen one Indian specimen without the white ear-lappet. The legs
are leaden blue in the Indian, whereas they show some tendency to be
yellowish in the Malayan and Javan specimens. In the former Mr. Blyth finds
the tarsus remarkably variable in length. According to Temminck (7/20.
'Coup-d'oeil general sur l'Inde Archipelagique' tome 3 1849 page 177; see
also Mr. Blyth in 'Indian Sporting Review' volume 2 page 5 1856.) the Timor
specimens differ as a local race from that of Java. These several wild
varieties have not as yet been ranked as distinct species; if they should,
as is not unlikely, be hereafter thus ranked, the circumstance would be
quite immaterial as far as the parentage and differences of our domestic
breeds are concerned. The wild G. bankiva agrees most closely with the
black-breasted red Game-breed, in colouring and in all other respects,
except in being smaller, and in the tail being carried more horizontally.
But the manner in which the tail is carried is highly variable in many of
our breeds, for, as Mr. Brent informs me, the tail slopes much in the
Malays, is erect in the Games and some other breeds, and is more than erect
in Dorkings, Bantams, etc. There is one other difference namely, that in G.
bankiva, according to Mr. Blyth, the neck-hackles when first moulted are
replaced during two or three months not by other hackles, as with our
domestic poultry, but by short blackish feathers. (7/21. Mr. Blyth 'Annals
and Mag. of Nat. Hist.' 2nd series volume 1 1848 page 455.) Mr. Brent,
however, has remarked that these black feathers remain in the wild bird
after the development of the lower hackles, and appear in the domestic bird
at the same time with them: so that the only difference is that the lower
hackles are replaced more slowly in the wild than in the tame bird; but as
confinement is known sometimes to affect the masculine plumage, this slight
difference cannot be considered of any importance. It is a significant fact
that the voice of both the male and female G. bankiva closely resembles, as
Mr. Blyth and others have noted, the voice of both sexes of the common
domestic fowl; but the last note of the crow of the wild bird is rather
less prolonged. Captain Hutton, well known for his researches into the
natural history of India, informs me that he has seen several crossed fowls
from the wild species and the Chinese bantam; these crossed fowls BRED
FREELY with bantams, but unfortunately were not crossed inter se. Captain
Hutton reared chickens from the eggs of the Gallus bankiva; and these,
though at first very wild, afterwards became so tame that they would crowd
round his feet. He did not succeed in rearing them to maturity; but as he
remarks, "no wild gallinaceous bird thrives well at first on hard grain."
Mr. Blyth also found much difficulty in keeping G. bankiva in confinement.
In the Philippine Islands, however, the natives must succeed better, as
they keep wild cocks to fight with their domestic game-birds. (7/22.
Crawfurd 'Desc. Dict. of Indian Islands' 1856 page 112.) Sir Walter Elliot
informs me that the hen of a native domestic breed of Pegu is
undistinguishable from the hen of the wild G. bankiva; and the natives
constantly catch wild cocks by taking tame cocks to fight with them in the
woods. (7/23. In Burmah, as I hear from Mr. Blyth, the wild and tame
poultry constantly cross together, and irregular transitional forms may be
seen.) Mr. Crawfurd remarks that from etymology it might be argued that the
fowl was first domesticated by the Malays and Javanese. (7/24. Ibid page
113.) It is also a curious fact, of which I have been assured by Mr. Blyth,
that wild specimens of the Gallus bankiva, brought from the countries east
of the Bay of Bengal, are far more easily tamed than those of India; nor is
this an unparalleled fact, for, as Humboldt long ago remarked, the same
species sometimes evinces a more tameable disposition in one country than
in another. If we suppose that the G. bankiva was first tamed in Malaya and
afterwards imported into India, we can understand an observation made to me
by Mr. Blyth, that the domestic fowls of India do not resemble the wild G.
bankiva of India more closely than do those of Europe.]
From the extremely close resemblance in colour, general structure, and
especially in voice, between Gallus bankiva and the Game fowl; from their
fertility, as far as this has been ascertained, when crossed; from the
possibility of the wild species being tamed, and from its varying in the
wild state, we may confidently look at it as the parent of the most typical
of all the domestic breeds, namely, the Game fowl. It is a significant
fact, that almost all the naturalists in India, namely Sir W. Elliot, Mr.
S.N. Ward, Mr. Layard, Mr. J.C. Jerdon, and Mr. Blyth (7/25. Mr. Jerdon in
the 'Madras Journ. of Lit. and Science' volume 22 page 2 speaking of G.
bankiva says "unquestionably the origin of most of the varieties of our
common fowls." For Mr. Blyth see his excellent article in 'Gardener's
Chronicle' 1851 page 619; and in 'Annals and Mag. of Nat. Hist.' volume 20
1847 page 388.), who are familiar with G. bankiva, believe that it is the
parent of most or all our domestic breeds. But even if it be admitted that
G. bankiva is the parent of the Game breed, yet it may be urged that other
wild species have been the parents of the other domestic breeds; and that
these species still exist, though unknown, in some country, or have become
extinct. The extinction, however, of several species of fowls, is an
improbable hypothesis, seeing that the four known species have not become
extinct in the most ancient and thickly peopled regions of the East. There
is, in fact, not one other kind of domesticated bird, of which the wild
parent-form is unknown, that is become 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 (7/26. 'Gardener's Chronicle' 1851 page 619.), generally have
a restricted range: we see this well illustrated in India, where the genus
Gallus inhabits the base of the Himalaya, and is succeeded higher up by
Gallophasis, and still higher up by Phasianus. Australia, with its islands,
is out of the question as the home for unknown species of the genus. It is,
also, as improbable that Gallus should inhabit South America (7/27. I have
consulted an eminent authority, Mr. Sclater, on this subject, and he thinks
that I have not expressed myself too strongly. I am aware that one ancient
author, Acosta, speaks of fowls as having inhabited S. America at the
period 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 volume 2 page 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 appearance was somewhat changed. Hence it
is not a little doubtful, notwithstanding the statement of the natives,
whether these birds really were fowls. That the fowl has become feral on
several islands is certain. Mr. Fry, a very capable judge, informed Mr.
Layard, in a letter, 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 unfortunately 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 page 62), and in the Ladrones
(Anson's Voyage). Those found in the Pellew 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.) as that a humming-bird
should be found in the Old World. From the character of the other
gallinaceous birds of Africa, it is not probable that Gallus is an African
genus. We need not look to the western parts of Asia, for Messrs. Blyth and
Crawfurd, who have attended to this subject, doubt whether Gallus ever
existed in a wild state even as far west as Persia. Although the earliest
Greek writers speak of the fowl as a Persian bird, this probably merely
indicates its line of importation. For the discovery of unknown species we
must look to India, to the Indo-Chinese countries, and to the northern
parts of the Malay Archipelago. The southern portion of China is the most
likely country; but as Mr. Blyth informs me, skins have been exported from
China during a long period, and living birds are largely kept there in
aviaries, so that any native species of Gallus would probably have become
known. Mr. Birch, of the British Museum, has translated for me passages
from a Chinese Encyclopaedia published in 1609, but compiled from more
ancient documents, in which it is said that fowls are creatures of the
West, and were introduced into the East (i.e. China) in a dynasty 1400 B.C.
Whatever may be thought of so ancient a date, we see that the Indo-Chinese
and Indian regions were formerly considered by the 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 discovered.
In considering whether the domestic breeds are descended 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
improbability 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 no 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 occasionally be met with, which
are almost identical in plumage with the wild G. bankiva. This is a fact
well deserving 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
breeding cocks of the golden-spangled variety is their tendency 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" (7/28. Mr. Hewitt in 'The
Poultry Book' by W.B. Tegetmeier 1866 page 248.), 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 individual bird. With Spanish, Polish, pencilled
Hamburgh, silver-spangled Hamburgh fowls, and with some other less common
breeds, I have never heard of a black-breasted red bird having appeared.
From my experience with pigeons, I made the following crosses. I first
killed all my own poultry, no others living near my house, and then
procured, by Mr. Tegetmeier'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 plumage is strongly inherited, and that the
belief in the prepotent 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-coloured instead of black. In this case, with advancing age
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
(7/29. 'Journal of Horticulture' January 14, 1862 page 325.) by a breeder,
that he crossed two silver-pencilled Hamburgh hens with a Spanish cock, and
reared a number of chickens, all of which were black, the cocks having
GOLDEN and the hens brownish hackles; so that in this instance likewise
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 male 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 Came cock;"--now this sub-breed can be
produced, as I am informed by Mr. Tegetmeier, by crossing a black-breasted
red Game cock with a white Game hen, and the "pile" sub-breed thus produced
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-
inheritance of this character has 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 gorgeous bird, so much so
that an acquaintance had it preserved and stuffed simply from its beauty.
When stalking about it closely resembled the wild Gallus bankiva, but with
the red feathers rather darker. On close comparison 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 Spanish breed has
long been known to breed true, and no instance 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 described 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, besides 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.
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, became
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 (7/30. 'Die Huhner- und Pfauenzucht' Ulm 1827 s. 17. For Mr.
Hewitt's statement with respect to the white Silk fowl see the 'Poultry
Book' by W.B. Tegetmeier 1866 page 222. I am indebted to Mr. Orton for a
letter on the same subject.) a distinct breed with black bones, and with
black, not silky plumage, has likewise been observed to degenerate.
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 hens of some other species of Gallus are 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 analogous variation, as neither
the wild rock pigeon nor any closely allied species has such feathers. The
frequent appearance of pencilling in crossed birds probably accounts 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 cuckoo-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
characteristic 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 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 crossing distinct breeds; but it is a case of analogous
variation, 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 inexplicable; 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. (7/31.
Dixon 'Ornamental and Domestic Poultry' pages 253, 324, 335. For game fowls
see Ferguson on 'Prize Poultry' page 260.) The editor of the 'Poultry
Chronicle' (7/32. 'Poultry Chronicle' volume 2 71.) remarks that all the
breeds which properly have red ear-lappets occasionally produce birds with
white ear-Tappets. This remark more especially 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 generally, but not invariably,
white in India.
In concluding this part of my subject, I may repeat that there exists one
widely-ranging, varying, and common species of Gallus, namely, G. bankiva,
which can be tamed, produces fertile offspring when crossed with common
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 domesticated breed. We have seen that there is much difficulty in
believing that other, now unknown, species have been the parents of the
other domestic breeds. We know 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 Cochins, with their
imperfect tail and small wings, bear in 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 were once
formed, crossing would come into play in changing their character and in
increasing their number. Brahma Pootras, 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
Malayan 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.
Rutimeyer found no remains of the fowl in the ancient Swiss lake-dwellings;
but, according to Jeitteles (7/33. 'Die vorgeschichtlichen Alterthumer' II.
Theil 1872 page 5. Dr. Pickering in his 'Races of Man' 1850 page 374 says
that the head and neck of a fowl is carried in a Tribute-procession to
Thoutmousis III. (1445 B.C.); but Mr. Birch of the British Museum doubts
whether the figure can be identified as the head of a fowl. Some caution 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
prejudice against this bird. I am informed by the Rev. S. Erhardt that on
the east coast of Africa, from 4 to 6 deg south of the equator, most of the
pagan tribes at the present day hold the fowl in aversion. The natives of
the Pellew Islands would not eat the fowl nor will the Indians in some
parts of S. America. For the ancient history of the fowl see also Volz
'Beitrage zur Culturgeschichte' 1852 s. 77; and Isid. Geoffroy St.-Hilaire
'Hist. Nat. Gen.' tome 3 page 61. Mr. Crawfurd has given an admirable
history of the fowl in his paper 'On the Relation of Domesticated Animals
to Civilisation' read before the Brit. Assoc. at Oxford in 1860 and since
printed separately. I quote from him on the Greek poet Theognis, and on the
Harpy Tomb described by Sir C. Fellowes. I quote from a letter of Mr.
Blyth's with respect to the Institutes of Manu.), such have certainly since
been found associated with extinct animals and prehistoric remains. It is,
therefore a strange fact that the fowl is not mentioned in the Old
Testament, nor figured on the ancient Egyptian monuments. It is not
referred to by 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, between the sixth and seventh centuries B.C., of
which Mr. Layard sent me an impression; and on the Harpy Tomb in Lycia,
about 600 B.C.: so that the fowl apparently reached Europe in a
domesticated condition somewhere about 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 Caesar. 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 authority 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 Encyclopaedia, 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
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 true Bantam, is
referred to in an old Japanese Encyclopaedia, as I am informed by Mr.
Birch. In the Chinese Encyclopaedia published in 1596, but compiled from
various sources, some of high antiquity, seven breeds are mentioned,
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 Aldrovandi "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 have been much improved and modified since his time;
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 feathers. Mr. Dixon (7/34. 'Ornamental and Domestic
Poultry' 1847 page 185; for passages translated from Columella see page
312. For Golden Hamburghs see Albin 'Natural History of Birds' 3 volumes
with plates 1731-38.) 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 anatomical 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
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 difference. Beginning
with the eggs and chickens, I will pass on to their secondary sexual
characters, and then to their differences in external structure and in the
skeleton. I enter on the following details chiefly to show how variable
almost every character has become under domestication.
[EGGS.
Mr. Dixon remarks (7/35. 'Ornamental and Domestic Poultry' page 152.) that
"to every hen belongs an individual 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 hand-writing 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 (7/36.
Ferguson on 'Rare Prize Poultry' page 297. This writer, I am informed,
cannot generally be trusted. He gives, however, figures and much
information on eggs. See pages 34 and 235 on the eggs of the Game fowl.);
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
(7/37. See 'Poultry Book' by Mr. Tegetmeier 1866 pages 81 and 78.) 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-breed of Spanish, are said to lay
harder eggs than true Spanish. (7/38. 'The Cottage Gardener' October 1855
page 13. On the thinness of the eggs of Game-fowls see Mowbray on 'Poultry'
7th edition page 13.) 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. 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 hens 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 similarly striped. (7/39. My information, which is very far from
perfect, on chickens in the down, is derived chiefly from Mr. Dixon's
'Ornamental 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 'Poultry Book' 1866 page 221.) 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 Bantams, display a new
character, for their chickens have their breasts and throats more or less
white, with sometimes a little white elsewhere. 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 Dorkings; by the partridge and grouse-coloured sub-breeds of Cochins
(Brent), but not, as we have seen, by the 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 young 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 stripes.]
With respect to the period of life at which the characters 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
protuberance of the anterior part of the skull is well developed before the
chickens come out of the egg (7/40. As I hear from Mr. Tegetmeier; see also
'Proc. Zoolog. Soc.' 1856 page 366. On the late development of the crest
see 'Poultry Chronicle' volume 2 page 132.) 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 magnificent comb, and this is developed at an unusually early 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. (7/41. On these points see 'Poultry Chronicle' volume
3 page 166; and Tegetmeier 'Poultry Book' 1866 pages 105 and 121.) Cochins
are characterised by a small tail, and in the young cocks the tail is
developed at an unusually late period. (7/42. Dixon 'Ornamental and
Domestic Poultry' page 273.) Game fowls are notorious for their pugnacity;
and the young cocks crow, clap their little wings, and fight obstinately
with each other, even whilst under their mother's care. (7/43. Ferguson on
'Rare and Prize Poultry' page 261.) "I have often had," says one author
(7/44. Mowbray on 'Poultry' 7th edition 1834 page 13.), "whole broods,
scarcely feathered, stone-blind from fighting; the rival couples moping in
corners, and renewing their battles on obtaining the first ray of light."
The weapons and pugnacity of all male gallinaceous birds evidently serve
the purpose of gaining possession of the females; so that the tendency in
our Game chickens to fight at an extremely early age is not only useless,
but injurious, as they suffer much from their wounds. The training for
battle during an early age may be natural to the wild Gallus bankiva; but
as man during many generations has gone on selecting the most obstinately
pugnacious cocks, it is more probable that their pugnacity has been
unnaturally increased, and unnaturally transferred to the young male
chickens. In the same manner, it is probable that the extraordinary
development 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 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
difference in plumage. Mr. Brent informs me that he has seen two strains of
black-breasted red Games, of 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 "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. (7/45. See the full description
of the varieties of the Game-breed in Tegetmeier 'Poultry Book' 1866 page
131. For Cuckoo Dorkings page 97.) 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, excepting of course in the
hackles, crest, and beard. In spangled Hamburghs, there is likewise a
considerable degree of similarity between the two sexes. In pencilled
Hamburghs, on the other hand, there is much dissimilarity; the pencilling
which is characteristic of the hens being 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 females, 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 (7/46. Mr. Hewitt in
Tegetmeier 'Poultry Book' 1866 pages 246 and 156. For hen-tailed game-cocks
see page 131.), 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 flowing sickle-like tail-feathers. A hen-tailed sub-breed
of Hamburghs was recently much esteemed. There is also a breed of Game-
fowls, in which the males and females resemble 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. (7/47. 'The Field'
April 20, 1861. The writer says he has seen half-a-dozen cocks thus
sacrificed.) 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
(7/48. 'Proceedings of Zoolog. Soc.' March 1861 page 102. The engraving of
the hen-tailed cock just alluded to was exhibited before the Society.) has
recorded the remarkable case of a brown-breasted red Game cock which, after
assuming its perfect masculine plumage, became hen-feathered in the autumn
of the following 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 celebrated
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 seasons 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." (7/49. 'The Field' April 20, 1861.)
I have remarked in my 'Origin of Species' that secondary sexual characters
are apt to differ much in the species of the same genus, and to be
unusually variable in the individuals of the same 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 (7/50. 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), and its form is eminently characteristic of each
kind, with the exception of the Dorkings, in which the form has not been as
yet determined 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 developed in Spanish fowls; and in a local breed called
Red-caps, it is sometimes "upwards of three inches in breadth at the front,
and more than four inches in length, measured to the end of the peak
behind." (7/51. The 'Poultry Book' by Tegetmeier 1866 page 234.) In some
breeds the comb is double, and when the two ends are cemented together it
forms a "cup-comb;" in the "rose-comb" it is depressed, covered with small
projections, and produced backwards; in the horned and creve-coeur 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 rise 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 developed, that I have seen birds which could hardly pick up their
food; and a German writer asserts (7/52. 'Die Huhner- und Pfauenzucht' 1827
s. 11.) that they are in consequence liable to be struck by hawks.
Monstrous 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; in certain Polish sub-breeds they are replaced 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 male birds
display extraordinarily-shaped feathers, such as naked shafts with discs at
the end, etc., 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 re-clothed 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 barbs. Hence the coloured tips appeared like little
separate 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 flowing 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 Dorkings and Gaines; but droop much in Malays and
in some Cochins. Sultans are characterised by an additional number 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
(7/53. 'Poultry Chronicle' volume 1 page 595. Mr. Brent has informed me of
the same fact. With respect to the position of the spurs in Dorkings see
'Cottage Gardener' September 18, 1860 page 380.); and in birds of this
breed the spur is often placed almost on the outside of the leg. Double
spurs are mentioned in an ancient Chinese Encyclopaedia. 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 distinguish the sexes in the
Gallinaceae, 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 plumage between the
male and female, the male is always the most beautiful; but in golden-
spangled Hamburghs the hen is equally beautiful with the cock, and
incomparably 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 gallinaceous 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 the males of certain other sub-breeds,
which from the hen having a small crest, are called lark-crested, "a single
upright comb sometimes almost entirely takes the place of the crest."
(7/54. Dixon 'Ornamental and Domestic Poultry' page 320.) From this latter
case, and more especially from some facts presently to be given with
respect to the protuberance of the skull in Polish fowls, the crest in this
breed must 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 transferred to the female, for her comb is
unusually large, though not upright. In Game fowls the bold and savage
disposition of the male has likewise been largely transferred to the female
(7/55. Mr. Tegetmeier informs me that Game hens have been found so
combative, that it is now generally the practice to exhibit each hen in a
separate pen.); and she sometimes even possesses the eminently masculine
character of spurs. Many cases are on record of fertile hens being
furnished with spurs; and in Germany, according to Bechstein (7/56.
'Naturgeschichte Deutschlands' b. 3 1793 s. 339, 407.), the spurs in the
Silk hen are sometimes very long. He mentions also another breed similarly
characterised, in which the hens are excellent layers, but are apt to
disturb and break their eggs owing to their spurs.
Mr. Layard (7/57. On the Ornithology of Ceylon in 'Annals and Mag. of Nat.
History.' 2nd series volume 14 1854 page 63.) 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 described 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 found the
same rule to hold 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 includes 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 (7/58. 'Handbuch der
vergleich. Anatomie' 1805 page 85 note. Mr. Tegetmeier, who gives in 'Proc.
Zoolog. Soc.' November 25, 1856 a very interesting account of the skulls of
Polish fowls, not knowing of Bechstein's account, has disputed the accuracy
of Blumenbach's statement. For Bechstein see 'Naturgeschichte Deutschlands'
b. 3 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 furnished with a
comb.), who particularly attended to abnormal peculiarities in domestic
animals, states, in 1805, that this was the case; and Bechstein had
previously, in 1793 observed the same fact. This latter author has
carefully described the effects on the skull of a crest not only in the
case of fowls, but of 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 on a bony
protuberance of variable size. He well describes the peculiarities of this
protuberance; he attended also 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 remarks that he never observed this
protuberance in male fowls. Hence there can be no doubt that this
extraordinary character in the skulls of Polish fowls was formerly in
Germany confined 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 variable 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! Crossing often causes strange
modification 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 results, as Mr.
Tegetmeier tells me, from crossing white Cochins with black Spanish fowls,
or white Dorkings with black Minorcas. (7/59. 'Cottage Gardener' January 3,
1860 page 218.) A good observer (7/60. Mr. Williams in a paper read before
the Dublin Nat. Hist. Soc. quoted in 'Cottage Gardener' 1856 page 161.)
states that a first-rate silver-spangled Hamburgh hen gradually lost the
most characteristic qualities of the breed, for the black lacing to her
feathers 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 marked manner; and
chickens produced from this latter hen were at first almost pure white,
"but on moulting acquired black colours 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 mocking, as M. Godron (7/61. 'De l'Espece' 1859 page 442. For
the occurrence of black-boned fowls in South America, see Roulin in 'Mem.
de l'Acad. des Sciences' tome 6 page 351; and Azara, 'Quadrupedes du
Paraguay' tome 2 page 324. A frizzled fowl sent to me from Madras had black
bones.) 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 differ much.
The beak varies slightly in length and curvature, but incomparably less
than with pigeons. In most crested fowls the nostrils 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 counted, 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 primary
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 feathers, 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 one which I possessed there was no oil-
gland; 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 told, the breed
had kept true for twenty years; but rumpless fowls often produce chickens
with tails. (7/62. Mr. Hewitt in Tegetmeier 'Poultry Book' 1866 page 231.)
An eminent physiologist (7/63. Dr. Broca in Brown-Sequard 'Journal de
Phys.' tome 2 page 361.) 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 (7/64. Dixon 'Ornamental Poultry' page 325.) 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 examined. Prof. Hoffmann has
sent me a sketch of the feet of a fowl of the common breed at Giessen, with
a web extending between the three toes, for about a third of their length.
In Cochins the middle toe is said (7/65. 'Poultry Chronicle' volume 1 page
485. Tegetmeier 'Poultry Book' 1866 page 41. On Cochins grazing ibid page
46.) 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
surprisingly 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 (7/66. Ferguson on 'Prize Poultry' page 87.) have a loud,
deep, somewhat prolonged crow, but with considerable individual difference.
Colonel 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 "prolonged howling screech" of the
cocks in Sikhim. (7/67. Col. Sykes in 'Proc. Zoolog. Soc.' 1832 page 151.
Dr. Hooker's 'Himalayan Journals' volume 1 page 314.) 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 Cochins. The latter, it has been asserted, "graze
to a much greater extent than any other varieties." The Spanish fowls
suffer more from frost than other breeds.]
Before we pass on to the skeleton, the degree of distinctness 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.
bankiva. 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, from 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 coloured 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 skulls I owe to
the kindness of Mr. Tegetmeier, and three of the skeletons to Mr. Eyton.
SKULL.
(FIGURE 33. OCCIPITAL FORAMEN, of natural size. A. Wild Gallus bankiva. B.
Cochin Cock.
FIGURE 34. SKULLS of natural size, viewed from above, a little obliquely.
A. Wild Gallus bankiva. B. White-crested Polish Cock.
FIGURE 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.
FIGURE 36. SKULL OF HORNED FOWL, of natural size, viewed from above, a
little obliquely. (In the possession of Tegetmeier.))
[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.
bankiva as the standard of comparison. 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 worth notice occur. 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. bankiva. In a Bantam or Jumper from Burmah these
same characters are more strongly pronounced, and the supra-occiput is more
pointed. In a black Bantam the skull is not so globular, and the occipital
foramen is very large, and has nearly the same sub-triangular outline
presently to be described in Cochins; and in this skull the two ascending
branches of the premaxillary are overlapped in a singular 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 developed in the hens. The
pterygoids, and the processes of the lower jaw, are broader, relatively to
the size of the head, than in G. bankiva; and this is likewise the case
with Dorkings when of large size. The 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 horizontal 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.
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 that of G. bankiva, the breadth
between the orbits was exactly double. Of Hamburghs I have examined four
skulls (male and female) 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 pre-maxillary 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, flattened 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 different sub-
breeds of English Polish fowls. The hemispherical protuberance of the
frontal bones (7/68. See Mr. Tegetmeier's account with woodcuts of the
skull of Polish fowls in 'Proc. Zoolog. Soc.' November 25, 1856. For other
references see Isid. Geoffroy Saint-Hilaire 'Hist. Gen. des Anomalies' tome
1 page 287. M. C. Dareste suspects ('Recherches sur les Conditions de la
Vie' etc. Lille 1863 page 36) that the protuberance is not formed by the
frontal bones, but by the ossification of the dura mater.) may be seen in
figure 34, 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 figure 35 longitudinal sections are given of the skull of a
Polish fowl, and, for comparison, of a Cochin of the same size. The
protuberance in all Polish fowls occupies the same position 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
surprisingly great. The brain is modified in a corresponding manner, as is
shown in the two longitudinal sections, which deserve attentive
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. This cavity, as I hear from Mr. Tegetmeier, is entirely filled with
brain. In the skull of the Cochin and of all ordinary fowls a strong
internal ridge of bone separates the anterior from the central cavity; but
this ridge is quite absent in the Polish skull here figured. The shape of
the central cavity is circular in the Polish, and lengthened in the Cochin
skull. 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 (figure
35), a perfect gradation in the configuration of each part of the internal
surface could be traced. In the Polish skull, with a small protuberance,
the ridge between the anterior and middle cavities was 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 modifications in the
form of the brain affect the intellect of Polish fowls; some writers have
stated that they are extremely stupid, but Bechstein and Mr. Tegetmeier
have shown that this is by no means generally the case. Nevertheless
Bechstein (7/69. 'Naturgeschichte Deutschlands' b. 3 1793 s. 400.) 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. (7/70. The 'Field' May 11, 1861. I have received
communications to a similar effect from Messrs. Brent and Tegetmeier.)
To return to the skull of Polish fowls. 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 processes 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 perpendicular, 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 differences. The palatine bones are a little more
curved upwards at their posterior ends. The frontal bones, anteriorly 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 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 crescentic outline.
I must still say a few words on some of the foreign Crested breeds. The
skull of a crested, rumpless, white Turkish fowl was very slightly
protuberant, and but little perforated; the ascending branches of the
premaxillary were well developed. In another Turkish breed, called
Ghoondooks, the skull was considerably protuberant and perforated; the
ascending branches of the premaxillary were so much aborted that they
projected only 1/15th of an inch; and the inner processes of the nasal bone
were so completely aborted, that the surface where they should have
projected was 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 nasal 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 by Mr.
Tegetmeier: it resembles a Polish skull in most of its characters, 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 medial 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
projections 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 protuberance 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. A series may 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 protuberance 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
selected with a large bony protuberance, when adult they will have a large
crest. There can be no doubt that in former times the breeder of Polish
fowls attended solely to the crest, and not to the skull; nevertheless, by
increasing the crest, in which he has been wonderfully successful, 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 premaxillary 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.
VERTEBRAE.
(FIGURE 37. SIXTH CERVICAL VERTEBRA, natural size, viewed laterally. A.
Wild Gallus bankiva. B. Cochin cock.)
In G. bankiva there are fourteen cervical, seven dorsal with ribs,
apparently fifteen lumbar and sacral, and six caudal vertebrae (7/71. It
appears that I have not correctly designated the several groups of
vertebrae, for a great authority, Mr. W.K. Parker ('Transact. Zoolog. Soc.'
volume 5 page 198) specifies 16 cervical, 4 dorsal, 15 lumbar, and 6 caudal
vertebrae in this genus. But I have used the same terms in all the
following descriptions.); 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 vertebrae in the several breeds difficult. I have spoken of
six caudal vertebrae, because the basal one is almost completely anchylosed
with the pelvis; but if we consider the number as seven, the caudal
vertebrae agree in all the skeletons. The cervical vertebrae 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
importance, for all the cervical vertebrae 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 vertebrae (7/72.
Macgillivray 'British Birds' volume 1 page 25.), we need feel no surprise
at the number of the cervical vertebrae in the fowl being, as it appears,
variable.
There are seven dorsal vertebrae 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 vertebrae 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 vertebrae. So
that the degree to which these middle dorsal vertebrae are anchylosed 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 lumbar in G. bankiva; the
sternal portion of both the seventh and eighth ribs did not reach the
sternum. In four skeletons in which ribs 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 skeletons; in this Game cock, as far as could be judged from the
appearance of the lumbar vertebrae, 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 extremely broad in Cochins, and is
completely ossified. As previously stated, it is scarcely possible to count
the lumbo-sacral vertebrae; but they certainly do not correspond in shape
or number in the several skeletons. The caudal vertebrae 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 vertebrae were a little
elongated. In three rumpless fowls the caudal vertebrae were few in number,
and anchylosed together into a misformed mass.
In the individual vertebrae the differences in structure are very slight.
In the atlas the cavity for the occipital condyle is either ossified into a
ring, or is, as in Bankiva, open on its upper margin. The upper arc of the
spinal canal is a little more arched in Cochins, in conformity with
the shape of the occipital foramen, than in G. bankiva. In several
skeletons a difference, but not of much importance, may be observed, which
commences at the fourth cervical vertebra, and is greatest at about the
sixth, seventh, or eighth vertebra; this consists in the haemal descending
processes being united 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 developed than in the corresponding vertebra of
the Game fowl or G. bankiva.
PELVIS.
This differs in some few points in the several skeletons. 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 vertebrae; the outline, however, does differ
in being more truncated in Bantams, and more rounded in certain breeds, as
in Cochins. The outline of the ischiadic 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.
(FIGURE 38. EXTREMITY OF THE FURCULA, of natural size, viewed laterally. A.
Wild Gallus bankiva. B. Spangled Polish Fowl. C. Spanish Fowl. D. Dorking
Fowl.)
STERNUM.
This bone is generally so much deformed that it is scarcely possible to
compare its shape strictly in the several breeds. The form of the
triangular extremity of the lateral processes differs considerably, being
either almost equilateral or much elongated. The front margin of the crest
is more or less perpendicular and varies greatly, as does the curvature of
the posterior end, and the flatness of the lower surface. The outline of
the manubrial process also varies, being wedge-shaped in the Bankiva, and
rounded in the Spanish breed. The FURCULUM differs in being more or less
arched, and greatly, as may be seen in the accompanying outlines, in the
shape of the terminal plate; but the shape of this part differed a little
in two skeletons of the wild Bankiva. The CORACOID presents no difference
worth notice. The SCAPULA varies in shape, being of nearly uniform breadth
in Bankiva, much broader in the middle in the Polish fowl, and abruptly
narrowed towards the apex in the two Sultan fowls.
I carefully compared each separate bone of the leg and wing, relatively 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 absolutely 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 varied 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 differences, 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 vertebrae, and the end of the scapula in both was somewhat
attenuated. In the skull, the deep medial furrow in the frontal bones and
the vertically elongated occipital 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 depressed, 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 before specified, are eminently characteristic of Polish and
other Crested fowls.
But the most striking result of my 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 structure;
fowls have been exposed to unnatural conditions of life, and their whole
organisation has thus been rendered variable; but the breeder is quite
indifferent to, and never intentionally selects, any modification in the
skeleton. 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,--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
modification in the skeleton is related to some external character 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 furculum, 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.
[EFFECTS OF THE DISUSE OF PARTS.
Judging from the habits of our European gallinaceous birds, Gallus 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-feathers, cannot fly at all; and there is
reason to believe that both these breeds are ancient, so that their
progenitors during many generations 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 Burmese
Jumper, which has unnaturally short legs, are slightly shortened relatively
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 measurements 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 Table 7.I. The calculated weight of the
wing-bones relatively to the 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. (7/73. 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-bones in G.
bankiva, Cochins, Dorkings, etc. And now taking 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.)
TABLE 7.I. (Weights in grains.)
COLUMN 1. Actual Weight of Femur and Tibia.
COLUMN 2. Actual Weight of Humerus and Ulna.
COLUMN 3. Weight of Wing-bones relatively to the Leg-bones in comparison
with these same bones in Gallus bankiva.
1. 2. 3.
Gallus bankiva -- wild male 86 54 100
1. Cochin -- male 311 162 83
2. Dorking -- male 557 248 70
3. Spanish (Minorca) -- male 386 183 75
4. Gold-Spangled Polish -- male 306 145 75
5. Game, black-breasted -- male 293 143 77
6. Malay -- female 231 116 80
7. Sultan -- male 189 94 79
8. Indian Frizzled -- male 206 88 67
9. Burmese Jumper -- female 53 36 108
10. Hamburgh (pencilled) -- male 157 104 106
11. Hamburgh (pencilled) -- female 114 77 108
12. Silk (black-boned) -- female 88 57 103
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 carried 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, relatively to the legs, are
slightly increased in weight; but it should be 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 nature 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 eyesight, but after
having calculated the weights of the leg-bones relatively 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 apprehend, safely
be attributed to disuse.
To make Table 7.I. quite satisfactory, it ought to have been shown that in
the eight first birds the leg-bones have not actually 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. (7/74.
Mr. Blyth (in 'Annals and Mag. of Nat. Hist.' 2nd series volume 1 1848 page
456) gives 3 1/4 pounds 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.) I am indeed inclined to suspect that the leg-bones in the Dorking,
No. 2 in the table, are proportionally too heavy; but this bird was a very
large one, weighing 7 pounds 2 ounces, 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.
TABLE 7.II.
COLUMN 1. Length of Sternum (in inches and decimals.)
COLUMN 2. Depth of Crest of Sternum (in inches and decimals.).
COLUMN 3. Depth of Crest relatively to the length of the Sternum, in
comparison with Gallus bankiva.
1. 2. 3.
Gallus bankiva -- male. 4.20 1.40 100
1. Cochin -- male. 5.83 1.55 78
2. Dorking -- male. 6.95 1.97 84
3. Spanish -- male. 6.10 1.83 90
4. Polish -- male. 5.07 1.50 87
5. Game -- male. 5.55 1.55 81
6. Malay -- female. 5.10 1.50 87
7. Sultan -- male. 4.47 1.36 90
8. Frizzled hen -- male. 4.25 1.20 84
9. Burmese Jumper -- female. 3.06 0.85 81
10. Hamburgh -- male. 5.08 1.40 81
11. Hamburgh -- female. 4.55 1.26 81
12. Silk fowl -- female. 4.49 1.01 66
In Table 7.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 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. (7/75. The third column is calculated
on the same principle as explained in footnote 7/73 above.)
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 furculum, and in
the shape of its sternal extremity. Medical men believe that the abnormal
form of the spine so commonly observed in women of the higher ranks results
from the attached muscles not being fully exercised. So it is with our
domestic 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 deformed to an
extreme degree. Mr. Romanes, however, believes that the malformation is due
to fowls whilst young resting their sternums on the sticks on which they
roost.]
Finally, we may conclude with respect to the various 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 important, subject. In Cochin and Game fowls there is perhaps some
relation between the colour of the plumage and the darkness of the egg-
shell. 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 absent 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 greatly 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 correlation between a crest of feathers and
the imperfectly ossified condition of the skull. Not only does this hold
good with nearly all crested fowls, but likewise with tufted ducks, and as
Dr. Gunther 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
amherstiae), a close relation in colour, as well as in structure, between
the plumes on the head and on the loins. It would therefore appear that the
same law has regulated the state of the feathers on the head and body, both
with species living under natural conditions, and with birds which have
varied under domestication.
CHAPTER 1.VIII.
DUCK--GOOSE--PEACOCK--TURKEY--GUINEA-FOWL--CANARY-BIRD--GOLD-FISH--HIVE-
BEES--SILK-MOTHS.
DUCKS, SEVERAL BREEDS OF.
PROGRESS OF DOMESTICATION.
ORIGIN OF FROM THE COMMON WILD-DUCK.
DIFFERENCES IN THE DIFFERENT BREEDS.
OSTEOLOGICAL DIFFERENCES.
EFFECTS OF USE AND DISUSE ON THE LIMB-BONES.
GOOSE, ANCIENTLY DOMESTICATED.
LITTLE VARIATION OF.
SEBASTOPOL BREED.
PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.
TURKEY, BREEDS OF.
CROSSED WITH THE UNITED STATES SPECIES.
EFFECTS OF CLIMATE ON.
GUINEA-FOWL, CANARY-BIRD, GOLD-FISH, HIVE-BEES.
SILK-MOTHS, SPECIES AND BREEDS OF.
ANCIENTLY DOMESTICATED.
CARE IN THEIR SELECTION.
DIFFERENCES IN THE DIFFERENT RACES.
IN THE EGG, CATERPILLAR, AND COCOON STATES.
INHERITANCE OF CHARACTERS.
IMPERFECT WINGS.
LOST INSTINCTS.
CORRELATED CHARACTERS.
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 dermal sack largely developed.
2. The Rouen, of great size, coloured like the wild duck, with green or
mottled beak; dermal sack largely developed.
3. Tufted Duck, with a large top-knot of fine downy feathers, supported on
a fleshy mass, with the skull perforated beneath. 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 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. (8/1. 'Poultry Chronicle' 1854 volume 2 page 91
and volume 1 page 330.) I presume it is this latter sub-variety which has
been described in France (8/2. Dr. Turral 'Bull. Soc. d'Acclimat.' tome 7
1860 page 541.) 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. (8/3. Willughby's 'Ornithology' by Ray page 381. This breed is
also figured by Albin in 1734 in his 'Nat. Hist. of Birds' volume 2 page
86.) It shows its prolonged domestication by almost incessantly laying
eggs, like the fowls which are called everlasting layers. (8/4. F. Cuvier
in 'Annales du Museum' tome 9 page 128 says that moulting and incubation
alone stops these ducks laying. Mr. B.P. Brent makes a similar remark in
the 'Poultry Chronicle' 1855 volume 3 page 512.)
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 archipelago. It walks with its body extremely erect, and
with its thin neck stretched straight upwards. Beak rather short. Tail
upturned, including only 18 feathers. Femur and metatarsus elongated.]
Almost all naturalists admit that the several breeds are descended from the
common wild duck (Anas boschas); most fanciers, on the other hand, take as
usual a very different view. (8/5. Rev. E.S. Dixon 'Ornamental and Domestic
Poultry' 1848 page 117. Mr. B.P. Brent in 'Poultry Chronicle' volume 3 1855
page 512.) Unless we deny that domestication, prolonged during centuries,
can affect even such unimportant characters as colour, size, and in a
slight degree proportional dimensions and mental disposition, 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 (8/6. Crawfurd on
the 'Relation of Domesticated Animals to Civilisation' read before the
Brit. Assoc. at Oxford 1860.) to the ancient Egyptians, to the Jews of the
Old Testament, and to the Greeks of the Homeric period. About eighteen
centuries ago Columella (8/7. Dureau de La Malle in 'Annales des Sciences
Nat.' tome 17 page 164; and tome 21 page 55. Rev. E.S. Dixon 'Ornamental
Poultry' page 118. Tame ducks were not known in Aristotle's time, as
remarked by Volz in his 'Beitrage zur Kulturgeschichte' 1852 s. 78.) 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 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 language 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 experiment 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
(8/8. I quote this account from 'Die Enten- und Schwanenzucht' Ulm 1828 s.
143. See Audubon 'Ornithological Biography' volume 3 page 168 on the taming
of ducks on the Mississippi. For the same fact in England see Mr. Waterton
in Loudon's 'Mag. of Nat. Hist.' volume 8 1835 page 542; and Mr. St. John
'Wild Sports and Nat. Hist. of the Highlands' 1846 page 129.), as is known
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 (8/9.
Mr. E. Hewitt in 'Journal of Horticulture' 1862 page 773; and 1863 page
39.) has described in detail his often repeated and successful experiments
in domesticating 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 brethren in
undisturbed possession of their foster-mother's care. The difference of
habit at the onset in the newly-hatched ducklings almost entails such a
result to a certainty." 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 always
changed and deteriorated in character in the course of two or three
generations; notwithstanding that great care was taken to prevent their
crossing with tame ducks. After the third generation his birds lost the
elegant carriage 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 destroyed nearly
the whole of his stock and procured fresh eggs from wild nests; so that he
never bred the same family for more than five or six generations. His birds
continued to pair together, and never became polygamous 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 other species
for the parentage of the more distinct breeds, namely, Penguin, Call, Hook-
billed, Tufted, and Labrador ducks. I will not repeat the arguments used in
the previous chapters on the improbability of man having in ancient times
domesticated 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 fertile together (8/10. 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 perfectly
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
Penguins and Labradors were again crossed with Penguins, and subsequently
bred by me inter se, and they were extremely fertile.);--on all the breeds
having the same general disposition, instinct, etc. 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 descended from
distinct species, we must assume that man formerly hit upon species all of
which had this now unique character. Moreover, sub-varieties of each breed
are coloured almost exactly like the wild duck, as I have seen with the
largest and smallest breeds, namely Rouens and Call ducks, and, as Mr.
Brent states (8/11. 'Poultry Chronicle' 1855 volume 3 page 512.), is the
case with Hook-billed ducks. This gentleman, as he informs me, crossed a
white Aylesbury drake and a black Labrador 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 specimens, 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 archipelago, where the wild species does not
exist, with exactly 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 tends to cause the duck to vary much, for Zollinger
(8/12. 'Journal of the Indian Archipelago' volume 5 page 334.), speaking of
the Penguin breed, says that in Lombok "there is an unusual and very
wonderful variety of ducks." One Penguin 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 chestnut-brown, thus more closely
resembling the Mallard.
From 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 resembling in general plumage the wild duck, we may conclude
with confidence that all the breeds are descended from A. boschas.
[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. A good
observer assured me that one year his ducks of this breed laid almost
perfectly white eggs. Another curious case shows what singular variations
sometimes occur and are inherited; Mr. Hansell (8/13. 'The Zoologist'
volumes 7, 8 1849-1850 page 2353.) 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.
(FIGURE 39. SKULLS, viewed laterally, reduced to two-thirds of the natural
size. A. Wild Duck. B. Hook-billed Duck.)
The Hook-billed duck is highly remarkable (see figure 39, of skull); and
its peculiar beak has been inherited at least since the year 1676. This
structure is evidently analogous with that described in the Bagadotten
carrier pigeon. Mr. Brent (8/14. 'Poultry Chronicle' 1855 volume 3 page
512.) says that, when Hook-billed ducks are crossed with common ducks,
"many young ones are produced with the upper mandible shorter than the
lower, which not unfrequently causes the death of the bird." With ducks 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 farm-yard kind, and in a
duck having no other peculiarity 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. But the voice varies in the
different breeds; Mr. Brent (8/15. 'Poultry Chronicle' volume 3 1855 page
312. With respect to Rouens see ditto volume 1 1854 page 167.) says that
Hook-billed ducks are very loquacious, and that Rouens utter a "dull, loud,
and monotonous cry, easily distinguishable by an experienced ear." As the
loquacity of the Call duck is highly serviceable, 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." (8/16. Col. Hawker
'Instructions to young Sportsmen' quoted by Mr. Dixon in his 'Ornamental
Poultry' page 125.) It has been erroneously asserted that Call ducks hatch
their eggs in less time than common ducks. (8/17. 'Cottage Gardener' April
9, 1861.)
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 middle toe, in three specimens, there were twenty-
seven or twenty-eight scutellae, 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 between one of
these birds and the Egyptian goose (Anser aegyptiacus) (8/18. These hybrids
have been described by M. Selys-Longchamps in the 'Bulletins (tome 12 No
10) Acad. Roy. de Bruxelles.'), and likewise with some mongrels which I
raised between the Penguin and Labrador duck. I am not much surprised that
some writers should maintain 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 descendant, much modified by
domestication under an unnatural climate, of Anas boschas.
OSTEOLOGICAL CHARACTERS.
The skulls of the several breeds differ from each other and from the skull
of the wild duck in very little except in the proportional length and
curvature of the premaxillaries. 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 (figure
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 premaxillaries are relatively shorter than in
the wild duck; and the inferior 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
lachrymal bones were produced much further backwards, so as to have a
different shape and nearly to 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 compared them in all the principal breeds;
but excepting in size they presented no difference.
(FIGURE 40.-CERVICAL VERTEBRA, of natural size. A. Eighth cervical vertebra
of Wild Duck viewed on haemal surface. B. Eighth cervical vertebra of Call
Duck, viewed as above. C. Twelfth cervical vertebra of Wild Duck viewed
laterally. D. Twelfth cervical vertebra of Aylesbury Duck, viewed
laterally.)
VERTEBRAE AND RIBS.
In one skeleton of the Labrador duck there were the usual fifteen cervical
vertebrae and the usual nine dorsal vertebrae bearing ribs; in the other
skeleton there were fifteen cervical and ten dorsal vertebrae with ribs;
nor, as far as could be judged, was this owing merely to a rib having been
developed on the first lumbar vertebra; for in both skeletons the lumbar
vertebrae 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 dorsal vertebrae; in a third skeleton small ribs were attached to
the so-called fifteenth cervical vertebra, making ten pairs of ribs; but
these ten ribs do not correspond, or arise from the same vertebra, with the
ten in the above-mentioned Labrador duck. In the Call duck, which had small
ribs attached to the fifteenth cervical vertebra, the haemal spines of the
thirteenth and fourteenth (cervical) and of the seventeenth (dorsal)
vertebrae corresponded with the spines on the fourteenth, fifteenth, and
eighteenth vertebrae of the wild duck: so that each of these vertebrae had
acquired a structure proper to one posterior to it in position. In the
eighth cervical vertebra of this same Call duck (figure 40, B), the two
branches of the haemal spine stand much closer together than in the wild
duck (A), and the descending haemal processes are much shortened. In the
Penguin duck the neck from its thinness and erectness falsely appears (as
ascertained by measurement) to be much elongated, but the cervical and
dorsal vertebrae present no difference; the posterior dorsal vertebrae,
however, are more completely anchylosed to the pelvis than in the wild
duck. The Aylesbury duck has fifteen cervical and ten dorsal vertebrae
furnished with ribs, but the same number of lumbar, sacral, and caudal
vertebrae, as far as could be traced, as in the wild duck. The cervical
vertebrae in this same duck (figure 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 twelfth cervical vertebra in
these two birds. From the foregoing statements we see that the fifteenth
cervical vertebra occasionally becomes modified into a dorsal vertebra, and
when this occurs all the adjoining vertebrae are modified. We also see that
an additional dorsal vertebra bearing a rib is occasionally developed, the
number of the cervical and lumbar vertebrae 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 elongated. In the sternum,
furculum, coracoids, and scapulae, the differences 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 the Penguin 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 upright 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 Table 8.I.
TABLE 8.I.a.
COLUMN 1. Length of Femur, Tibia, and Metatarsus together (inches).
COLUMN 2. Length of Humerus, Radius, and Metacarpus together (inches).
COLUMN 3. Or as (ratio).
Name of Breed. 1. 2. 3.
Wild mallard. 7.14 9.28 100:129
Aylesbury. 8.64 10.43 100:120
Tufted (Dutch). 8.25 9.83 100:119
Penguin. 7.12 8.78 100:123
Call. 6.20 7.77 100:125
TABLE 8.I.b.
COLUMN 1. Length of Femur, Tibia, and Metatarsus together (inches).
COLUMN 2. Length of all the bones of the wing (inches).
COLUMN 3. Or as (ratio).
Name of Breed. 1. 2. 3.
Wild duck (another specimen). 6.85 10.07 100:147
Common domestic duck. 8.15 11.26 100:138
In table 8.I we see, by comparison with the wild duck, that the reduction
in the length of the bones of the wing, relatively to those of the legs,
though slight, is universal. The reduction 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 Table 8.II:
TABLE 8.II.a.
COLUMN 1. Weight of Femur, Tibia, and Metatarsus (grains).
COLUMN 2. Weight of Humerus, Radius, and Metacarpus (grains).
COLUMN 3. Or as (ratio).
Name of Breed. 1. 2. 3.
Wild mallard. 54 97 100:179
Aylesbury. 164 204 100:124
Hooked-bill. 107 160 100:149
Tufted (Dutch). 111 148 100:133
Penguin. 75 90.5 100:120
Labrador. 141 165 100:117
Call. 57 93 100:163
TABLE 8.II.b.
COLUMN 1. Weight of all the Bones of the Leg and Foot (grains).
COLUMN 2. Weight of all the Bones of the Wing (grains).
COLUMN 3. Or as (ratio).
Name of Breed. 1. 2. 3.
Wild (another specimen). 66 115 100:173
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 actual decrease in the wing-
bones, but from the increased weight and length of the bones of the legs.
TABLE 8.III.a.
COLUMN 1. Weight of entire Skeleton (grains). (N.B. One Metatarsus and Foot
was removed from each skeleton, as it had been accidentally lost in two
cases.)
COLUMN 2. Weight of Femur, Tibia, and Metatarsus (grains).
COLUMN 3. Or as (ratio).
Name of Breed. 1. 2. 3.
Wild mallard. 839 54 1000:64
Aylesbury. 1925 164 1000:85
Tufted (Dutch). 1404 111 1000:79
Penguin. 871 75 1000:86
Call (from Mr. Fox). 717 57 1000:79
TABLE 8.III.b.
COLUMN 1. Weight of entire Skeleton (grains). (N.B. One Metatarsus and Foot
was removed from each skeleton, as it had been accidentally lost in two
cases.)
COLUMN 2. Weight of Humerus, Radius and Metacarpus (grains).
COLUMN 3. Or as (ratio).
Name of Breed. 1. 2. 3.
Wild mallard. 839 97 1000:115
Aylesbury. 1925 204 1000:105
Tufted (Dutch). 1404 148 1000:105
Penguin. 871 90 1000:103
Call (from Mr. Baker). 914 100 1000:109
Call (from Mr. Fox). 717 92 1000:129
Table 8.III.a shows that the leg-bones relatively to the weight of the
entire skeleton have really increased in weight; but Table 8.III.b shows
that according to the same standard the wing-bones have also really
decreased in weight; so that the relative disproportion 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 Tables 8.III.a and b, I may first state that I tested them
by taking another skeleton of a wild duck and of a 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 the second table we see,
with the exception of one case, a plain reduction 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 is consequent on
the remarkable lightness and thinness of all the bones of the skeleton.
Lastly, I weighed the furculum, 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 hundred 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 different 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
had probably been caused by gradual disuse. Hence, during the earlier
stages of the process of reduction, such birds would probably have
resembled our domesticated ducks in the state of their organs of flight.
This is the case with the water-hen (Gallinula nesiotis) of Tristan
d'Acunha, which "can flutter a little, but obviously uses its legs, and not
its wings, as a mode of escape." Now Mr. Sclater (8/19. 'Proc. Zoolog.
Soc.' 1861 page 261.) 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. chloropus).
On the other hand, the thigh-bones and pelvis are increased in length, the
former by four lines, relatively 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 presume no one will dispute that they have resulted from
the lessened use of the wings and the increased use of the legs.
THE GOOSE.
This bird deserves some notice, as hardly any other anciently domesticated
bird or quadruped has varied so little. That geese were anciently
domesticated we know from certain verses in Homer; and from these birds
having been kept (388 B.C.) in the Capitol at Rome as sacred to Juno, which
sacredness implies great antiquity. (8/20. 'Ceylon' by Sir J.E. Tennent
1859 volume 1 page 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 page 132. The goose figured on
the Egyptian monuments seems to have been the Red goose of Egypt.) 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. (8/21. Macgillivray's 'British Birds' volume 4 page 593.) A large
majority of capable judges are convinced that our geese are descended from
the wild Grey-leg goose (Anser ferus); the young of which can easily be
tamed. (8/22. Mr. A. Strickland, 'Annals and Mag. of Nat. Hist.' 3rd series
volume 3 1859 page 122, reared some young wild geese, and found them in
habits and in all characters identical with the domestic goose.) This
species, when crossed with the domestic goose, produced in the Zoological
Gardens, as I was assured in 1849, perfectly fertile offspring. (8/23. See
also Hunter 'Essays' edited by Owen volume 2 page 322.) Yarrell (8/24.
Yarrell's 'British Birds' volume 3 page 142.) has observed that the lower
part of the trachea of the domestic goose is sometimes flattened, and that
a ring of white feathers sometimes surrounds the base of the beak. These
characters seem at first sight 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 little flexible in its organisation under long-
continued domestication, the amount of variation which it has undergone may
be worth giving. It has increased in size and in productiveness (8/25. L.
Lloyd 'Scandinavian Adventures' 1854 volume 2 page 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.); and varies from white to a dusky colour.
Several observers (8/26. The Rev. L. Jenyns (Blomefield) seems first to
have made this observation in his 'British Animals.' See also Yarrell, and
Dixon in his 'Ornamental Poultry' (page 139), and 'Gardener's Chronicle'
1857 page 45.) 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 almost snow-white male
of the Rock goose (Bernicla antarctica) standing on the sea-shore by his
dusky partner is a sight well known to those who have traversed the sounds
of Tierra del Fuego and the Falkland Islands. Some geese have top-knots;
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. (8/27. Mr. Bartlet exhibited the head and neck of a bird thus
characterised before the Zoological Soc. February 1860.) 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. (8/28. W. Thompson
'Natural Hist. of Ireland' 1851 volume 3 page 31. The Rev. E.S. Dixon gave
me some information on the varying colour of the beak and legs.) This
latter fact deserves attention, because the colour of the legs and beak is
highly serviceable in discriminating the several closely allied wild forms.
(8/29. Mr. A. Strickland in 'Annals and Mag. of Nat. Hist.' 3rd series
volume 3 1859 page 122.) At our Shows two breeds are exhibited; viz., the
Embden and Toulouse; but they differ in nothing except colour. (8/30.
'Poultry Chronicle' volume 1 1854 page 498; volume 3 page 210.) Recently a
smaller and singular variety has been imported from Sebastopol (8/31. 'The
Cottage Gardener' September 4, 1860 page 348.), with the scapular feathers
(as I hear from Mr. Tegetmeier, who sent me specimens) greatly elongated,
curled, and even spirally 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
structure of the feathers is transmitted to half-bred birds. In Gallus
sonneratii the barbs and barbules blend together, 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 somewhat from any known wild
species, yet the amount of variation which it has undergone, as compared
with that of 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
languages 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 (8/32. 'L'Hist. de la Nature des Oiseaux' par
P. Belon 1555 page 156. With respect to the livers of white geese being
preferred by the Romans see Isid. Geoffroy St.-Hilaire 'Hist. Nat. Gen.'
tome 3 page 58.) 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 managers 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 domestication, 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 believes will
hereafter be found wild in some country, but not in India, where it is
certainly unknown. The males of these japanned birds differ conspicuously
from the common peacock in the colour of their secondary wing-feathers,
scapulars, wing-coverts, and thighs, and are I think more beautiful; they
are rather smaller than the common sort, and are always beaten by them in
their battles, as I hear from the Hon. A.S.G. Canning. The females are much
paler coloured than those of the common kind. Both sexes, as Mr. Canning
informs me, are white when they leave the egg, and they differ from the
young of the white variety only in having a peculiar pinkish tinge on their
wings. These japanned birds, though appearing suddenly in flocks of the
common kind, propagate their kind quite truly. Although they do not
resemble the hybrids which have been raised between P. cristatus and
muticus, nevertheless 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. (8/33. Mr. Sclater
on the black-shouldered peacock of Latham 'Proc. Zoolog. Soc.' April 24,
1860. Mr. Swinhoe at one time believed, 'Ibis' July 1868, that this kind of
peafowl was found wild in Cochin China, but he has since informed me that
he feels very doubtful on this head.)
On the other hand, Sir H. Heron states (8/34. 'Proc. Zoolog. Soc.' April
14, 1835.) that this breed suddenly appeared within his memory in Lord
Brownlow's large stock of pied, white, and common peacocks. 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, though a
smaller and weaker bird, increased, "to the extinction of the previously
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
peacocks 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. Mr. Jenner Weir informs me
that a peacock at Blackheath whilst young was white, but as it became older
gradually assumed the characters of the black-shouldered variety; both its
parents were common peacocks. Lastly, Mr. Canning has given a case of a
female of this same variety appearing in Ireland in a flock of the ordinary
kind. (8/35. 'The Field' May 6, 1871. I am much indebted to Mr. Canning for
information with respect to his birds.) Here, then, we have seven well
authenticated cases in Great Britain of japanned birds, having suddenly
appeared within recent times in flocks of the common peafowl. This variety
must also have formerly appeared in Europe, for Mr. Canning has seen an old
picture, and another is referred to in the 'Field,' with this variety
represented. These facts seem to me to indicate that the japanned peacock
is a strongly marked variety or "sport," which tends at all times and in
many places to reappear. This view is supported by the young being at first
white like the young of the white breed, which is undoubtedly a variation.
If, on the other hand, we believe the japanned peacock to be a distinct
species, we must suppose that in all the above cases the common breed had
at some former period been crossed by it, but had lost every trace of the
cross; yet that the offspring of these birds suddenly and completely
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 improbability 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 seven
instances in the same country, within no great length of time, to a wolf
perfect in every character; and we must further suppose 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 bird as the P. nigripennis, when first imported, would have
realised a large price; it is therefore improbable that it should have been
silently introduced and its history subsequently lost. On the whole the
evidence seems to me, as it did to Sir R. Heron, to be decisive in favour
of the japanned or black-shouldered breed being a variation, induced by
some unknown cause. On this view, the case is the most remarkable one 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 experienced of living
ornithologists.
THE TURKEY.
It seems fairly well established by Mr. Gould (8/36. 'Proc. Zoolog. Soc.'
April 8, 1856 page 61. Prof. Baird believes (as quoted in Tegetmeier
'Poultry Book' 1866 page 269) that our turkeys are descended 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.), that the turkey, in accordance with the history of its
first introduction, is descended from a wild Mexican form, which had been
domesticated by the natives before the discovery of America, and which is
now generally ranked as a local race, and not as a distinct species.
However this may be, the case deserves notice because in the 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." (8/37. Audubon 'Ornithological Biography' volume 1 1831 pages 4-
13; and 'Naturalist's Library' volume 14 'Birds' page 138.) Several
accounts have likewise been published 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 wild race or species. F. Michaux (8/38.
F. Michaux 'Travels in N. America' 1802 English translation page 217.)
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, occasionally with white
patches about the head. The other breeds scarcely differ except in colour,
and their chickens are generally mottled all over with brownish-grey.
(8/39. 'Ornamental Poetry' by the Rev. E.S. Dixon 1848 page 34.) The
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. (8/40. Bechstein 'Naturgesch. Deutschlands' b. 3 1793 s. 309.) Albin
in 1738, and Temminck within a much later period, describe a beautiful
breed, dusky-yellowish, brown above and white beneath, with a large top-
knot of soft plumose feather. The spurs of the male were rudimentary. This
breed has been for a long time extinct in Europe; but a living specimen has
lately been imported from the east coast of Africa, which still retains the
top-knot and the same general colouring and rudimentary spurs. (8/41. Mr.
Bartlett in 'Land and Water' October 31, 1868 page 233; and Mr. Tegetmeier
in the 'Field' July 17, 1869 page 46.) Mr. Wilmot has described (8/42.
'Gardener's Chronicle' 1852 page 699.) a white turkey-cock having 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 inherited this
kind of crest, but afterwards it 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 top-knot 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 differing
slightly from each other, present an analogous case with the wild cattle
kept in the several British parks. We must suppose that such differences
have resulted 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. In India the climate has apparently wrought a
still greater change in the turkey, for it is described by Mr. Blyth (8/43.
E. Blyth 'Annals and Mag. of Nat. Hist.' 1847 volume 20 page 391.) 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 some naturalists 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. (8/44. Roulin makes this remark
in 'Mem. de divers Savans, 1'Acad. des Sciences' tome 6 1835 page 349. Mr.
Hill of Spanish Town in a letter to me describes five varieties of the
Guinea fowl in Jamaica. I have seen singular pale-coloured varieties
imported from Barbadoes and Demerara.) The Guinea fowl has become
thoroughly feral in Jamaica and in St. Domingo (8/45. For St. Domingo see
M. A. Salle, in 'Proc. Zoolog. Soc.' 1857 page 236. Mr. Hill remarks to me,
in his letter, on the colour of the legs of the feral birds in Jamaica.),
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 notice. It has been crossed with nine or
ten other species of Fringillidae, and some of the hybrids are almost
completely fertile; but we have no evidence that any distinct breed has
originated from such crosses. Notwithstanding the modern domestication of
the canary, many varieties have been produced; even before the year 1718 a
list of twenty-seven varieties was published in France (8/46. Mr. B.P.
Brent 'The Canary, British Finches' etc. pages 21, 30.), and in 1779 a long
schedule of the desired qualities was printed by the London Canary Society,
so that methodical selection has been practised during a considerable
period. The greater number of the varieties differ only in colour and in
the markings of their plumage. Some breeds however, differ in shape, such
as the hooped or bowed canaries, and the Belgian canaries with their much
elongated bodies. Mr. Brent (8/47. 'Cottage Gardener' December 11, 1855
page 184: an account is here given of all the varieties. For many
measurements of the wild birds, see Mr. E. Vernon Harcourt ibid December
25, 1855 page 223.) 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 top-knotted canaries, and it is a singular fact that, if two top-
knotted birds are matched, the young, instead of having very fine top-
knots, are generally bald, or even have a wound on their heads. (8/48.
Bechstein 'Naturgesch. der Stubenvogel' 1840 s. 243; see s. 252 on the
inherited song of Canary-birds. With respect to their baldness see also W.
Kidd 'Treatise on Song-Birds.') It would appear as if the top-knot were due
to some morbid condition, 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 moulted, the peculiarity ceases." (8/49. W. Kidd
'Treatise on Song-Birds' page 18.) Canaries differ much in disposition and
character, and in some small degree in song. They produce eggs three or
four times during the year.
GOLD-FISH.
Besides mammals and birds, only a 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 they have been kept in confinement from an ancient
period in China. Mr. Blyth (8/50. The 'Indian Field' 1858 page 255.)
suspects, from the analogous variation of other fishes, that golden-
coloured fish do not occur in a state of nature. These fishes frequently
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 described and given coloured drawings of no less than eighty-nine
varieties. (8/51. Yarrell 'British Fishes' volume 1 page 319.) Many of the
varieties, however, such as triple tail-fins, etc., ought to be called
monstrosities; 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" (8/52.
Mr. Blyth in the 'Indian Field' 1858 page 255.), it might have been
predicted that selection would have been largely practised in the formation
of new breeds; and this is the case. In an old Chinese work it is said that
fish with vermilion scales were first raised in confinement during the Sung
dynasty (which commenced A.D. 960), "and now they are cultivated in
families everywhere for the sake of ornament." In another and more ancient
work, it is said that "there is not a household where the gold-fish is not
cultivated, in RIVALRY as to its colour, and as a source of profit," etc.
(8/53. W.F. Mayers 'Chinese Notes and Queries' August 1868 page 123.)
Although many breeds exist, it is a singular fact that the variations are
often not inherited. Sir R. Heron (8/54. 'Proc. Zoolog. Soc.' May 25,
1842.) kept many of these fishes, and placed all the deformed ones, namely,
those destitute of dorsal fins and those furnished with a double anal fin,
or triple tail, in a pond by themselves; but they did "not produce a
greater proportion of deformed offspring than the perfect fishes."
Passing over an almost infinite diversity of colour, we meet with the most
extraordinary modifications of structure. 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 latter deviation
of structure seems generally to occur "at the expense of the whole or part
of some other fin (8/55. Yarrell 'British Fishes' volume 1 page 319.); but
Bory de Saint-Vincent (8/56. 'Dict. Class. d'Hist. Nat.' tome 5 page 276.)
saw at Madrid gold-fish furnished with a dorsal fin and a triple tail. One
variety is characterised by a hump on its back near the head; and the Rev.
L. Jenyns (Blomefield) (8/57. 'Observations in Nat. Hist.' 1846 page 211.
Dr. Gray has described in 'Annals and Mag. of Nat. Hist.' 1860 page 151 a
nearly similar variety but destitute of a dorsal fin.) 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 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 (8/58. 'De l'Espece' 1859
page 459. With respect to the bees of Burgundy see M. Gerard, art. 'Espece'
in 'Dict. Univers. d'Hist. Nat.') 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 (8/59. See a
discussion on this subject, in answer to a question of mine, in 'Journal of
Horticulture' 1862 pages 225-242; also Mr. Bevan Fox in ditto 1862 page
284) 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. Woodbury states (8/60. This excellent observer may be
implicitly trusted; see 'Journal of Horticulture' July 14, 1863 page 39.)
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
(8/61. 'Journal of Horticulture' September 9, 1862 page 463; see also Herr
Kleine on same subject November 11 page 643, who sums up, that, though
there is some variability in colour, no constant or perceptible differences
can be detected in the bees of Germany.), 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 swarming and honey-gathering bees, this is a habit
which has 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 Luneburg
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 difference.
This remarkable uniformity in the hive-bee, wherever kept, may probably be
accounted for by the great difficulty, 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.
(8/62. Mr. Woodbury has published several such accounts in 'Journal of
Horticulture' 1861 and 1862.) 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 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. Gerstacker (8/63. 'Annals and Mag. of Nat. Hist.' 3rd series
volume 11 page 339.) but not by other highly competent judges, as
geographical races; he grounds his conclusion in chief part on the fact
that in certain districts, as in the Crimea and Rhodes, they vary 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 (8/64. 'The Cottage Gardener' May 1860
page 110; and ditto in 'Journal of Hort.' 1862 page 242.) procured some
bees from a cottager a few miles from Edinburgh, and perceived that they
differed from the common bee in the hairs on the head and thorax being
lighter 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 selection 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 an early period 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 structure
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 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 silkworms. Several distinct species 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 (8/65. 'Transact. Entomolog. Soc.' 3rd series volume 3 pages 143-173
and pages 295-331.) 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 opinion of
several capable judges who have particularly attended 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.
(8/66. Godron 'De l'Espece' 1859 tome 1 page 460. The antiquity of the
silkworm in China is given on the authority of Stanislas Julien.)
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. (8/67. See the remarks of Prof. Westwood, Gen.
Hearsey and others at the meeting of the Entomolog. Soc. of London July
1861.) Disuse has apparently aided in checking the development of the
wings. But the most important element in the production 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 cocoons and moths for
breeding is notorious (8/68. See for instance M. A. de Quatrefages 'Etudes
sur les Maladies actuelles du Ver a Soie' 1859 page 101.), and the
production 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 careful 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. (8/69. My
authorities for the statements will be given in the chapter on Selection.)
[The following details on the differences between the several breeds are
taken, when not stated to the contrary, from M. Robinet's excellent work
(8/70. 'Manuel de l'Educateur de Vers a Soie' 1848.), which bears every
sign of care and large experience. 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
gradually acquire the character of quick development, as in the Trevoltini
races. (8/71. Robinet ibid pages 12, 318. I may add that the eggs of N.
American silkworms taken to the Sandwich Islands produced moths at very
irregular periods; and the moths thus raised yielded 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 character would
ultimately have been acquired. See review in 'Athenaeum' 1844 page 329 of
J. Jarves 'Scenes in the Sandwich Islands.')
CATERPILLARS.
These vary greatly in size and colour. The skin is generally white,
sometimes mottled with black or grey, and occasionally quite black. The
colour, however, as M. Robinet asserts, is not constant, even in perfectly
pure breeds; except in the race tigree, so called from being marked with
transverse black stripes. As the general colour of the caterpillar is not
correlated with that of the silk (8/72. 'The Art of rearing Silkworms'
translated from Count Dandolo 1825 page 23.), 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 (pages 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 (8/73.
'Transact. Ent. Soc.' ut supra pages 153, 308.) 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 silkworms 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 "a 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 (8/74. Robinet ibid page 317.) 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
a trois mues, que nous avons experimentees, ont fait quatre mues a la
seconde ou a la troisieme annee, ce qui semble prouver qu'il a suffi de les
placer dans des conditions favorables pour leur rendre une faculte 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 different 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 constriction 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, but 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 arrivee a un tel etat de purete, qu'on ne voit pas un seul cocon jaune
dans des millions de cocons blancs." (8/75. Robinet ibid pages 306-317.)
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 received a similar statement from the eminent
naturalist, M. de Quatrefages. Captain Hutton also says (8/76. 'Transact.
Ent. Soc.' ut supra page 317.) 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 perpetuating any particular variation.
The males of the wild Bombycidae "fly swiftly in the day-time and evening,
but the females are usually very sluggish and inactive." (8/77. Stephen's
Illustrations, 'Haustellata' volume 2 page 35. See also Capt. Hutton
'Transact. Ent. Soc.' ibid page 152.) 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
perpetuated. In the silk-moth both sexes have imperfect, crumpled wings,
and are incapable of flight; but still there is a trace of the
characteristic difference in the two sexes; for though, on comparing a
number of males and females, I could detect no difference in the
development 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. Quatrefages
(8/78. 'Etudes sur les Maladies du Ver a Soie' 1859 pages 304, 209.) 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 la un veritable arret de
developpement partiel." On the other hand, he describes the female moths of
the Andre Jean breed as having "leurs ailes larges et etalees. Un seul
presente quelques courbures irregulieres et des plis anormaux." 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 (8/79. Quatrefages 'Etudes' etc. page 214.) but this
proceeds, according to Capt. Hutton (8/80. 'Transact. Ent. Soc.' ut supra
page 151.), 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 (8/81. 'Manuel de l'Educateur' etc. page 26.) of again crawling
up the trunk. Even this capacity sometimes fails, for M. Martins (8/82.
Godron 'De l'Espece' page 462.) placed some caterpillars on a tree, and
those which fell were not able to remount and perished of hunger; they were
even incapable of passing from leaf to leaf.
Some of the modifications which the silk-moth has undergone stand in
correlation with one another. Thus, the eggs of the moths which produce
white cocoons and of those which produce yellow cocoons differ slightly in
tint. The abdominal feet, also, of the caterpillars which yield white
cocoons are always white, whilst those which give yellow cocoons are
invariably yellow. (8/83. Quatrefages 'Etudes' etc. pages 12, 209, 214.) 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
(8/84. Robinet 'Manuel' etc. page 303.) that in France the caterpillars of
the races which produce 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. (8/85. Robinet ibid page
15.)]
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 a corresponding period. And finally we see that insects are
amenable to the great principle of Selection.
CHAPTER 1.IX.
CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS.
PRELIMINARY REMARKS ON THE NUMBER AND PARENTAGE OF CULTIVATED PLANTS.
FIRST STEPS IN CULTIVATION.
GEOGRAPHICAL DISTRIBUTION OF CULTIVATED PLANTS.
CEREALIA.
DOUBTS ON THE NUMBER OF SPECIES.
WHEAT: VARIETIES OF.
INDIVIDUAL VARIABILITY.
CHANGED HABITS.
SELECTION.
ANCIENT HISTORY OF THE VARIETIES.
MAIZE: GREAT VARIATION OF.
DIRECT ACTION OF CLIMATE ON.
CULINARY PLANTS.
CABBAGES: VARIETIES OF, IN FOLIAGE AND STEMS, BUT NOT IN OTHER PARTS.
PARENTAGE OF.
OTHER SPECIES OF BRASSICA.
PEAS: AMOUNT OF DIFFERENCE IN THE SEVERAL KINDS, CHIEFLY IN THE PODS AND
SEED.
SOME VARIETIES CONSTANT, SOME HIGHLY VARIABLE.
DO NOT INTERCROSS.
BEANS.
POTATOES: NUMEROUS VARIETIES OF.
DIFFERING LITTLE EXCEPT IN THE TUBERS.
CHARACTERS INHERITED.
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 unknown 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 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 monstrosities are of
little significance for our purpose. Many varieties are propagated solely
by grafts, buds, layers, bulbs, etc., and frequently it is not known how
far their peculiarities can be transmitted by seminal generation.
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 many characters in our cultivated plants have
become variable.
Before entering on details a few general remarks on the origin of
cultivated plants may be introduced. M. Alph. De Candolle (9/1. 'Geographie
botanique raisonnee' 1855 pages 810 to 991.) 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 cultivated plants. Of these he believes that
85 are almost certainly known in their wild state; but on this head other
competent judges (9/2. Review by Mr. Bentham in 'Hort. Journal' volume 9
1855 page 133 entitled 'Historical Notes on cultivated Plants' by Dr. A.
Targioni-Tozzetti. See also 'Edinburgh Review' 1866 page 510.) 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 include
flowers; and several of the more anciently cultivated flowers, such as
certain roses, the common Imperial lily, the tuberose, and even the lilac,
are said (9/3. 'Hist. Notes' as above by Targioni-Tozzetti.) 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 remote 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. The
difficulty would also be removed if they have been exterminated during the
progress of civilisation; but M. De Candolle has shown that this probably
has seldom occurred. As soon as a plant was 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 countries, where the whole face of the land has been
greatly changed, it can hardly be doubted that some plants have become
extinct; 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 (9/4. 'Considerations sur les Cereales' 1842
page 37. 'Geographie Bot.' 1855 page 930. "Plus on suppose l'agriculture
ancienne et remontant a une epoque d'ignorance, plus il est probable que
les cultivateurs avaient choisi des especes offrant a l'origine meme un
avantage incontestable.") and De Candolle have 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 authors 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 Australia cooking, during
a dearth, many vegetables in various 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 (9/5. Dr. Hooker has given me this information. See also his
'Himalayan Journals' 1854 volume 2 page 49.), 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 stomachs, so as to relieve the pangs of hunger. They looked
like walking skeletons, and suffered fearfully from constipation. Sir
Andrew Smith also informs me that on such occasions the natives observe as
a guide for themselves, 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 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 savages 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 compelled 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 (9/6. 'Travels in Central Africa' English translation
volume 1 pages 529 and 390; volume 2 pages 29, 265, 270. Livingstone
'Travels' page 551.) 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 meadow-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 seed 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. (9/7. For instance in both North
and South America. Mr. Edgeworth 'Journal Proc. Linn. Soc.' vol 6 Bot. 1862
page 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 grasses, namely,
of Agrostis, Panicum, Cenchrus, and Pennisetum, as well as the seeds of
four other genera belonging to distinct families.)
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, etc.,
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 Switzerland during the Stone-period largely collected
wild crabs, sloes, bullaces, hips of roses, elderberries, beechmast, and
other wild berries and fruit. (9/8. Prof. O. Heer 'Die Pflanzen der
Pfahlbauten' 1866 aus dem Neujahr. Naturforsch. Geselschaft' 1866; and Dr.
H. Christ in Rutimeyer's 'Die Fauna der Pfahlbauten' 1861 s. 226.) 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 (9/9. 'Travels' page
535. Du Chaillu 'Adventures in Equatorial Africa' 1861 page 445.) 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
forethought, would be to sow the seeds of useful plants; and as the soil
near the hovels of the natives (9/10. In Tierra del Fuego the spot where
wigwams had formerly stood could be distinguished at a great distance by
the bright green tint of the native vegetation.) 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 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. (9/11. 'American
Acad. of Arts and Sciences' April 10th, 1860 page 413. Downing 'The Fruits
of America' 1845 page 261.) 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 cultivation. 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 (9/12. 'Journals of Expeditions in
Australia' 1841 volume 2 page 292.) 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 killing water-fowl whilst very
young will be followed by "much rain, snow, blow much." (9/13. Darwin
'Journal of Researches' 1845 page 215.) I may add, as showing 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 (9/14. De Candolle has tabulated the facts in
the most interesting manner in his 'Geographie Bot.' page 986.) 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 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 be indeed a surprising 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 (9/15. 'Flora of Australia' Introduction page 110.)
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 cultivated several plants; but all inquirers
believe, in accordance with the traditions of the natives, that the early
Polynesian colonists brought with them seeds and roots, as well as the dog,
which had been wisely preserved during their long voyage. The Polynesians
are so frequently 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 colonists, would not have had any strong inducement
to cultivate 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 civilised state of the inhabitants, as shown by the fact of
their having practised artificial irrigation and made tunnels through hard
rocks without the use of iron or gunpowder, and who, as we shall see in a
future chapter, fully recognised, as far as animals were concerned, and
therefore probably in the case of plants, the important principle 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 (9/16. 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 references Asa Gray in the 'American Journal of Science' volume 24
November 1857 page 441. For the traditions of the natives of New Zealand
see Crawfurd 'Grammar and Dict. of the Malay Language' 1852 page 260.) 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 civilised 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.' (9/17. See for example Mr. Hewett C. Watson's
remarks on our wild plums and cherries and crabs: 'Cybele Britannica'
volume 1 pages 330, 334, etc. Van Mons (in his 'Arbres Fruitiers' 1835 tome
1 page 444) declares that he has found the types of all our cultivated
varieties in wild seedlings, but then he looks on these seedlings as so
many aboriginal stocks.)
[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 (9/18. See A. De Candolle 'Geograph. Bot.' 1855 page 928 et
seq. Godron 'De l'Espece' 1859 tome 2 page 70; and Metzger 'Die
Getreidearten' etc. 1841.) 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 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 (9/19. Mr. Bentham in his
review entitled 'Hist. Notes on cultivated Plants' by Dr. A. Targioni-
Tozzetti in 'Journal of Hort. Soc.' volume 9 1855 page 133. He informs me
that he still retains the same opinion.), "We ourselves have no hesitation
in stating our conviction, 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 (9/20. 'Geograph. Bot.' page 928. The whole subject is discussed
with admirable fulness and knowledge.) has adduced abundant evidence that
common wheat (Triticum vulgare) has been found wild in various parts of
Asia, where it is not likely to have escaped from cultivation: and there is
some force in M. Godron's remark, that, supposing these plants to be
escaped seedlings (9/21. Godron 'De l'Espece' tome 2 page 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 Aegilops; but M.
Godron (tome 1 page 165) has shown by careful experiments that the first
step in the series, viz. Aegilops triticoides, is a hybrid between wheat
and Ae. ovata. The frequency with which these hybrids spontaneously arise,
and the gradual manner in which the Ae. triticoides becomes converted into
true wheat, alone leave any doubt with respect to M. Godron's
conclusions.), as 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 character.
But the strong tendency to inheritance, which most of the varieties of
wheat evince, as we shall presently see, is here greatly undervalued. Much
weight must also be attributed to a remark by Professor Hildebrand (9/22.
'Die Verbreitungsmittel der Pflanzen' 1873 page 129.) that when the seeds
or fruit of cultivated plants possess qualities disadvantageous to them as
a means of distribution, we may feel almost sure that they no longer retain
their aboriginal condition. On the other hand, M. De Candolle insists
strongly on the frequent occurrence in the Austrian dominions of rye and of
one kind of oats in an apparently 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 cereals. With
respect to oats, according to Mr. Buckmann (9/23. Report to British
Association for 1857 page 207.), the wild English Avena fatua can be
converted by a few years of careful cultivation and selection into forms
almost identical with two very distinct 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 the more distant
parts of the world; for Loiseleur-Deslongchamps (9/24. Considerations sur
les Cereales' 1842-43 page 29.) speaks of three new species or varieties,
sent to Europe in 1822 from Chinese Mongolia, which he considers as being
there indigenous. Moorcroft (9/25. 'Travels in the Himalayan Provinces'
etc. 1841 volume 1 page 224.) 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
important 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 have arisen so strongly marked, that they
have been considered by capable judges as specifically 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 (9/26. Col. J. Le Couteur on the 'Varieties of
Wheat' pages 23, 79.); 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 (9/27. Loiseleur-Deslongchamps 'Consid.
sur les Cereales' page 11.) differ in colour and in shape, being
quadrangular, 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 Gramineae serves even as a generic character
(9/28. See an excellent review in Hooker 'Journ. of Botany' volume 8 page
82 note.); although, as remarked by Godron (9/29. 'De l'Espece' tome 2 page
73.) 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 colour; 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 (9/30. Ibid tome 2
page 75.) has remarked, in an exactly parallel manner,--in the seed being
downy or glabrous, and in colour,--and in the florets being barbed or not
barbed, etc. 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.
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.
(9/31. For Dalbret and Philippar see Loiseleur-Deslongchamps 'Consid. sur
les Cereales' pages 45, 70. Le Couteur on Wheat pages 6, 14-17.) 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, he found that there was only one "secure
mode to ensure the growth of pure sorts, namely, to grow them from single
grains or from single ears, and to follow up the plan by afterwards sowing
only the produce of the most productive so as to form a stock." But Major
Hallett (9/32. See his Essay on 'Pedigree in Wheat' 1862; also paper read
before the British Association 1869 and other publications.) has gone much
farther, and by the continued selection of plants from the grains of the
same ear, during successive generations, has made his 'Pedigree in Wheat'
(and other cereals) now famous in many quarters of the world. 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 (9/33.
'Varieties of Wheat' Introduction page 6. Marshall in his 'Rural Economy of
Yorkshire' volume 2 page 9 remarks that "in every field of corn there is as
much variety as in a herd of cattle.") 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 Professor 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. Shirreff has had the good fortune to
raise in his lifetime seven new varieties, which are now extensively grown
in many parts of Britain. (9/34. 'Gardener's Chronicle' and 'Agricult.
Gazette' 1862 page 963.)
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. On the other hand Major
Hallett (9/35. 'Gardener's Chronicle' November 1868 page 1199.) has shown
how wonderfully constant some varieties are, although not ancient ones, and
although cultivated in various countries. With respect to the tendency to
vary, Metzger (9/36. 'Getreidearten' 1841 s. 66, 91, 92, 116, 117.) 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 summers; 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 variability, and they further show that a variety may become
habituated to new conditions. One is at first inclined to conclude with
Loiseleur-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 Linnaeus as distinct species; but M. Monnier (9/37. Quoted by
Godron 'De l'Espece' volume 2 page 74. So it is, according to Metzger
'Getreidearten' s. 18, with summer and winter barley.) has proved that the
difference between them is only temporary. 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 converted 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
(9/38. Loiseleur-Deslongchamps 'Cereales' part 2 page 224. Le Couteur page
70. Many other accounts could be added.) differently from our European
varieties. In Canada the first settlers, according to Kalm (9/39. 'Travels
in North America' 1753-1761 English translation volume 3 page 165.), found
their winters too severe for winter-wheat brought from France, and their
summers often too short for summer-wheat; and they thought that their
country was useless for corn crops until they procured summer-wheat from
the northern parts of Europe, which succeeded well. It is notorious that
the proportion of gluten differs much under different climates. The weight
of the grain is also quickly affected by climate: Loiseleur-Deslongchamps
(9/40. 'Cereales' part 2 pages 179-183.) 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 (9/41. 'On the
Varieties of Wheat' Introduction page 7. See Marshall 'Rural Econ. of
Yorkshire' volume 2 page 9. With respect to similar cases of adaptation in
the varieties of oats see some interesting papers in the 'Gardener's
Chronicle and Agricult. Gazette' 1850 pages 204, 219.) 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 habituated to its conditions of life, as Metzger
has shown certainly occurs, but it is probably in main part due to innate
differences between the several varieties.
Much has been written on the deterioration of wheat; that the quality of
the flour, size of grain, time of flowering, and hardness, may be modified
by climate and soil, seems nearly certain; but that the whole body of any
one sub-variety ever becomes changed into another and distinct sub-variety,
there is no reason to believe. What apparently does take place, according
to Le Couteur (9/42. 'On the Varieties of Wheat' page 59. Mr. Shirreff and
a higher authority cannot be given ('Gardener's Chronicle and Agricult.
Gazette' 1862 page 963), says "I have never seen grain which has either
been improved or degenerated by cultivation, so as to convey the change to
the succeeding crop.), 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 supplants the variety which was first sown.
With respect to the natural crossing of distinct varieties the evidence is
conflicting, but preponderates against its frequent occurrence. Many
authors maintain that impregnation takes place in the closed flower, but I
am sure from my own observation 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 varieties of wheat have
arisen; but their differences are unimportant, 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. (9/43. Alph. De Candolle 'Geograph. Bot.' page 930. )
It is an important fact, which we have recently learnt from the admirable
researches of Heer (9/44. 'Pflanzen der Pfahlbauten' 1866.), that the
inhabitants of Switzerland, even so early as the Neolithic period,
cultivated 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 remarked, agriculture even at the
Neolithic period, had already made considerable progress; for, besides the
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-inhabitants either still kept up commercial
intercourse with some southern people or had originally proceeded as
colonists from the South.
Loiseleur-Deslongchamps (9/45. 'Les Cereales' page 94.) has argued that, if
our cereal plants have 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
overlooked. 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 variations, which can be selected and separately
propagated; and that occasionally more strongly pronounced variations
appear, which, as Mr. Shirreff has proved, are well worthy of extensive
cultivation. Not until equal attention be paid to the variability and
selection of weeds, can the argument from their constancy under
unintentional culture be of any value. In accordance with the principles of
selection we can understand how it is that in the several cultivated
varieties of wheat the organs of vegetation 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 (9/46. Quoted by Le Couteur
page 16.) 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 laborious the work has been found by Le Coutour and
Hallett. Although the principle of selection is so important, yet the
little which man has effected, by incessant efforts (9/47. A. De Candolle
'Geograph. Bot.' page 932.) 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 contained a
sufficient supply of the necessary chemical elements.
We now know that man was sufficiently civilised to cultivate the ground at
an immensely remote period; so that wheat might have been improved long ago
up to 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 improvement of our cereals. In the most ancient
lake-habitations of Switzerland, when men employed only flint-tools, the
most extensively cultivated wheat was a peculiar kind, with remarkably
small ears and grains. (9/48. O. Heer 'Die Pflanzen der Pfahlbauten' 1866.
The following passage is quoted from Dr. Christ in 'Die Fauna der
Pfahlbauten, von Dr. Rutimeyer' 1861 s. 225.) "Whilst the grains of the
modern forms are in section from seven to eight millimetres 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 1/2 lines long, and
scarcely 1 1/2 broad, whilst those now grown have a length of three lines,
and almost the same in breadth." (9/49. Heer as quoted by Carl Vogt
'Lectures on Man' English translation page 355.) 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 appearance and final
disappearance of the several plants which were cultivated in greater or
less abundance in Switzerland during former successive periods, and which
generally differed more or less from our existing varieties. The peculiar
small-eared and small-grained wheat, already 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 exactly 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. monococcum) is known to have existed during the
Stone period only by the presence of a single ear. A sixth kind, the common
T. spelta, was not introduced into Switzerland until the Bronze age. Of
barley, besides the short-eared and small-grained kind, two others were
cultivated, 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 somewhat smaller than those produced by our existing
varieties. 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 became 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 descriptions given by palaeontologists of the
first appearance, the increasing rarity, and final extinction or
modification of fossil species, embedded in the successive 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 from between four and eight species,
which may have either closely resembled our present cultivated forms, or
have been so widely different 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 OR INDIAN CORN: Zea mays.
Botanists are nearly unanimous that all the cultivated kinds belong to the
same species. It is undoubtedly (9/50. See Alph. De Candolle's long
discussion in his 'Geograph. Bot.' page 942. With respect to New England
see Silliman's 'American Journal' volume 44 page 99.) 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
(9/51. 'Travels in Peru' English translation page 177.) describes two
kinds, now extinct or not known in Peru, which were taken from tombs
apparently prior to the dynasty of the Incas. 'But there is even stronger
evidence of antiquity, for I found on the coast of Peru (9/52. 'Geolog.
Observ. on S. America' 1846 page 49.) 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 (9/53. This maize is figured in Bonafous' magnificent work,
'Hist. Nat. du Mais' 1836 P1. v. bis, and in the 'Journal of Hort. Soc.'
volume 1 1846 page 115 where an account is given of the result of sowing
the seed. A young Guarany Indian, on seeing this kind of maize, told
Auguste St. Hilaire (see De Candolle 'Geograph. Bot.' page 951) that it
grew wild in the humid forests of his native land. Mr. Teschemacher in
'Proc. Boston Soc. Hist.' October 19, 1842 gives an account of sowing the
seed.), in which the grains, instead of being naked, are concealed by husks
as much as eleven lines in length, has been stated, but on insufficient
evidence, to grow wild in Brazil. It is almost certain that the aboriginal
form would have had its grains thus protected (9/54. Moquin-Tandon
'Elements de Teratologie' 1841 page 126.); 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 cultivated, should vary so quickly and in
so great a degree.
Maize has varied in an extraordinary and conspicuous manner. Metzger (9/55.
'Die Getreidearten' 1841 s. 208. I have modified a few of Metzger's
statements in accordance with those made by Bonafous in his great work
'Hist. Nat. du Mais' 1836.), 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 are coloured--white, pale-yellow, orange,
red, violet, or elegantly streaked with black (9/56. Godron 'De l'Espece'
tome 2 page 80; Al. De Candolle ibid page 951.); 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, and which is extensively
cultivated in the United States as sweet corn) 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
mais a 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. (9/57. 'Transact. Bot.
Soc. of Edinburgh' volume 8 page 60.) Azara describes (9/58. 'Voyages dans
l'Amerique Meridionale' tome 1 page 147.) 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. (9/59. Bonafous 'Hist. Nat. du Mais' page 31.) Some of the
foregoing differences would certainly be considered of specific value with
plants in a state of nature.
Le Comte Re states that the grains of all the varieties which he cultivated
ultimately assumed a yellow colour. But Bonafous (9/60. Ibid page 31.)
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. (9/61. Metzger 'Getreidearten' s. 206.) Peter Kalm (9/62.
'Description of Maize' by P. Kalm 1752 in 'Swedish Acts' volume 4. I have
consulted an old English MS. translation.), who particularly 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 deg
in Virginia, and sown in lat. 43-44 deg 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 deg in Canada. By
taking great care at first, the southern kinds after some years' culture
ripen their seed perfectly in their northern homes, so that this is an
analogous 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 seeds 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 (9/63. 'Getreidearten' s. 208.) in one case, namely, with a tall
kind (Breit-korniger mais, Zea altissima) brought from the warmer parts of
America. During the first year the plants were twelve feet high, and a 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
resemblance 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. Analogous 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. In the above cases the seeds were carried from
a warm to a colder climate. But Fritz Muller informs me that a dwarf
variety with small rounded seeds (papa-gaien-mais), introduced from Germany
into S. Brazil, produces plants as tall, with seeds as flat, as those of
the kind commonly cultivated there.]
These facts afford the most remarkable instance known to me of the direct
and prompt action of climate on a plant. It might have been expected that
the tallness of the stem, the period of 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,
however, flowers, with their product the seed, are formed by the
metamorphosis of the stem and leaves, any modification 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 Jersey, 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 (9/64. 'Cabbage Timber' 'Gardener's Chronicle' 1856 page 744,
quoted from Hooker 'Journal of Botany.' A walking-stick made from a
cabbage-stalk is exhibited in the Museum at Kew.) and as walking-sticks. We
are thus reminded that in certain countries plants belonging to the
generally herbaceous order of the Cruciferae 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 kails, which come nearest to the wild parent-form. There are
also various frizzled and laciniated kinds, some of such beautiful colours
that Vilmorin in his Catalogue of 1851 enumerates ten varieties which are
valued solely for ornament. 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
(9/65. 'Journal de la Soc. Imp. d'Horticulture' 1855 page 254 quoted from
'Gartenflora' April 1855.) of the 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 the
flower-stems in the broccoli and cauliflower, it is remarkable that the
flowers themselves, the seed-pods and seeds, present extremely slight
differences or none at all. (9/66. Godron 'De l'Espece' tome 2 page 52;
Metzger 'Syst. Beschreibung der Kult. Kohlarten' 1833 s. 6.) 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 Portsmouth
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 collection of the seeds of twenty-
eight different kinds, and most of them were undistinguishable; when there
was any difference it was excessively slight; thus, the seeds of various
broccolis and cauliflowers, 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
difference 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 neglected 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. (9/67. Regnier 'De l'Economie Publique des Celtes' 1818 page 438.)
It would be useless to give a classified description (9/68. See the elder
De Candolle in 'Transact. of Hort. Soc.' volume 5; and Metzger 'Kohlarten'
etc.) of the numerous races, sub-races, and varieties of the cabbage; but
it may be mentioned that Dr. Lindley has lately proposed (9/69. 'Gardener's
Chronicle' 1859 page 992.) a system founded on the state of development of
the terminal and lateral leaf-buds. Thus:
I. All the leaf-buds active and open, as in the wild-cabbage, kail, etc.
II. All the leaf-buds active, but forming heads, as in Brussel-sprouts,
etc.
III. Terminal leaf-bud alone active, forming a head as in common cabbages,
savoys, etc.
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 abortive and
succulent, as in the sprouting-broccoli. This latter variety is a new one,
and bears the same relation to common broccoli, as 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 sixteenth
century (9/70. Alph. De Candolle 'Geograph. Bot.' pages 842 and 989.), 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 proof of this: I
raised 233 seedlings from cabbages of different kinds, which had purposely
been planted near each other, and of the seedlings no less than 155 were
plainly deteriorated and mongrelised; 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 "Cottager's
Kail," has lately been produced by crossing common kail and Brussel-
sprouts, recrossed with purple broccoli (9/71. 'Gardener's Chronicle'
February 1858 page 128.), and is said to be true; but plants raised by me
were not nearly so constant in character as any common kind of 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 (9/72. 'Kohlarten' s. 22.) 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 countries, and the same thing has been observed with
an English variety grown during an extremely warm and damp autumn near
Paris. (9/73. Godron 'De l'Espece' tome 2 page 52; Metzger 'Kohlarten' s.
22.) 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 (9/74.
'Geograph. Bot.' page 840.) 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 all commingled together, of the various
cultivated kinds. In the same manner as we have often seen with
domesticated animals, the supposed multiple origin of the cabbage throws no
light on the characteristic differences between the cultivated forms. If
our cabbages are the descendants of three or four distinct species, every
trace of any sterility which may originally have existed between them is
now lost, for none of the varieties can be kept distinct without scrupulous
care to prevent intercrossing.
The other cultivated forms of the genus Brassica are descended, according
to the view adopted by Godron and Metzger (9/75. Godron 'De l'Espece' tome
2 page 54; Metzger 'Kohlarten' s. 10.), 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, namely, Swedish turnips (believed to be of hybrid origin)
(9/76. 'Gardener's Chronicle and Agricult. Gazette' 1856 page 729. See more
especially ibid 1868 page 275: the writer asserts that he planted a variety
of cabbage (B. oleracea) close to turnips (B. rapa) and raised from the
crossed seedlings true Swedish turnips. These latter plants ought,
therefore, to be classed with cabbages or turnips, and not under B. napus.)
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 Godron 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. (9/77.
'Gardener's Chronicle and Agricult. Gazette' 1855 page 730.) Metzger by
culture converted the biennial or winter rape into the annual or summer
rape,--varieties which have been thought by some authors to be specifically
distinct. (9/78. Metzger, 'Kohlarten' s. 51.)
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 bow quickly the roots of the wild
parsnip can be enlarged, as Vilmorin formerly proved in the case of the
carrot. (9/79. These experiments by Vilmorin have been quoted by many
writers. An eminent 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. Carriere has lately
stated ('Gardener's Chronicle' 1865 page 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.)
This latter plant, in its cultivated state, differs in scarcely any
character from the wild English carrot, except in general luxuriance and in
the size and quality of its roots; but ten varieties, differing in the
colour, shape, and quality of the root, are cultivated in England and come
true by seed. (9/80. Loudon's 'Encyclop. of Gardening' page 835.) 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 modifications have
been again and again selected, until at last a great amount of change has
been effected.
With respect to the radish, M. Carriere, by sowing the seed of the wild
Raphanus raphanistrum in rich soil, and by continued selection during
several generations, raised many varieties, closely like the cultivated
radish (R. sativus) in their roots, as well as the wonderful Chinese
variety, R. caudatus: (see 'Journal d'Agriculture pratique' tome 1 1869
page 159; also a separate essay 'Origine des Plantes Domestiques' 1869.)
Raphanus raphanistrum and sativus have often been ranked as distinct
species, and owing to differences in their fruit even as distinct genera;
but Professor Hoffman ('Bot. Zeitung' 1872 page 482) has now shown that
these differences, remarkable as they are, graduate away, the fruit of R.
caudatus being intermediate. By cultivating R. raphanistrum during several
generations (ibid 1873 page 9), Professor Hoffman also obtained plants
bearing fruits like those of R. sativus.
PEA (Pisum sativum).
Most botanists look at the garden-pea as specifically distinct from the
field-pea (P. arvense). The latter exists 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. (9/81. Alph. De Candolle
'Geograph. Bot.' 960. Mr. Bentham 'Hort. Journal' volume 9 1855 page 141
believes that garden and field peas belong to the same species, and in this
respect he differs from Dr. Targioni.) Andrew Knight crossed, as I am
informed 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. Alefield has recently studied (9/82. 'Botanische Zeitung' 1860
s. 204.) the genus with care, and, after having cultivated about fifty
varieties, concludes that certainly they all belong to the same species. It
is an interesting fact already alluded to, that, according to O. Heer
(9/83. 'Die Pflanzen der Pfahlbauten' 1866 s. 23.) 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. arvense or the
field-pea. The varieties of the common garden-pea are numerous, and differ
considerably from one another. For comparison I planted at the same time
forty-one, English and French varieties. They differed greatly in height,--
namely from between 6 and 12 inches to 8 feet (9/84. A variety called the
Rounciva attains this height, as is stated by Mr. Gordon in 'Transact.
Hort. Soc.' 2nd series volume 1 1835 page 374 from which paper I have taken
some facts.),--in manner of growth, and in period of maturity. Some 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:--HAIR'S
DWARF MONMOUTH has very large leaves, and the POIS NAIN HATIF, and the
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 GEANT 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, two, or several flowers in a small
cluster, are borne on the same peduncle; and this is a difference which is
considered of specific value in some of the Leguminosae. 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 the POIS NAIN HATIF; but
HAIR'S 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 constant
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, which, 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 GEANT is much pointed
at the extremity; and in the variety "A 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 remarkably 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.
(FIGURE 41. PODS AND PEAS. I. Queen of Dwarfs. II. American Dwarf. III.
Thurston's Reliance. IV. Pois Geant sans parchemin. a. Dan O'Rourke Pea. b.
Queen of Dwarfs Pea. c. Knight's Tall White Marrow. a. Lewis's Negro Pea.)
In the pea itself we have every tint between almost pure white, brown,
yellow, and intense green; in the varieties of the SUGAR PEAS we have these
same tints, together with red passing through fine purple into a dark
chocolate tint. These colours are either uniform or distributed in dots,
striae, 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 different 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
THE 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
distinct species.
Andrew Knight (9/85. 'Phil. Tract.' 1799 page 196.) 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
Canterbury, 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. (9/86. 'Gardener's
Magazine' volume 1 1826 page 153.) But the greater number of varieties have
a singularly short existence: thus Loudon remarks (9/87. 'Encyclopaedia of
Gardening' page 823.) 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. Masters 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 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
Leguminosae, 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 it could hardly fail to be left
on the stigma of the next flower which was visited. Nevertheless, distinct
varieties growing closely together rarely cross; and I have reason to
believe that this is due to their stigmas being prematurely fertilised in
this country by pollen from the same flower. The horticulturists who raise
seed-peas are thus enabled to plant distinct varieties close together
without any bad consequences; and it is certain, as I have myself found,
that true seed may be saved during at least several generations under these
circumstances. (9/88. See Dr. Anderson to the same effect in the 'Bath Soc.
Agricultural Papers' volume 4 page 87.) Mr. Fitch raised, as he informs me,
one variety for twenty years, and it always came true, though grown close
to other varieties. From the analogy of kidney-beans I should have expected
(9/89. I have published full details of experiments on this subject in the
'Gardener's Chronicle' 1857 October 25.) that varieties thus circumstanced
would have occasionally crossed; and I shall give in the eleventh chapter
two cases of this having occurred, as shown (in a manner hereafter to be
explained) by the pollen of the one variety having acted directly on the
seeds of the other. Whether many of the new varieties which incessantly
appear are due to such occasional and accidental crosses, I do not know.
Nor do I know whether the short existence of almost all the numerous
varieties is the result of mere change of fashion, or of their having a
weak constitution, from being the product of long-continued self-
fertilisation. It may, however, be noticed that several of Andrew Knight's
varieties, 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 vigorous in 1860; but now, in 1865, a writer, speaking
(9/90. 'Gardener's Chronicle' 1865 page 387.) 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. Alefield
has given (9/91. 'Bonplandia' 10, 1862 s. 348.) short diagnostic characters
of forty varieties. Everyone who has seen a collection must have been
struck with the great difference in shape, thickness, proportional length
and 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 (9/92. Heer 'Die
Pflanzen der Pfahlbauten' 1866 s. 22.) by a peculiar and now extinct
variety producing very small beans. (9/93. Mr. Bentham informs me that in
Poitou and the adjoining parts of France, varieties of Phaseolus vulgaris
are extremely numerous, and so different that they were described by Savi
as distinct species. Mr. Bentham believes that all are descended from an
unknown eastern species. Although the varieties differ so greatly in
stature and in their seeds, "there is a remarkable sameness in the
neglected characters of foliage and flowers, and especially in the
bracteoles, an insignificant character in the eyes even of botanists.")
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.
(9/94. Darwin 'Journal of Researches' 1845 page 285. Sabine in 'Transact.
Hort. Soc.' volume 5 page 249.) The varieties cultivated in Britain are
numerous; thus Lawson (9/95. 'Synopsis of the Vegetable Products of
Scotland' quoted in Wilson's 'British Farming' page 317.) gives a
description 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 a difference between the individuals of the same variety as between
the different varieties. The flower varied 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. (9/96. Sir G. Mackenzie, in 'Gardener's
Chronicle' 1845 page 790.) The fruit or berries also differ, but only in a
slight degree. (9/97. Putsche und Vertuch 'Versuch einer Monographie der
Kartoffeln' 1819 s. 9, 15. See also Dr. Anderson 'Recreations in
Agriculture' volume 4 page 325.) The varieties are liable in very different
degree to the attack of the Colorado potato-beetle. (9/98. Walsh 'The
American Entomologist' 1869 page 160. Also S. Tenney 'The American
Naturalist' May 1871 page 171.)
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 (9/99.
'Gardener's Chronicle' 1862 page 1052.) 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 or rhizomes is different; thus,
in the gurken-kartoffeln they form a pyramid with the apex downwards, 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, 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, etc., 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 (9/100. 'Bath Society Agricult. Papers'
volume 5 page 127. And 'Recreations in Agriculture' volume 5 page 86.)
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, produced extremely dissimilar tubers;
and some tubers which externally 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. Kidney
potatoes must be ranked amongst the most highly cultivated and artificial
races; nevertheless their peculiarities can often be strictly propagated by
seed. A great authority, Mr. Rivers (9/101. 'Gardener's Chronicle' 1863
page 643.) states that "seedlings 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."
CHAPTER 1.X.
PLANTS continued--FRUITS--ORNAMENTAL TREES--FLOWERS.
FRUITS.
GRAPES.
VARY IN ODD AND TRIFLING PARTICULARS.
MULBERRY.
THE ORANGE GROUP.
SINGULAR RESULTS FROM CROSSING.
PEACH AND NECTARINE.
BUD-VARIATION.
ANALOGOUS VARIATION.
RELATION TO THE ALMOND.
APRICOT.
PLUMS.
VARIATION IN THEIR STONES.
CHERRIES.
SINGULAR VARIETIES OF.
APPLE.
PEAR.
STRAWBERRY.
INTERBLENDING OF THE ORIGINAL FORMS.
GOOSEBERRY.
STEADY INCREASE IN SIZE OF THE FRUIT.
VARIETIES OF.
WALNUT.
NUT.
CUCURBITACEOUS PLANTS.
WONDERFUL VARIATION OF.
ORNAMENTAL TREES.
THEIR VARIATION IN DEGREE AND KIND.
ASH-TREE.
SCOTCH-FIR.
HAWTHORN.
FLOWERS.
MULTIPLE ORIGIN OF MANY KINDS.
VARIATION IN CONSTITUTIONAL PECULIARITIES.
KIND OF VARIATION.
ROSES.
SEVERAL SPECIES CULTIVATED.
PANSY.
DAHLIA.
HYACINTH.
HISTORY AND VARIATION OF.
[THE VINE (Vitis vinifera) (Grapes).
The best authorities consider all our grapes as the descendants of one
species which now grows wild in western Asia, which grew wild during the
Bronze age in Italy (10/1. Heer 'Pflanzen der Pfahlbauten' 1866 s. 28.),
and which has recently been found fossil in a tufaceous deposit in the
south of France. (10/2. Alph. De Candolle 'Geograph. Bot.' page 872; Dr. A.
Targioni-Tozzetti in 'Jour. Hort. Soc.' vol 9 page 133. For the fossil vine
found by Dr. G. Planchon see 'Nat. Hist. Review' 1865 April page 224. See
also the valuable works of M. de Saporta on the 'Tertiary Plants 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 described by Clemente (10/3.
Godron 'De l'Espece' tome 2 page 100.) in a forest in Spain; but as the
grape sows itself freely in Southern Europe, and as several of the chief
kinds transmit their characters by seed (10/4. See an account of M.
Vibert's experiments by Alex. Jordan in 'Mem. de l'Acad. de Lyon' tome 2
1852 page 108.), 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 propagated by seed, we may infer from the largely
increased number of varieties since the earlier historical records. New
hot-house varieties are produced almost every year; for instance (10/5.
'Gardener's Chronicle' 1864 page 488.) a golden-coloured variety has been
recently raised in England from a black grape without the aid of a cross.
Van Mons (10/6. 'Arbres Fruitiers' 1836 tome 2 page 290.) reared a
multitude of varieties 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 fruits 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 Horticultural Gardens of London,
published in 1842, 99 varieties are enumerated. Wherever the grape is grown
many varieties occur: Pallas 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
peculiarities, all taken from Odart's highly esteemed work (10/7. Odart
'Ampelographie Universelle' 1849.) 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 (page 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 page 397) presents a
peculiarity by which it can be at once recognised amongst a host of other
varieties, namely, 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 (page 426), amongst others, "by some of the
leaves, and it is always the lowest on the branches, suddenly becoming 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
(page 71), which often produces small round, and large oblong, berries in
the same bunch. Certain grapes called Nebbiolo (page 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
(page 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 (page 243) is valued for well sustaining
prolonged humidity. This latter variety sprouts late in the spring, yet
matures its fruit early; other varieties (page 362) have the fault of being
too much excited by the April sun, and in consequence suffer from frost. A
Styrian variety (page 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 characters 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 old groups of varieties (10/8. M. Bouchardat in 'Comptes Rendus'
December 1, 1851 quoted in 'Gardener's Chronicle' 1852 page 435. See also
C.V. Riley on the manner in which some few of the varieties of the American
Labruscan Vine escape the attacks of the Phylloxera: 'Fourth Annual Report
on the Insects of Missouri' 1872 page 63 and 'Fifth Report' 1873 page 66.)
have suffered far more from mildew than others. Thus "the group of
Chasselas, so rich in varieties, did not afford a single 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 American 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 silkworm, in a manner not observed with other plants; but
this has arisen simply from such variations in the mulberry having been
attended to, selected, and rendered more or less constant. M. de
Quatrefages (10/9. 'Etudes sur les Maladies actuelles du Ver a Soie' 1859
page 321.) briefly describes six kinds cultivated in one valley in France:
of these the AMOUROUSO produces excellent leaves, but is 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 because the leaves can be easily
collected: lastly, the ROSO bears strong hardy leaves, produced in large
quantity, but with the one inconvenience, 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
caterpillars, 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 (10/10. 'Productive Resources of India' page 130.), "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 (10/11. 'Traite du Citrus' 1811. '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.), who almost devoted his life-time to the subject, considers that
there are four species, namely, sweet and bitter oranges, lemons, and
citrons, each of which has given rise to whole groups of varieties,
monsters, and supposed hybrids. One high authority (10/12. Mr. Bentham
'Review of Dr. A. Targioni-Tozzetti, Journal of Hort. Soc.' volume 9 page
133.) 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 (10/13. 'Geograph. Bot.' page 863.),
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
mentions two other forms cultivated in Japan and Java, which he ranks
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 bergamotte, as probably hybrids.
I have briefly abstracted these opinions for the sake of showing those who
have never attended to such subjects, how perplexing 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 characters almost perfectly by seed. Sweet and
bitter oranges differ in no important respect except in the flavour of
their fruit, but Gallesio (10/14. 'Teoria della Riproduzione' pages 52-57.)
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, etc. 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 a soft-shelled pine-tree into the dignity of an aboriginally
created species. The positive assertion made by Macfayden (10/15. Hooker
'Bot. Misc.' volume 1 page 302; volume 2 page 111.) that the pips of sweet
oranges produced 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,
partly resembling their parent-forms. I can adduce another case: the myrtle
leaved orange is ranked by all authors as a variety, but is very distinct
in general aspect: in my father's greenhouse, during many years, it rarely
yielded any fruit, 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 (10/16. 'Teoria della Riproduzione'
page 53.) they largely intercross without artificial aid; thus he
positively states that seeds taken from lemon-trees (C. lemonum) growing
mingled with the citron (C. medica), which is generally considered as a
distinct species, produced a graduated series of varieties between these
two forms. Again, an 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 determining whether to rank these forms as species or varieties; for
it is now known that undoubted species of Verbascum, Cistus, Primula,
Salix, etc., 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 statement by another
(10/17. Gallesio 'Teoria della Riproduzione' page 69.) namely, that when he
impregnated the flowers of the common orange with the pollen taken from
undoubted VARIETIES 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 remarkable
facts in vegetable physiology: Gallesio (10/18. Ibid page 67.) 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 of species, is a subject which I
shall fully discuss in the following chapter.
The second remarkable fact is, that two supposed hybrids (10/19. Gallesio
'Teoria della Riproduzione' pages 75, 76.) (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
(10/20. 'Gardener's Chronicle' 1841 page 613.), is an analogous case, I
know not. But to this subject I shall have to recur.
I will conclude by giving from A. Risso (10/21. 'Annales du Museum' tome 20
page 188.) 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
embroidered with yellow, borne on footstalks 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.
PEACH AND NECTARINE (Amygdalus persica).
The best authorities are nearly unanimous that the peach has never been
found wild. It was introduced from Persia into Europe a little before the
Christian era, and at this period few varieties existed. Alph. De Candolle
(10/22. 'Geograph. Bot.' page 882.), 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 character 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.
(FIGURE 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. 8. Smyrna Almond.)
Andrew Knight (10/23. 'Transactions of Hort. Soc.' volume 3 page 1 and
volume 4 page 396 and note to page 370. A coloured drawing is given of this
hybrid.), 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 almond; and in this he
has been followed by various authors. (10/24. 'Gardener's Chronicle' 1856
page 532. A writer, it may be presumed Dr. Lindley, remarks on the perfect
series which may be formed between the almond and the peach. Another high
authority, Mr. Rivers, who has had such wide experience, strongly suspects
('Gardener's Chronicle' 1863 page 27) that peaches, if left to a state of
nature, would in the course of time retrograde into thick-fleshed almonds.)
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, slightly furrowed,
much flattened, and elongated stone, protected by a tough, greenish layer
of bitter flesh. Mr. Bentham (10/25. 'Journal of Hort. Soc.' volume 9 page
168.) has particularly 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
compressed, in size, shape, strength, and in the depth of the furrows, as
may be seen in figure 42 (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
(No. 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
horticultural 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 swollen, with
the aspect of a peach, including a hard stone surrounded by a fleshy
covering, which is sometimes eatable. (10/26. Whether this is the same
variety as one lately mentioned ('Gardener's Chronicle' 1865 page 1154) by
M. Carriere under the name of persica intermedia, I know not; this variety
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.) A remarkable statement by M. Luizet has recently appeared in the
'Revue Horticole' (10/27. Quoted in 'Gardener's Chronicle' 1866 page 800.),
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. Carriere, 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 succession 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 instance, 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 (10/28. Quoted in 'Journal
de La Soc. Imp. d'Horticulture' 1855 page 238.) 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 most 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
marvellous manner.
One fact, however, is opposed to this conclusion. A hybrid, raised by
Knight from the sweet almond by the pollen of the peach, produced 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 sterility cannot be accounted for by the youth of the
trees (and this 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 good 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 (10/29. 'Teoria della Riproduzione
Vegetale' 1816 page 86.) says he has verified this with respect to eight
races of the peach. Mr. Rivers (10/30. 'Gardener's Chronicle' 1862 page
1195.) has given some striking instances from his own experience, and it is
notorious that good peaches are constantly raised in North America from
seed. Many of the American sub-varieties come true or nearly true to their
kind, such as the white-blossom, several of the yellow-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.
(10/31. Mr. Rivers 'Gardener's Chronicle' 1859 page 774.) 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. (10/32. Downing 'The Fruits of America' 1845 pages
475, 489, 492, 494, 496. See also F. Michaux 'Travels in N. America'
English translation page 228. For similar cases in France see Godron 'De
l'Espece' tome 2 page 97.) With respect to nectarines, both cling and
freestones are known in North America to reproduce themselves by seed.
(10/33. Brickell 'Nat. Hist. of N. Carolina' page 102 and Downing 'Fruit
Trees' page 505.) In England the new white nectarine was a seedling of the
old white, and Mr. Rivers (10/34. 'Gardener's Chronicle' 1862 page 1196.)
has recorded several similar cases. From this strong tendency to
inheritance, which both peach and nectarine trees exhibit,--from certain
slight constitutional differences (10/35. The peach and nectarine do not
succeed equally well in the some soil: see Lindley 'Horticulture' page
351.) 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 cannot 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 identity: and an eminent botanist has quite recently (10/36.
Godron 'De l'Espece' tome 2 1859 page 97.) maintained that the nectarine
"probably constitutes a distinct species."
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 (10/37. 'Transact. Hort. Soc.' volume 6 page 394.) that the
Boston nectarine was produced from a peach-stone, and this nectarine
reproduced itself by seed. (10/38. Downing's 'Fruit Trees' page 502.) Mr.
Rivers states (10/39. 'Gardener's Chronicle' 1862 page 1195.) that from
stones of three distinct varieties of the peach he raised three varieties
of nectarine; and in one of these cases no nectarine grew near the parent
peach-tree. In another instance Mr. Rivers raised a nectarine from a peach,
and in the succeeding generation another nectarine from this nectarine.
(10/40. 'Journal of Horticulture' February 5, 1866 page 102.) 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 instances recorded by Mr. Rivers;
and in two of these instances the parent nectarines had been seedlings from
other nectarines. (10/41. Mr. Rivers in 'Gardener's 'Chronicle' 1859 page
774, 1862 page 1195; 1865 page 1059; and 'Journal of Hort.' 1866 page 102.)
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 nectarines, 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 producing a
nectarine (10/42. 'Correspondence of Linnaeus' 1821 pages 7, 8, 70.) and in
1766 he added two other instances. In the same work, the editor, Sir J.E.
Smith, describes the more remarkable 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 (10/43. 'Transact. Hort. Soc.' volume 1 page 103.)
records six other cases of peach-trees producing nectarines. Three of the
varieties are named; viz., the Alberge, 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 (10/44. Loudon's 'Gardener's Mag.' 1826 volume 1.
page 471.) a clingstone peach, purchased as the Chancellor, was planted in
1815, and in 1824, after having previously produced peaches alone, bore on
one branch twelve nectarines; in 1825 the same branch yielded twenty-six
nectarines, and in 1826 thirty-six nectarines, together with eighteen
peaches. One of the peaches was almost as smooth on one side as a
nectarine. The nectarines were as dark as, but smaller than, the Elruge.
At Beccles a Royal George peach (10/45. Loudon's 'Gardener's Mag.' 1828
page 53.) produced a fruit, "three parts of it being peach and one part
nectarine, quite distinct in appearance as well as in flavour." The lines
of division were longitudinal, as represented in the woodcut. A nectarine-
tree grew five yards from this tree.
Professor Chapman states (10/46. Ibid 1830 page 597.) 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 (10/47. 'Gardener's Chronicle' 1841 page 617.)
produced this year a nectarine between two peaches; a nectarine-tree grew
close by.
In 1844 (10/48. 'Gardener's Chronicle' 1844 page 589.) a Vanguard peach-
tree produced, in the midst of its ordinary fruit, a single red Roman
nectarine.
Mr. Calver is stated (10/49. 'Phytologist' volume 4 page 299.) to have
raised in the United States a seedling peach which produced a mixed crop of
both peaches and nectarines.
Near Dorking (10/50. 'Gardener's Chronicle' 1856 page 531.) a branch of the
Teton de Venus peach, which reproduces itself truly by seed (10/51. Godron,
'De l'Espece' tome 2 page 97.), 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 nectarines, but
at Carclew (10/52. 'Gardener's Chronicle' 1856 page 531.) the unique case
occurred, of a nectarine-tree, raised twenty years before from seed and
never grafted, producing a fruit half peach and half nectarine;
subsequently bore a perfect peach.
To sum up the foregoing facts; we have excellent evidence of peach-stones
producing nectarine-trees, and of nectarine-stones producing peach-trees,--
of the same tree bearing peaches and nectarines,--of peach-trees suddenly
producing by bud-variation nectarines (such nectarines reproducing
nectarines by seed), as well as fruit in part nectarine and in part peach,-
-and, lastly, of one nectarine-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 have given birth by bud-variation or by seed to peaches,
oftener than peaches to nectarines; but this is by no means the case.
Two explanations have been suggested to account for these conversions.
First, that the parent trees have been in every case hybrids (10/53. Alph.
De Candolle 'Geograph. Bot.' page 886.) 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 hative nectarine (10/54. Thompson in Loudon's 'Encyclop. of
Gardening' page 911.), produces peaches, but these are said SOMETIMES to
partake of the smoothness and flavour of the nectarine. But let it be
observed that in the previous list no less than six well-known varieties
and several unnamed varieties of the peach have once suddenly produced
perfect nectarines by bud variation: 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: although this certainly is possible, it cannot here apply; for
we have not a shadow of evidence that a branch which has borne fruit
directly affected by foreign pollen is so profoundly modified as afterwards
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 Carclew 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
following chapter many analogous cases of bud-variation will he given.
The varieties of the peach and the 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. (10/55. 'Catalogue of Fruit in Garden of Hort.
Soc.' 1842 page 105.) 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.
The varieties of the peach have largely increased in number since the
Christian era, when from two to five varieties were known (10/56. Dr. A.
Targioni-Tozzetti 'Journal Hort. Soc.' volume 9 page 167. Alph. de Candolle
'Geograph. Bot.' page 885.) 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 (10/57. 'Transact. Hort. Soc.' volume 5 page
554. See also Carriere 'Description et Class. des Varietes de Pechers.')
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 separating 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 margin of the petal is 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
reniform glands (10/58. Loudon's 'Encyclop. of Gardening' page 907.) and
some few peaches, such as the Brugnen, bear on the same tree both globular
and kidney-shaped glands. (10/59. M. Carriere in 'Gardener's Chronicle'
1865 page 1154.) According to Robertson (10/60. 'Transact. Hort. Soc.'
volume 3 page 332. See also 'Gardener's Chronicle' 1865 page 271 to same
effect. Also 'Journal of Horticulture' September 26, 1865 page 254.) 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 maturity, in the fruit keeping well, and in hardiness,--the
latter circumstance being especially attended to in the United States.
Certain 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.
(10/61. 'Transact. Hort. Soc.' volume 4 page 512.) Another Chinese variety,
called the Honey-peach, is remarkable from the fruit terminating in a long
sharp point; its leaves are glandless and widely dentate. (10/62. 'Journal
of Horticulture' September 8, 1853 page 188.) The Emperor of Russia peach
is a third singular variety, having deeply double-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.
(10/63. 'Transact. Hort. Soc.' volume 6 page 412.)
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. (10/64. 'Gardener's Chronicle' 1857 page 216.) One of these
varieties, called the camellia-flowered, bears flowers above 2 1/4 inches
in diameter, whilst those of the fruit-bearing kinds do not at most exceed
1 1/4 inch in diameter. The flowers of the double-flowered peaches have the
singular property (10/65. 'Journal of Hort. Soc.' volume 2 page 283.) 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
characterised, 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 Caucasian region. (10/66. Alph. de Candolle
'Geograph. Bot.' page 879.) On this view the varieties deserve notice,
because they illustrate differences supposed by some botanists to be of
specific value in the almond and plum. The best monograph on the apricot is
by Mr. Thompson (10/67. 'Transact. Hort. Soc.' 2nd series volume 1 1835
page 56. See also 'Cat. of Fruit in Garden of Hort. Soc.' 3rd edition
1842.) 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, sometimes 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 clinging to the stone, as in the
last-mentioned kind, or in readily separating from it, as in the Turkey-
apricot. In all these differences we see the closest analogy with the
varieties of the peach and nectarine. 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 Moorpark, and generally in the Hemskirke, the stone
presents a singular character in being perforated, with a bundle of fibres
passing 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 bitter
than in some other kinds; slightly bitter in the Royal; and "sweet like a
hazel-nut" 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." (10/68. Downing 'The Fruits of
America' 1845 page 157: with respect to the Alberge apricot in France see
page 153.) According to Mr. Rivers (10/69. 'Gardener's Chronicle' 1863 page
364.), seedling apricots deviate but little from the character of their
race: in France the Alberge is constantly reproduced from seed with but
little variation. In Ladakh, according to Moorcroft (10/70. 'Travels in the
Himalayan Provinces' volume 1 1841 page 295.) 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).
(FIGURE 43. PLUM STONES, of natural size, viewed laterally. 1. Bullace
Plum. 2. Shropshire Damson. 3. Blue Gage. 4. Orleans. 5. Elvas. 6. Denyers
Victoria. 7. Diamond.)
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
naturalised in England. (10/71. See an excellent discussion on this subject
in Hewett C. Watson 'Cybele Britannica' volume 4 page 80.) It is not at all
improbable, in accordance with some observations made by Mr. Rivers (10/72.
'Gardener's Chronicle' 1865 page 27.), 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 (10/73. 'De l'Espece'
tome 2 page 94. On the parentage of our plums see also Alph. De Candolle
'Geograph. Bot.' page 878. Also Targioni-Tozzetti 'Journal Hort. Soc.'
volume 9 page 164. Also Babington 'Manual of Brit. Botany' 1851 page 87.)
that the cultivated varieties may be divided into two main groups, which he
supposes to be descended from two 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 diversified 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 (10/74. 'Fruits
of America' pages 276, 278, 284, 310, 314. Mr. Rivers raised ('Gardener's
Chronicle' 1863 page 27) from the Prune-peche, which bears large, round,
red plums on stout, robust shoots, a seedling which bears oval, smaller
fruit on shoots that are so slender as to be almost pendulous.) 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 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 collection 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 surprising 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, of which seven of first-rate quality
have been recently introduced into England. (10/75. 'Gardener's Chronicle'
1855 page 726.) Varieties occasionally arise having an innate adaptation
for certain soils, almost as strongly pronounced as with natural species
growing on the most distinct geological formations; thus in America the
imperial gage, differently from almost all other kinds, "is peculiarly
fitted for DRY LIGHT soils where many sorts drop their fruit," whereas on
rich heavy soils the fruit is often insipid. (10/76. Downing 'Fruit Trees'
page 278.) 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 native Yorkshire it bears abundantly: one of my
relations also repeatedly tried in vain to grow this variety in a sandy
district in Staffordshire.
Mr. Rivers has given (10/77. 'Gardener's Chronicle' 1863 page 27. Sageret
in his 'Pomologie Phys.' page 346 enumerates five kinds which can be
propagated in France by seed: see also Downing 'Fruit Trees of America'
page 305, 312, etc.) a number of interesting facts, showing 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 instances 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, etc.).
Botanists believe that our cultivated cherries are descended from one, two,
four, or even more wild stocks. (10/78. Compare Alph. De Candolle
'Geograph. Bot.' page 877. Bentham and Targioni-Tozzetti in 'Hort. Journal'
volume 9 page 163; Godron 'De l'Espece' tome 2 page 92.) 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 fertilised by pollen of the
Elton cherry; for these hybrids produced in all only five cherries, and one
alone of these contained a seed. (10/79. 'Transact. Hort. Soc.' volume 5
1824 page 295.) Mr. Thompson (10/80. Ibid second series volume 1 1835 page
248.) 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 cherries are 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. (10/81. ) 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 cherry several flower-
peduncles arise from a common peduncle, upwards 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 could 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 extraordinary statement is made that
all the leaf-bearing shoots spring from old flower-buds. Lastly, there is
an important physiological distinction 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. (10/82.
These several statements are taken from the four following works, which
may, I believe, be trusted: Thompson in 'Hort. Transact.' see above;
Sageret 'Pomologie Phys.' 1830 pages 358, 364, 367, 379; 'Catalogue of the
Fruit in the Garden of Hort. Soc.' 1842 pages 57, 60; Downing 'The Fruits
of America' 1845 pages 189, 195, 200.)
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 praecox or paradisiaca, do not deserve to
be ranked as distinct species. The P. praecox is supposed by some authors
(10/83. Mr. Lowe states in his 'Flora of Madeira' quoted in 'Gardener's
Chronicle' 1862 page 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 Canaries, 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.) 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
stocks, it is asserted (10/84. See 'Journ. of Hort. Tour, by Deputation of
the Caledonian Hort. Soc.' 1823 page 459.) cannot be propagated true by
seed. The common wild crab varies considerably in England; but many of the
varieties are believed to be escaped seedlings. (10/85. H.C. Watson 'Cybele
Britannica' volume 1 page 334.) 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 various 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." (10/86. Loudon's 'Gardener's Mag.' volume
6 1830 page 83.) Another Russian apple, the white Astracan, possesses the
singular property of becoming transparent, when ripe, like some sorts of
crabs. The API ETOILE has five prominent ridges, hence its name; the API
NOIR is nearly black: the TWIN CLUSTER PIPPIN often bears fruit joined in
pairs. (10/87. See 'Catalogue of Fruit in Garden of Hort. Soc.' 1842 and
Downing 'American Fruit Trees.') The trees of the several sorts differ
greatly in their periods of leafing and flowering; in my orchard the COURT
PENDU PLAT produces leaves so late, that during several springs I thought
that it was 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. (10/88. Loudon's Gardener's
Magazine' volume 4 1828 page 112.) In some kinds the fruit ripens in mid-
summer; in others, late in the autumn. These several differences in
leafing, flowering, and fruiting, are not at all necessarily correlated;
for, as Andrew Knight has remarked (10/89. 'The Culture of the Apple' page
43. Van Mons makes the same remark on the pear 'Arbres Fruitiers' tome 2
1836 page 414.), 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 (10/90. Lindley's
'Horticulture' page 116. See also Knight on the Apple-Tree, in 'Transact.
of Hort. Soc.' volume 6 page 229.), which is the glory of the orchards near
New York; and so it is with several varieties which we have imported from
the Continent. On the other hand, our Court of Wick succeeds well under the
severe climate of Canada. The Caville 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 a few fruit even during
the first year. (10/91. Transact. Hort. Soc.' volume 1 1812 page 120.) Mr.
Rivers has recently described (10/92. 'Journal of Horticulture' March 13,
1866 page 194.) 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 certain soils. But perhaps
the strangest constitutional peculiarity is that the Winter Majetin is not
attacked by the mealy bug or coccus; Lindley (10/93. 'Transact. Hort. Soc.'
volume 4 page 68. For Knight's case see volume 6 page 547. When the coccus
first appeared in this country it is said (volume 2 page 163) that it was
more injurious to crab-stocks than to the apples grafted on them. The
Majetin apple has been found equally free of the coccus at Melbourne in
Australia ('Gardener's Chronicle' 1871 page 1065). The wood of this tree
has been there analysed, and it is said (but the fact seems a strange one)
that its ash contained over 50 per cent of lime, while that of the crab
exhibited not quite 23 per cent. In Tasmania Mr. Wade ('Transact. New
Zealand Institute' volume 4 1871 page 431) raised seedlings of the Siberian
Bitter Sweet for stocks, and he found barely one per cent of them attacked
by the coccus. Riley shows ('Fifth Report on Insects of Missouri' 1873 page
87) that in the United States some varieties of apples are highly
attractive to the coccus and others very little so. Turning to a very
different pest, namely, the caterpillar of a moth (Carpocapsa pomonella),
Walsh affirms ('The American Entomologist' April 1869 page 160) that the
maiden-blush "is entirely exempt from apple-worms." So, it is said, are
some few other varieties; whereas others are "peculiarly subject to the
attacks of this little pest.") 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 insects just
above the stock, but that three days afterwards they entirely disappeared;
this apple, however, was raised from a cross between the Golden Harvey 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. (10/94. 'Mem. de La Soc. Linn. de Paris'
tome 3 1825 page 164; and Seringe 'Bulletin Bot.' 1830 page 117.) 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 (10/95. Gardener's Chronicle' 1849 page 24.)
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. (10/96. R.
Thompson, in 'Gardener's Chronicle' 1850 page 788.) Yet it was a mistake to
suppose that with most varieties the characters are not to a certain extent
inherited. 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, etc. (10/97. Sageret
'Pomologie Physiologique' 1830 page 263. Downing 'Fruit Trees' pages 130,
134, 139, etc. Loudon's 'Gardener's Mag.' volume 8 p. 317. Alexis Jordan
'De l'Origine des diverses Varietes' in 'Mem. de l'Acad. Imp. de Lyon' tome
2 1852 pages 95, 114. 'Gardener's Chronicle' 1850 pages 774, 788.), 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 celebrated botanists in Europe, M. Decaisne, has
carefully studied the many varieties (10/98. 'Comptes Rendus' July 6,
1863.); although he formerly believed that they were derived from more than
one species, he now thinks that all belong 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. Decaisne raised many seedlings from four distinct kinds, and has
carefully recorded the variations in each. Notwithstanding this extreme
degree of variability, it is now positively known that many kinds reproduce
by seed the leading characters of their race. (10/99. 'Gardener's
Chronicle' 1856 page 804; 1857 page 820; 1862 page 1195.)
STRAWBERRIES (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. (10/100. 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 (Downing 'Fruits' page
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. vesca, or our common
wood-strawberry.) 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 cultivated, but only five varieties of
Fragaria vesca, with some sub-varieties, 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 authority, M. Gay, to be merely a
strongly marked race of F. chiloensis. (10/101. 'Le Fraisier' par le Comte
L. de Lambertye 1864 page 50.) These five or six forms have been ranked by
most botanists as specifically distinct; but this may be doubted, for
Andrew Knight (10/102. 'Transact. Hort. Soc.' volume 3 1820 page 207.) who
raised no less than 400 crossed strawberries, asserts that the F.
virginiana, chiloensis and grandiflora "may be made to breed together
indiscriminately," and he found, in accordance with the principle of
analogous variation, "that similar varieties could be obtained from the
seeds of any one of them."
Since Knight's time there is abundant and additional evidence (10/103. See
an account by Prof. Decaisne, and by others in 'Gardener's Chronicle' 1862
page 335 and 1858 page 172; and Mr. Barnet's paper in 'Hort. Soc.
Transact.' volume 6 1826 page 170.) of the extent to which the American
forms spontaneously cross. We owe indeed to such crosses most of our
choicest existing varieties. 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 offspring
from the Hautbois, though fruiting well, never produced seed, with the
exception of a single one, which reproduced the parent hybrid form.
(10/104. 'Transact. Hort. Soc.' volume 5 1824 page 294.) 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
barren. Mr. W. Smith, of York, has raised similar hybrids with equally poor
success. (10/105. 'Journal of Horticulture' December 30, 1862 page 779. See
also Mr. Prince to the same effect ibid 1863 page 418.) We thus see
(10/106. For additional evidence see 'Journal of Horticulture' December 9,
1862 page 721.) 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 be classed under some one of the
above 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. (10/107. 'Le Fraisier' par le
Comte L. de Lambertye pages 221, 230.) 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 see now 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 remarkable from "its
peculiar dark and polished surface, and from presenting an appearance
entirely unlike that of any other kind." (10/108. 'Transact. Hort. Soc.'
volume 6 page 200.) Although 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 embedded in the pulp, is, according to De Jonghe
(10/109. 'Gardener's Chronicle' 1858 page 173.) absolutely 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 Linnaeus 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,
or present intermediate states. (10/110. Godron 'De l'Espece' tome 1 page
161.) A variety raised by Mr. Myatt (10/111. 'Gardener's Chronicle' 1851
page 440.), apparently belonging to one of the American forms presents a
variation 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. (10/112. F. Gloede in 'Gardener's
Chronicle' 1862 page 1053.) A strawberry of the American Pine section is
also said to make but few runners. (10/113. Downing 'Fruits' page 532.)
Much has been written on the sexes of strawberries; the true Hautbois
properly bears the male and female organs on separate plants (10/114.
Barnet in 'Hort. Transact.' volume 6 page 210.), and was consequently named
by Duchesne dioica; but it frequently produces hermaphrodites; and Lindley
(10/115. 'Gardener's Chronicle' 1847 page 539.), by propagating such plants
by runners, at the same time destroying the males, soon raised a self-
prolific stock. The other species often showed a tendency towards an
imperfect separation of the sexes, as I have noticed with plants forced in
a hot-house. Several English varieties, which in this country are free from
any such tendency, when cultivated in rich soils under the climate of North
America (10/116. For the several statements with respect to the American
strawberries see Downing 'Fruits' page 524; 'Gardener's Chronicle' 1843
page 188; 1847 page 539; 1861 page 717.) commonly produce plants with
separate sexes. Thus a whole acre of Keen's Seedlings in the United States
has been observed to be almost 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 investigate this subject, report that "few varieties have the flowers
perfect in both sexual organs," etc. 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 apparently depends more on the nature
of the soil than on the climate; a famous gardener says that "no mortal
could grow the British Queen at Shrubland Park unless the whole nature of
the soil was altered." (10/117. Mr. D. Beaton in 'Cottage Gardener' 1860
page 86. See also 'Cottage Gardener' 1855 page 88 and many other
authorities. For the Continent see F. Gloede in 'Gardener's Chronicle' 1862
page 1053.) La Constantine is one of the hardiest kinds, and can withstand
Russian winters, but it is easily burnt by the sun, so that it will not
succeed in certain soils either in England or the United States. (10/118.
Rev. W.F. Radclyffe in 'Journal of Hort.' March 14, 1865 page 207.) 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." (10/119. Mr. H. Doubleday in 'Gardener's Chronicle' 1862 page
1101.) 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 exactly
the same manner, were not at all infested by this fungus. (10/120.
'Gardener's Chronicle' 1854 page 254.) 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 Europe; 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 mentions it in 1573, and Parkinson
specifies eight varieties in 1629; the Catalogue of the Horticultural
Society for 1842 gives 149 varieties, and the lists of the Lancashire
nurserymen are said to include above 300 names. (10/121. Loudon's
'Encyclop. of Gardening' page 930; and Alph. De Candolle 'Geograph. Bot.'
page 910.) In the 'Gooseberry Grower's Register' for 1862 I find that 243
distinct varieties have won prizes at various periods, 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 this shows that all the kinds, numerous as they are, can be
recognised with certainty.
The bushes differ in their manner of growth, being erect, or spreading, or
pendulous. The periods of leafing and flowering differ both absolutely 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. (10/122. Loudon's 'Gardener's Magazine' volume 4
1828 page 112.) 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 different varieties the fruit varies in
abundance, in the period of maturity, 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. (10/123.
The fullest account of the gooseberry is given by Mr. Thompson in
'Transact. Hort. Soc.' volume 1 2nd series 1835 page 218 from which most of
the foregoing facts are taken.)
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 bracteae;
and this is the most singular deviation of structure which I have observed.
These same flowers also varied much in the 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 Pastime gooseberry "extra bracts are often
attached to the sides of the fruit." (10/124. 'Catalogue of Fruits of Hort.
Soc. Garden' 3rd edition 1842.)
The most interesting point in the history of the gooseberry is the steady
increase in the size of the fruit. Manchester is the metropolis of the
fanciers, and prizes from five shillings to five or ten pounds are yearly
given for the heaviest fruit. The 'Gooseberry Growers 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. (10/125. Mr. Clarkson of Manchester on the Culture of the
Gooseberry in Loudon's 'Gardener's Magazine' volume 4 1828 page 482.) 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 culture has been carried on. The fruit of the wild gooseberry is said
(10/126. Downing 'Fruits of America' page 213.) to weigh about a quarter of
an ounce or 5 dwts., that is, 120 grains; 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 the same variety reached the astonishing weight
of 37 dwts. 7 grs. (10/127. 'Gardener's Chronicle' 1844 page 811 where a
table is given; and 1845 page 819. For the extreme weights gained see
'Journal of Horticulture' July 26, 1864 page 61.) or 896 grs.; that is,
between seven or eight times the weight of the wild fruit. I find that a
small apple, 6 1/2 inches in circumference, has exactly this same weight.
The "London" gooseberry (which in 1852 had altogether gained 333 prizes)
has, up to the present year of 1875, 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 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 made, the soil is mulched, and
only a few berries are left on each bush (10/128. Mr. Saul of Lancaster in
Loudon's 'Gardener's Mag.' volume 3 1828 page 421; and volume 10 1834 page
42.) but the increase no doubt is in main part due to the continued
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 therefore here noticed. The walnut grows wild on
the Caucasus and in the Himalaya, where Dr. Hooker (10/129. 'Himalayan
Journals' 1854 volume 2 page 334. Moorcroft 'Travels' volume 2 page 146
describes four varieties cultivated in Kashmir.) found the fruit of full
size, but "as hard as a hickory-nut." It has been found fossil, as M. de
Saporta informs me, in the tertiary formation, of France.
In England the walnut presents considerable 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 titmice.
(10/130. 'Gardener's Chronicle' 1850 page 723.) 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. (10/131. Paper translated in Loudon's
'Gardener's Mag.' 1829 volume 5 page 202.) M. Cardan has minutely described
(10/132. Quoted in 'Gardener's Chronicle' 1849 page 101.) some singular
physiological peculiarities in the June-leafing variety, which produces its
leaves and flowers four or five weeks later than the common varieties; and
although in August it is apparently in exactly the same state of
forwardness as the other kinds, it retains its leaves and fruit much later
in the autumn. These constitutional peculiarities are strictly inherited.
Lastly, walnut-trees, which are properly monoicous, sometimes entirely fail
to produce male flowers. (10/133. 'Gardener's Chronicle' 1847 pages 541 and
558.)
NUTS (Corylus avellana).
Most botanists rank all the varieties under the same species, the common
wild nut. (10/134. The following details are taken from the 'Catalogue of
Fruits, 1842 in Garden of Hort. Soc.' page 103; and from Loudon's
'Encyclop. of Gardening' page 943.) The husk, or involucre, differs
greatly, being extremely short in Barr's Spanish, and extremely long in
filberts, in which it is contracted so as to prevent the nut falling out.
This kind of husk also protects the nut from birds, for titmice (Parus)
have been observed (10/135. 'Gardener's Chron.' 1860 page 956.) 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 varieties, but is thin in Cosford's-nut, and in
one variety is of a bluish 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
Cosford'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 (10/136. 'Annales des Sc. Nat. Bot.' 4th
series volume 6 1856 page 5.), a flood of light has recently been thrown on
this group of plants. M. Naudin, during many years, observed and
experimented on above 1200 living specimens, collected from all quarters of
the world. Six species are now recognised in the genus Cucurbita; but three
alone have been cultivated and concern us, namely, C. maxima and pepo,
which include all pumpkins, gourds, squashes, and the vegetable marrow, and
C. moschata. These three species are not known in a wild state; but Asa
Gray (10/137. 'American Journ. of Science' 2nd series volume 24 1857 page
442.) 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 (page 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 he 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 (pages 6, 16, 35) urges, "douees d'une
stabilite presque comparable a celle des especes les mieux caracterisees."
One variety, l'Orangin (pages 43, 63), has such prepotency in transmitting
its character, that when crossed with other varieties a vast majority of
the seedlings come true. Naudin, referring (page 47) to C. pepo, says that
its races "ne different des especes veritables qu'en ce qu'elles peuvent
s'allier les unes aux autres par voie d'hybridite, sans que leur
descendance perde la faculte de se perpetuer." 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 Gartner and Kolreuter. (10/138. Gartner
'Bastarderzeugung' 1849 s. 87 and s. 169 with respect to Maize; on
Verbascum ibid s. 92 and 181; also his 'Kenntniss der Befruchtung' s. 137.
With respect to Nicotiana see Kolreuter 'Zweite Forts.' 1764 s. 53; though
this is a somewhat different case.)
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 (pages 33, 46) exceeds
in value that of another by more than two thousand fold! When the fruit is
of very large size, the number produced is few (page 45); when of small
size, many are produced. No less astonishing (page 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 (page 34), namely, from six or seven to more than
twenty-five millimetres in length.
In the varieties which grow upright or do not run and climb, the tendrils,
though useless (page 31), are either present or are represented by various
semi-monstrous organs, or are quite absent. The tendrils are even absent in
some running varieties in which the stems are much elongated. It is a
singular fact that (page 31) in all the varieties with dwarfed stems, the
leaves closely resemble each other in shape.]
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 (10/139. Godron 'De l'Espece' tome 2 page 64.) 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 etamines restent invariables dans chacune
d'elles." Yet M. Naudin, in describing Cucurbita pepo (page 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. Neanmoins, je crois qu'on
la distinguera toujours facilement des deux autres especes, si l'on veut ne
pas perdre de vue les caracteres differentiels que je m'efforce de faire
ressortir. Ces caracteres sont quelquefois peu marques: il arrive meme que
plusieurs d'entre eux s'effacent presque entierement, mais ii 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 important 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 Gasparini to separate certain pumpkins as a
DISTINCT GENUS; but Naudin says (page 20), these parts have no constancy,
and in the flowers of the Turban varieties of C. maxima they sometimes
resume their ordinary structure. Again, in C. maxima, the carpels (page 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 varieties,
and graduates into the common form in which the carpels are almost entirely
enveloped within the receptacle. In C. moschata the ovarium (page 50)
varies greatly in shape, being oval, nearly spherical, or cylindrical, more
or less swollen in the upper part, or constricted round the middle, and
either straight or curved. When the ovarium is short and oval the interior
structure does not differ from that of C. maxima and pepo, but when it is
elongated the carpels occupy only the terminal and swollen portion. I may
add that in one variety of the cucumber (Cucumis sativus) the fruit
regularly contains five carpels instead of three. (10/140. Naudin 'Annal.
des Sc. Nat.' 4th series Bot. tome 11 1859 page 28.) 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 (10/141. 'Memoire sur les Cucurbitacees' 1826 pages 6, 24.) 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 (10/142. 'Flore des Serres' October 1861 quoted in
'Gardener's Chronicle' 1861 page 1135. I have often consulted and taken
some facts from M. Naudin's Memoir on Cucumis in 'Annal. des Sc. Nat.' 4th
series Bot. tome 11 1859 page 5.) 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 had not come to the end of them: he divides them
into ten sections, including numerous sub-varieties which all intercross
with perfect ease. (10/143. See also Sageret 'Memoire' page 7.) Of the
forms considered by Naudin to be varieties, 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 modified 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 sometimes 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 assuming the
characteristic features of distinct species and even of distinct though
allied genera: thus the serpent-melon has some resemblance to the fruit of
Trichosanthes anguina; we have seen that other varieties closely resemble
cucumbers; some Egyptian varieties 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 spontaneous 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 "extraordinary
production of races and varieties by a single species and their permanence
when not interfered with by crossing, are phenomena well calculated to
cause reflection."
USEFUL AND ORNAMENTAL TREES.
Trees deserve a passing notice on account of the numerous varieties which
they present, differing in their precocity, in their manner of growth,
their 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.
(10/144. Loudon's 'Arboretum et Fruticetum' volume 2 page 1217.) Of hollies
no less than eighty-four varieties are grown alongside each other in Mr.
Paul's nursery. (10/145. 'Gardener's Chronicle' 1866 page 1096.) 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 (10/146.
'Geograph. Bot.' page 1096.) 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 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 weeping habit is
sometimes inherited, though in a singularly capricious manner. In the
Lombardy poplar, and in certain fastigiate or pyramidal varieties of
thorns, junipers, oaks, etc., we have an opposite kind of growth. The
Hessian oak (10/147. 'Gardener's Chronicle' 1842 page 36.), which is famous
from its fastigiate habit and size, bears hardly any resemblance in general
appearance 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 fastigiate 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. (10/148. Loudon's
'Arboretum et Fruticetum' volume 3 page 1731.) The fastigiate Juniper (J.
suecica) likewise transmits its character by seed. (10/149. Ibid volume 4
page 2489.) Dr. Falconer informs me that in the Botanic Gardens at Calcutta
the great heat caused apple-trees to become fastigiate; and we thus see the
same result following from the effects of climate and from some unknown
cause. (10/150. Godron 'De l'Espece' tome 2 page 91 describes four
varieties of Robinia remarkable from their manner of growth.)
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 sometimes weakly inherited
(10/151. 'Journal of a Horticultural Tour, by Caledonian Hort. Soc.' 1823
page 107. Alph. De Candolle 'Geograph. Bot.' page 1083. Verlot 'Sur La
Production des Varietes' 1865 page 55 for the Barberry.); 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. (10/152.
Loudon's 'Arboretum et Fruticetum' volume 2 page 508.) In fact, nearly all
the peculiar varieties evince a tendency, more or less strongly marked, to
reproduce themselves by seed. (10/153. Verlot 'Des Varietes' 1865 page 92.)
This is to a certain extent the case, according to Bosc (10/154. Loudon's
'Arboretum et Fruticetum' volume 3 page 1376.), 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 (Carpinus betulus), which bears on each twig leaves of two shapes,
"several plants raised from seed all retained "the same peculiarity."
(10/155. 'Gardener's Chronicle' 1841 page 687.) 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 character by seed. (10/156. Godron 'De l'Espece'
tome 2 page 89. In Loudon's 'Gardener's Mag.' volume 12 1836 page 371, a
variegated bushy ash is described and figured, as having simple leaves; it
originated in Ireland.) The occurrence, in trees belonging to widely
different orders, of weeping and fastigiate 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 considered 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. (10/157. 'Gardener's Chronicle' 1863 page 575.)
Every one must have noticed how certain individual trees regularly put
forth and shed their leaves earlier or later than others of the same
species. There is a famous horse-chestnut 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.
(10/158. Quoted from Royal Irish Academy in 'Gardener's Chronicle' 1841
page 767.) There is a Cornish variety of the elm which is almost an
evergreen, and is so tender that the shoots are often killed by the frost;
and the varieties of the Turkish oak (Q. cerris) may be arranged as
deciduous, sub-evergreen, and evergreen. (10/159. Loudon 'Arboretum et
Fruticetum:' for Elm see volume 3 page 1376; for Oak page 1846.)
SCOTCH FIR (Pinus sylvestris).
I allude to this tree as it bears on the question of the greater
variability of our hedgerow trees compared with those under strictly
natural conditions. A well-informed writer (10/160. 'Gardener's Chronicle'
1849?, page 822.) states that the Scotch fir presents few varieties in its
native Scotch forests; but that it "varies much in figure and foliage, and
in the size, shape, and colour of its cones, when several generations 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." (10/161. 'Arboretum et Fruticetum' volume 4 page 2150.) I may
mention one rather important point in which this tree occasionally varies;
in the classification of the Coniferae, 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 enclosed, but specimens have been
observed with groups of three leaves in a sheath. (10/162. 'Gardener's
Chronicle' 1852 page 693.) Besides these differences in the semi-cultivated
Scotch fir, there are in several parts of Europe natural or geographical
races, which have been ranked by some authors as distinct species. (10/163.
See 'Beitrage zur Kenntniss Europaischer Pinus-arten von Dr. Christ: Flora,
1864.' He shows that in the Ober-Engadin P. sylvestris and montana are
connected by intermediate links.) Loudon (10/164. 'Arboretum et Fruticetum'
volume 4 pages 2159 and 2189.) considers P. pumilio, with its several sub-
varieties, as mughus, nana, etc., 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 oxyacantha).
The Hawthorn 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 (10/165. Ibid volume 2 page 830; Loudon's 'Gardener's Mag.'
volume 6 1830 page 714.) 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 the hawthorn has yielded more varieties than most other trees, is that
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. (10/166.
Loudon's 'Arboretum et Fruticetum' volume 2 page 834.) 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.
(10/167. Loudon's 'Gardener's Mag.' volume 9 1833 page 123.) 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
(10/168. Ibid volume 11 1835 page 503.) 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 circumstance alone would
render it difficult to detect the difference due to variation. For
instance, our Roses, Petunias, Calceolarias, Fuchsias, Verbenas, Gladioli,
Pelargoniums, etc., 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 (10/169. 'Gardener's
Chronicle' 1845 page 623.) a famous cultivator of this plant: some
varieties require 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 commencement of the season, others
at the close; one variety is known (10/170. D. Beaton in 'Cottage Gardener'
1860 page 377. See also Mr. Beck on the habits of Queen Mab in 'Gardener's
Chronicle' 1845 page 226.), which will stand "even pine-apple top and
bottom heat, 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 enable a plant in a state of nature to
become adapted to 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 colour, 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 flowers 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 Compositae, the corollas
of the central florets are greatly modified, and the modifications 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. (10/171. Moquin-Tandon 'Elements de Teratologie' 1841 page 213.) In
the "hose in hose" primulae, 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 (10/172. See also 'Cottage Gardener' 1860 page
133.) 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 developed from buds in the axils of
the scales of the involucre. A wonderful poppy has been described, in which
the stamens are converted into pistils; and so strictly was this
peculiarity inherited that, out of 154 seedlings, one alone reverted to the
ordinary and common type. (10/173. Quoted by Alph. de Candolle 'Bibl.
Univ.' November 1862 page 58.) 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 (10/174.
Knight 'Transact. Hort. Soc.' volume 4 page 322.) actually eighteen inches
in breadth. Peloric races of Gloxinia speciosa and Antirrhinum majus can be
propagated by seed, and they differ in a wonderful manner from the typical
form both in structure and appearance.
A much more remarkable modification has been recorded by Sir William and
Dr. Hooker (10/175. 'Botanical Magazine' tab. 5160 figure 4; Dr. Hooker in
'Gardener's Chronicle' 1860 page 190; Prof. Harvey in 'Gardener's
Chronicle' 1860 page 145; Mr. Crocker in 'Gardener's Chron.' 1861 page
1092.) 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 perfect hermaphrodite structure; and in these
flowers the perianth was inferior. To show the importance of this
modification under a classificatory point of view, I may quote what Prof.
Harvey says, namely, that had it "occurred in a state of nature, and had a
botanist collected a plant with such flowers, he would not only have placed
it in a distinct genus from Begonia, but would probably have considered 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 Saxifragae and Aristolochiaceae. 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. (10/176. Alph.
de Candolle 'Geograph. Bot.' page 1083; 'Gardener's Chronicle' 1861 page
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 page 64.) 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 this point of view such
works as Professor Moquin-Tandon's 'Teratologie' are highly instructive.
ROSES.
These flowers offer an instance of a number of forms generally ranked as
species, namely, R. centifolia, gallica, alba, damascena, spinosissima,
bracteata, indica, semperflorens, moschata, etc., 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 doubtful; thus, with respect to the British forms,
Babington makes seventeen, 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 (10/177. 'Rose Amateur's Guide' T. Rivers
1837 page 21.) 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 involved in much uncertainty." (10/178. 'Journal Hort. Soc.' volume 9
1855 page 182.) Nevertheless, Mr. Rivers in referring to R. indica (page
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 bud; such modified buds being
often called by gardeners sports. In the following chapter I shall fully
discuss this latter subject, and shall show that bud-variations can be
propagated not only by grafting and budding, but often by seed. Whenever a
new rose appears with any peculiar character, however produced, if it
yields seed, Mr. Rivers (page 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 page 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 between the
various kinds, but some constitutional peculiarities may be mentioned.
Several French roses (Rivers page 12) will not succeed in England; and an
excellent horticulturist (10/179. The Rev. W.F. Radclyffe in 'Journal of
Horticulture' March 14, 1865 page 207.) 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 nothing under a south wall." Many roses can be
forced, "many are totally unfit for forcing, among which is General
Jacqueminot." (10/180. 'Gardener's Chronicle' 1831 page 46.) From the
effects of crossing and variation Mr. Rivers enthusiastically anticipates
(page 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 (10/181. Mr. Sabine in 'Transact. Hort. Soc.'
volume 4 page 285.); 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 (10/182.
'An Encyclop. of Plants' by J.C. Loudon 1841 page 443.) it is said that
three hundred varieties could be procured in the nursery-gardens near
Glasgow; and these are described as blush, crimson, purple, red, marbled,
two-coloured, white, and yellow, and as differing much in the size and
shape of the flower.
PANSY OR HEARTSEASE (Viola tricolor, etc.).
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 nursery-man,
energetically commenced their culture; and in the course of a few years
twenty varieties could be purchased. (10/183. Loudon's 'Gardener's
Magazine' volume 11 1835 page 427; also Journal of Horticulture' April 14,
1863 page 275.) At about the same period, namely in 1813 or 1814, Lord
Gambier collected some wild plants, and his gardener, Mr. Thomson,
cultivated them, together with some common garden varieties, and soon
effected a great improvement. The first great change was the conversion 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 contrast 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
variously coloured, which are exhibited at our shows. But when I came to
enquire more closely, I found that, though the varieties were so modern,
yet that much confusion and doubt prevailed about their parentage. Florists
believe that the varieties (10/184. Loudon's 'Gardener's Magazine' volume 8
page 575: volume 9 page 689.) are descended from several wild stocks,
namely, V. tricolor, lutea, grandiflora, amoena, and altaica, more or less
intercrossed. And when I looked to botanical works to ascertain whether
these forms ought to be ranked 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 amoena (10/185. Sir J.E. Smith 'English Flora'
volume 1 page 306. H.C. Watson 'Cybele Britannica' volume 1 1847 page 181.)
is now looked at 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 (10/186.
Quoted from 'Annales des Sciences' in the Companion to the 'Bot. Mag.'
volume 1 1835 page 159.) who has paid particular 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 matters, Mr. H.C. Watson (10/187. 'Cybele Britannica' volume 1 page
173. See also Dr. Herbert on the changes of colour in transplanted
specimens, and on the natural variations of V. grandiflora, in 'Transact.
Hort. Soc.' volume 4 page 19.) says that, "while V. tricolor passes into V.
arvensis on the one side, it approximates so much towards V. lutea 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 proper season, they produced
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 inheritance, however, does hold good to a certain extent even with the
fleeting varieties of the Heartsease, for to gain good sorts it is
indispensable to sow the seed of good sorts. Nevertheless, in almost every
large seed-bed a few, almost wild seedlings reappear through reversion. 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 sometimes 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 number 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
extremity was considerably enlarged; and lastly, in some the basal part was
depressed, becoming, as usual, laterally compressed towards 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 the 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. tricolor,
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
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. (10/188. Salisbury in 'Transact. Hort.
Soc.' volume 1 1812 pages 84, 92. A semi-double variety was produced in
Madrid in 1790.) Mr. Sabine remarks that "it seems as if some period of
cultivation had been required before the fixed qualities of the native
plant gave way and began to sport into those changes which now so delight
us." (10/189. 'Transact. Hort. Soc.' volume 3 1820 page 225.) The flowers
have been greatly modified in shape from a flat to a globular form. Anemone
and ranunculus-like races (10/190. Loudon's 'Gardener's Mag.' volume 6 1830
page 77.) 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 (10/191. Loudon's 'Encyclop. of Gardening' page
1035.) have been raised from the same plant; yet, as Mr. Sabine has
remarked, "many of the seedlings 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 (10/192. 'Transact. Hort. Soc.' volume 1
page 91; and Loudon's 'Gardener's Mag.' volume 3 1828 page 179.); 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
(10/193. Mr. Wildman in 'Gardener's Chronicle' 1843 page 87. 'Cottage
Gardener' April 8, 1856 page 33.) and it has been noticed that some
varieties require much more moisture than others. (10/194. M. Faivre has
given an interesting account of the successive variations of the Chinese
primrose, since its introduction into Europe about the year 1820: 'Revue
des Cours Scientifiques' June 1869 page 428.)
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, etc., 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. (10/195. 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
page 342.) 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. 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,
however, 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
(10/196. 'Des Jacinthes, de leur Anatomie, Reproduction, et Culture'
Amsterdam 1768.) in 1768, it is stated that nearly 2,000 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
(10/197. Alph. de Candolle 'Geograph. Bot.' page 1082.) almost always yield
white hyacinths, and the yellow kinds come nearly true. The hyacinth is
remarkable from having 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 treatise asserts (page 43) that some
experienced florists, such as the famous G. Voorhelm, 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
(page 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 Coryphee, invariably produces (page 116) two
flower-stems, united together and covered by one skin. The flower-stem in
another kind (page 128) comes out of the ground in a coloured sheath,
before the appearance of the leaves, and is consequently liable 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 (page 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 cultivated for various purposes. But some of the
most variable plants, such as Kidney-beans, Capsicum, Millets, Sorghum,
etc., have been passed over; for botanists are not at all agreed which
kinds ought to rank as species and which as varieties; and the wild parent-
species are unknown. (10/198. Alph. De Candolle 'Geograph. Bot.' page 983.)
Many plants long cultivated in tropical countries, such as the Banana, have
produced numerous varieties; but as these have never been described with
even moderate care, they are here also passed over. Nevertheless, a
sufficient, and perhaps more than sufficient, number of cases have been
given, so that the reader may be enabled to judge for himself on the nature
and great amount of variation which cultivated plants have undergone.
CHAPTER 1.XI.
ON BUD-VARIATION, AND ON CERTAIN ANOMALOUS MODES OF REPRODUCTION AND
VARIATION.
BUD-VARIATION IN THE PEACH, PLUM, CHERRY, VINE, GOOSEBERRY, CURRANT, AND
BANANA, AS SHOWN BY THE MODIFIED FRUIT.
IN FLOWERS: CAMELLIAS, AZALEAS, CHRYSANTHEMUMS, ROSES, ETC.
ON THE RUNNING OF THE COLOUR IN CARNATIONS.
BUD-VARIATIONS IN LEAVES.
VARIATIONS BY SUCKERS, TUBERS, AND BULBS.
ON THE BREAKING OF TULIPS.
BUD-VARIATIONS GRADUATE INTO CHANGES CONSEQUENT ON CHANGED CONDITIONS OF
LIFE.
GRAFT-HYBRIDS.
ON THE SEGREGATION OF THE PARENTAL CHARACTERS IN SEMINAL HYBRIDS BY BUD-
VARIATION.
ON THE DIRECT OR IMMEDIATE ACTION OF FOREIGN POLLEN ON THE MOTHER-PLANT.
ON THE EFFECTS IN FEMALE ANIMALS OF A PREVIOUS IMPREGNATION ON THE
SUBSEQUENT OFFSPRING.
CONCLUSION AND SUMMARY.
This 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 appearance 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 difference 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 individuals are produced through the formation of
various 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, etc., 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, etc., had been
subjected to a long course of domestication, they would have varied by
buds; for they resemble plants in many respects. For instance, any new or
peculiar character presented by a compound animal is propagated by budding,
as occurs with differently coloured Hydras, and as Mr. Gosse has shown to
be the case with a singular variety of a true coral. Varieties 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 variations which I have
been able to collect, and afterwards show their importance. (11/1. Since
the publication of the first edition of this work, I have found that M.
Carriere, Chef des Pepinieres au Mus. d'Hist. Nat. in his excellent Essay
'Production et Fixation des Varietes' 1865 has given a list of bud-
variations far more extensive than mine; but as these relate chiefly to
cases occurring in France I have left my list as it stood, adding a few
facts from M. Carriere and others. Any one who wishes to study the subject
fully should refer to M. Carriere's Essay.) These cases prove that those
authors who, like Pallas, attribute all variability to the crossing either
of distinct races, or of distinct individuals belonging 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 cases of a peach-almond and a double-
flowered almond which suddenly produced fruit closely resembling true
peaches. I have also given 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 million, 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 remarkable, 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 production 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 sporting branch"
the grosse mignonne tardive, "a most excellent variety," which ripens its
fruit a fortnight later than the parent tree, and is equally good. (11/2.
'Gardener's Chronicle' 1854 page 821.) This same peach has likewise
produced by bud-variation the early grosse mignonne. Hunt's large tawny
nectarine "originated from Hunt's small tawny nectarine, but not through
seminal reproduction." (11/3. Lindley 'Guide to Orchard' as quoted in
'Gardener's Chronicle' 1852 page 821. For the Early mignonne peach see
'Gardener's Chronicle' 1864 page 1251.)
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. (11/4. 'Transact. Hort. Soc.' volume 2 page 160.) Mr.
Rivers, of Sawbridgeworth, informs me (January 1863) that a single 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 the other trees, but were unlike any other
known kind of yellow plum. (11/5. See also 'Gardener's Chronicle' 1863 page
27.)
CHERRY (Prunus cerasus).
Mr. Knight has recorded (ibid) the case of a branch of a May-Duke cherry,
which, though certainly never grafted, always produced fruit, ripening
later, and more oblong 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. (11/6.
'Gardener's Chronicle' 1852 page 821.) M. Carriere gives (page 37) numerous
analogous cases, and one of the same tree bearing three kinds of fruit.
GRAPES (Vitis vinifera).
The black or purple Frontignan in one case produced during two successive
years (and no doubt permanently), spurs which bore white Frontignan grapes.
In another case, on the same footstalk, the lower berries "were well-
coloured black Frontignans; those next the stalk were white, with the
exception 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 produced in the same
cluster. (11/7. 'Gardener's Chronicle' 1852 page 629; 1856 page 648; 1864
page 986. Other cases are given by Braun 'Rejuvenescence' in 'Ray Soc. Bot.
Mem.' 1853 page 314.) Count Odart describes a variety which often bears on
the same stalk small round and large oblong berries; though the shape of
the berry is generally a fixed character. (11/8. 'Ampelographie' etc. 1849
page 71.) Here is another striking case given on the excellent authority of
M. Carriere (11/9. 'Gardener's Chronicle' 1866 page 970.): "a black Hamburg
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 described by Dr. Lindley (11/10. 'Gardener's
Chronicle' 1855 pages 597, 612.) 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 latter 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 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 between red and white, produced during fourteen
years on separate branches and mingled on the same branch, berries of the
red, white, and champagne kinds. (11/11. 'Gardener's Chronicle' 1842 page
873; 1855 page 646. In the 'Chronicle' page 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.) 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.
(11/12. 'Revue Horticole' quoted in 'Gardener's Chronicle' 1844 page 87.)
Alexander Braun (11/13. 'Rejuvenescence in Nature' 'Bot. Memoirs Ray Soc.'
1853 page 314.) also has often seen branches on white currant-trees bearing
red berries.
PEAR (Pyrus communis).
Dureau de la Malle states that the flowers on some trees of an ancient
variety, the doyenne 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 doyenne
blanc, but in shape were like the bon-chretien: it was not ascertained
whether this new variety could be propagated by budding or grafting. The
same author grafted a bon-chretien on a quince, and it produced, besides
its proper fruit, an apparently new variety, of a peculiar form with thick
and rough skin. (11/14. 'Comptes Rendus' tome 41 1855 page 804. The second
case is given on the authority of Gaudichaud ibid tome 34 1852 page 748.)
APPLE (Pyrus malus).
In Canada, a tree of the variety called Pound Sweet, produced (11/15. This
case is given in the 'Gardener's Chronicle' 1867 page 403.), between two of
its proper fruit, an apple which was well russeted, 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. M. Carriere (page 38) mentions an analogous
instance. 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 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 ripening
earlier than the proper fruit. The abnormal fruit closely resembled, except
in being smaller, that of the Musa chinensis or cavendishii, which has
generally been ranked as a distinct species. (11/16. 'Journal of Proc.
Linn. Soc.' volume 2 Botany page 131.)
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. (11/17. 'Gardener's Chronicle' 1847 page 207.) The
Pompon 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."
(11/18. Herbert 'Amaryllidaceae' 1838 page 369.)
Crataegus oxyacantha.
A dark pink hawthorn has been known to throw out a single tuft of pure
white blossoms (11/19. 'Gardener's Chronicle' 1843 page 391.); and Mr. A.
Clapham, nurseryman, of Bedford, 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.
A. indica is well known often to produce new varieties by buds. 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. variegata 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. (11/20.
Exhibited at Hort. Soc. London. Report in 'Gardener's Chronicle' 1844 page
337.)
HIBISCUS (Paritium tricuspis).
A seedling of this plant, when some years old, produced, at Saharunpore
(11/21. Mr. W. Bell 'Bot. Soc. of Edinburgh' May 1863.), 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
considerably 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." Dr. King, who subsequently had charge of
these Gardens, informs me that a tree of Paritium tricuspis (probably the
very same plant) growing there, had a branch buried in the ground,
apparently by accident; and this branch changed its character wonderfully,
growing like a bush, and producing flowers and leaves, resembling in shape
those of another species, viz., P. tiliaceum. A small branch springing from
this bush near the ground, reverted to the parent-form. Both forms were
extensively propagated during several years by cuttings and kept perfectly
true.
Althaea rosea.
A double yellow Hollyhock 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. (11/22.
'Revue Horticole' quoted in 'Gardener's Chronicle' 1845 page 475.)
PELARGONIUM.
These highly cultivated plants seem eminently liable to bud-variation. I
will give only a few well-marked cases. Gartner has seen (11/23.
'Bastarderzeugung' 1849 s. 76.) a plant of P. zonale with a branch having
white edges, 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 (11/24. 'Journal of
Horticulture' 1861 page 336.) 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. (11/25. W.P. Ayres 'Gardener's
Chronicle' 1842 page 791.) 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. (11/26. W.P. Ayres ibid.) Here is a
better case of reversion: a variety produced from a complicated cross,
after having been propagated for five generations by seed, yielded by bud-
variation three very distinct varieties which were undistinguishable from
plants, "known to have been at some time ancestors of the plant in
question." (11/27. Dr. Maxwell Masters 'Pop. Science Review' July 1872 page
250.) 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 Hammersmith, states (11/28. 'Gardener's Chronicle'
1861 page 968.) that he has himself known this purple variety 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 occasionally produce flowers of the original
colour. This year coccineum 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 (11/29. Ibid 1861 page
945.); but one at least of these did not arise through bud-variation, but
is believed to be a seedling from Rollisson's Unique. (11/30. W. Paul
'Gardener's Chronicle' 1861 page 968.) There are, also, in the trade
(11/31. Ibid page 945.) two other slightly different varieties, of unknown
origin, of Rollisson's Unique: so that altogether we have a curiously
complex case of variation both by buds and seeds. (11/32. For other cases
of bud-variation in this same variety see 'Gardener's Chronicle' 1861 pages
578, 600, 925. For other distinct cases of bud-variation in the genus
Pelargonium see 'Cottage Gardener' 1860 page 194.) Here is a still more
complex case: M. Rafarin states that a pale rose-coloured variety produced
a branch bearing deep red flowers. "Cuttings were taken from this 'sport,'
from which 20 plants were raised, which flowered in 1867, when it was found
that scarcely two were alike." Some resembled the parent-form, some
resembled the sport, some bore both kinds of flowers; and even some of the
petals on the same flower were rose-coloured and others red. (11/33. Dr.
Maxwell Masters 'Pop. Science Review' July 1872 page 254.) 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. (11/34. Rev. W.T. Bree in Loudon's 'Gardener's Mag.' volume 8 1832
page 93.)
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. (11/35. 'The Chrysanthemum: its History and Culture'
by J. Salter 1865 page 41 etc.) A variety called cedo nulli bears small
yellow flowers, but habitually produces branches with white flowers; and a
specimen was exhibited, which Prof. T. Dyer saw, before the Horticultural
Society. The varieties which were first introduced from China were so
excessively 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 fluctuating 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 different 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 informs 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 chrysanthemums, which have all produced branches with
yellow flowers; but 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 cultivated in a
garden, not unfrequently produces on the same root flowers of four
different colours, viz., blue, white, dark-purple, and parti-coloured.
(11/36. Bree in Loudon's 'Gardener's Mag.' volume 8 1832 page 93.) The
flowers of Anthemis also vary on the same plant. (11/37. Bronn 'Geschichte
der Natur' b. 2 s. 123.)
ROSES.
Many varieties of the Rose are known or are believed to have originated by
bud-variation. (11/38. T. Rivers 'Rose Amateur's Guide' 1837 page 4.) The
common double moss-rose was imported into England from Italy about the year
1735. (11/39. Mr. Shailer quoted in 'Gardener's Chronicle' 1848 page 759.)
Its origin is unknown, but from analogy it probably arose from the Provence
rose (R. centifolia) by bud-variation; for the branches of the common moss-
rose have several times been known to produce Provence roses, wholly or
partially destitute of moss: I have seen one such instance, and several
others have been recorded. (11/40. 'Transact. Hort. Soc.' volume 4 1822
page 137; 'Gardener's Chronicle' 1842 page 422.) Mr. Rivers also informs me
that he raised two or three roses of the Provence class from seed of the
old single moss-rose (11/41. See also Loudon's 'Arboretum' volume 2 page
780.); 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." (11/42. 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
propagation. See 'Gard. Chronicle' 1852 page 759.) A white moss-rose has
been seen to bear a flower half white and half pink. (11/43. 'Gardener's
Chronicle' 1845 page 564.) 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-variation of the double
moss-rose originally imported from Italy. That the original moss-rose was
the product of bud-variation is probable, from the facts above given and
from the de Meaux moss-rose (also a variety of R. centifolia) (11/44.
'Transact. Hort. Soc.' volume 2 page 242.) having appeared as a sporting
branch on the common rose de Meaux. Prof. Caspary has carefully described
(11/45. 'Shriften der Phys. Oekon. Gesell. zu Konigsberg' February 3, 1865
s. 4. See also Dr. Caspary's paper in 'Transactions of the Hort. Congress
of Amsterdam' 1865.) 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 originated in this way. (11/46. 'Gardener's
Chronicle' 1852 page 759.) M. Carriere states (page 36) that he himself
knows of five varieties thus produced by the Baronne Prevost. The double
and highly-coloured Belladonna rose has produced by suckers both semi-
double and almost single white roses (11/47. 'Transact. Hort. Soc.' volume
2 page 242.); whilst suckers from one of these semi-double white roses
reverted to perfectly characterised Belladonnas. In St. Domingo, varieties
of the China rose propagated by cuttings often revert after a year or two
into the old China rose. (11/48. Sir R. Schomburgk 'Proc. Linn. Soc. Bot.'
volume 2 page 132.) Many cases have been recorded of roses suddenly
becoming striped or changing their character by segments: some plants of
the Comtesse de Chabrillant, which is properly rose-coloured, were
exhibited in 1862 (11/49. 'Gardener's Chronicle' 1862 page 619.), 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.
lutea) not rarely (11/50. Hopkirk 'Flora Anomala' 167.) 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 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
(11/51. 'Sur La Production et la Fixation des Varietes' 1865 page 4.), a
variety called Rosa cannabifolia, which has peculiarly shaped leaflets, and
differs 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 observed by Mr. H.
Curtis (11/52. 'Journal of Horticulture' March 1865 page 233.) on the old
Aimee Vibert Noisette, and he budded it on Celine; thus a climbing Aimee
Vibert was first produced and afterwards propagated.
DIANTHUS.
It is quite common with the Sweet William (D. barbatus) 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, etc.) occasionally vary by layers; and some kinds are so
little certain in character that they are called by floriculturists "catch-
flowers." (11/53. 'Gardener's Chronicle' 1843 page 135.) 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." (11/54. Ibid 1842 page 55.) 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; and many
others 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 variety. 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
purple 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 (11/55.
'Gardener's Chronicle' 1867 page 235.) has been observed to bear white and
pink flowers of two forms, the one resembling the Persicum strain, and the
other the Coum strain. Oenothera biennis has been seen (11/56. Gartner
'Bastarderzeugung' s. 305.) bearing flowers of three different colours. The
hybrid Gladiolus colvilii occasionally bears uniformly coloured flowers,
and one case is recorded (11/57. Mr. D. Beaton in 'Cottage Gardener' 1860
page 250.) of all the flowers on a plant thus changing colour. A Fuchsia
has been seen (11/58. 'Gardener's Chronicle' 1850 page 536.) bearing two
kinds of flowers. Mirabilis jalapa is eminently sportive, sometimes bearing
on the same root pure red, yellow, and white flowers, and others striped
with various combinations of these three colours. (11/59. Braun 'Ray Soc.
Bot. Mem.' 1853 page 315; Hopkirk 'Flora Anomala' page 164; Lecoq
'Geograph. Bot. de l'Europe' tome 3 1854 page 405; and 'De la Fecondation'
1862 page 303.) 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; incidentally some remarkable modifications in the leaves and
shoots of the rose and Paritium, and in a lesser degree in the foliage of
the Pelargonium and Chrysanthemum, have been noticed. I will now add a few
more cases of variation in leaf-buds. Verlot (11/60. 'Des Varietes' 1865
page 5.) states that on Aralia trifoliata, which properly has leaves with
three leaflets, branches frequently appear bearing simple leaves of various
forms; these can be propagated by buds or by grafting, and have given rise,
as he states, to several 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. (11/61. W. Mason in 'Gardener's Chronicle' 1843 page
878.) The varieties of some trees with cut leaves, as the oak-leaved
laburnum, the parsley-leaved vine, and especially the fern-leaved beech,
are apt to revert by buds to the common forms. (11/62. Alex. Braun 'Ray
Soc. Bot. Mem.' 1853 page 315; 'Gardener's Chron.' 1841 page 329.) The
fern-like leaves of the beech sometimes revert only partially, and the
branches display here and there sprouts bearing common leaves, fern-like,
and variously shaped leaves. Such cases differ but little from the so-
called heterophyllus varieties, in which the tree habitually bears 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. (11/63. Dr. M.T. Masters 'Royal
Institution Lecture' March 16, 1860.)
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 exposure 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
often display remarkable deviations of structure. Spores, which are of the
nature of buds, taken from such abnormal fronds, reproduce, with remarkable
fidelity, the same variety, after passing through the sexual stage. (11/64.
See Mr. W.K. Bridgeman's curious paper in 'Annals and Mag. of Nat. Hist.'
December 1861; also Mr. J. Scott 'Bot. Soc. Edinburgh' June 12, 1862.)
With respect to colour, leaves often become by bud-variation zoned,
blotched, or spotted with white, yellow, and red; and this occasionally
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. (11/65. 'Journal of Horticulture' 1861
page 336; Verlot 'Des Varietes' page 76.) There have been endless disputes
whether variegation should be considered as a disease. In a future chapter
we shall see that it is much influenced, both in the case of seedlings and
of mature plants, by the nature of the soil. Plants which have become
variegated as seedlings, generally transmit their character by seed to a
large proportion of their progeny; and Mr. Salter has given me a list of
eight genera in which this occurred. (11/66. See also Verlot 'Des Varietes'
page 74.) Sir F. Pollock has given me more precise information: he sowed
seed from a variegated plant of Ballota 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
variegated by bud-variation, and the variety is attempted to be propagated
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 worth keeping. Variegated
plants, whether originally produced from seeds or buds, can generally be
propagated by budding, grafting, etc.; 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." (11/67.
'Gardener's Chron.' 1844 page 86.) 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 generally 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 stature
still remains. (11/68. Ibid 1861 page 963.) It is remarkable that plants
propagated from branches which have reverted from variegated to plain
leaves (11/69. Ibid 1861 page 433; 'Cottage Gardener' 1860 page 2.) 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 hitherto 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 variegated 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 safely propagated (11/70. M. Lemoine quoted in
'Gardener's Chronicle' 1867 page 74 has lately observed that the Symphytum
with variegated leaves cannot be propagated by division of the roots. He
also found that out of 500 plants of a Phlox with striped flowers, which
had been propagated by root-division, only seven or eight produced striped
flowers. See also on striped Pelargoniums 'Gardener's Chronicle' 1867 page
1000.); but this latter plant may have originated as a variegated seedling,
which would account for its greater fixedness of 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. (11/71. Anderson 'Recreations in Agriculture' volume 5 page 152.) My
father repeatedly tried this experiment, and always with the same result. I
may here mention that maize and wheat sometimes produce new varieties from
the stock or root, as does the sugar-cane. (11/72. For wheat see
'Improvement of the Cereals' by P. Shirreff 1873 page 47. For maize and
sugar-cane Carriere ibid pages 40, 42. With respect to the sugar-cane Mr.
J. Caldwell of Mauritius says ('Gardener's Chronicle' 1874 page 316) the
Ribbon cane has here "sported into a perfectly green cane and a perfectly
red cane from the same head. I verified this myself, and saw at least 200
instances in the same plantation, and the fact has completely upset all our
preconceived ideas of the difference of colour being permanent. The
conversion of a striped cane into a green cane was not uncommon, but the
change into a red cane universally disbelieved, and that both events should
occur in the same plant incredible. I find, however, in Fleischman's
'Report on Sugar Cultivation in Louisiana for 1848 by the American Patent
Office, the circumstance is mentioned, but he says he never saw it
himself.")
Turning now to tubers: in the common Potato (Solanum tuberosum) a single
bud or eye sometimes varies and produces a new variety; or, occasionally,
and this is a much more remarkable circumstance, 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 observed (11/73.
'Gardener's Chronicle' 1857 page 662.) 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 Lancashire 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. (11/74. 'Gard. Chronicle' 1841 page 814.)
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. (11/75. Ibid 1857 page 613.) Several cases
have been recorded of large portions of whole rows of potatoes slightly
changing their character. (11/76. Ibid 1857 page 679. See also Philips
'Hist. of Vegetables' volume 2 page 91 for other and similar accounts.)
Dahlias propagated by tubers under the hot climate of St. Domingo vary
much; Sir R. Schomburgk gives the case of the "Butterfly 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." (11/77.
'Journal of Proc. Linn. Soc.' volume 2 Botany page 132.) 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
intermixed." (11/78. Loudon's 'Gardener's Mag.' volume 8 1832 page 94.)
Another case is described of a dahlia with purple flowers which bore a
white flower streaked with purple. (11/79. 'Gard. Chronicle' 1850 page 536;
and 1842 page 729.)
Considering how long and extensively many Bulbous plants have been
cultivated, and how numerous are the varieties produced from seed, these
plants have not perhaps varied so much by offsets,--that is, by the
production of new bulbs,--as might have been expected. With the Hyacinth,
however, several instances have been given by M. Carriere. A case also has
been recorded of a blue variety which for three successive years gave
offsets producing white flowers with a red centre. (11/80. 'Des Jacinthes'
etc. Amsterdam 1768 page 122.) Another hyacinth bore (11/81. 'Gardener's
Chronicle' 1845 page 212.) on the same truss a perfectly pink and a
perfectly blue flower. I have seen a bulb producing at the same time one
stalk or truss with fine blue flowers, another with fine red flowers, and a
third with blue flowers on one side and red on the other; several of the
flowers being also longitudinally striped red and blue.
Mr. John Scott informs me that in 1862 Imatophyllum miniatum, in the
Botanic Gardens of Edinburgh, threw up a sucker which differed from the
normal form, in the leaves being two-ranked instead of four-ranked. The
leaves were also smaller, with the upper surface 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." (11/82. Loudon's 'Encyclopaedia of Gardening' page
1024.) The 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 seedling. This bud-variation, in
accordance with the views of MM. Vilmorin and Verlot (11/83. 'Production
des Varietes' 1865 p. 63.) is probably an attempt to revert to that uniform
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 (11/84. 'Gardener's Chronicle' 1841 page 782; 1842 page
55.) that this can no more be accounted for than the variation of any other
plant. He believes that English growers, from care in choosing 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. Iris xiphium, according to M. Carriere (page 65), behaves in
nearly the same manner, as do so many tulips.
During two consecutive years all the early flowers in a bed of Tigridia
conchiflora (11/85. 'Gardener's Chron.' 1849 page 565.) 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 (11/86. 'Transact. Lin. Soc.' volume 2 page 354.) of two forms of
Hemerocallis, which have been universally considered 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 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 bud-variation, that the change is
effected through buds and not through seminal reproduction. But, on the
other hand, there is this difference--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. With the potato, we have
seen an intermediate case, for all the eyes in one tuber 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 conditions of life. When
the common Hepatica is transplanted from its native woods, the flowers
change colour, even during the first year. (11/87. Godron 'De l'Espece'
tome 2 page 84.) 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 produced 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 (11/88. M. Carriere has
lately described in the 'Revue Horticole' December 1, 1866 page 457, an
extraordinary case. He twice inserted grafts of the Aria vestita on thorn-
trees (epines) growing in pots; and the grafts, as they grew, produced
shoots with bark, buds, leaves, petioles, petals, and flower-stalks, all
widely different from those of the Aria. The grafted shoots were also much
hardier, and flowered earlier, than those on the ungrafted Aria.) were
considered by Andrew Knight (11/89. 'Transact. Hort. Soc.' volume 2 page
160.) 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; seedling 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 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, whilst they
closely resemble each other in old age; and as with certain oaks, and with
some varieties of the lime and hawthorn. (11/90. For the cases of oaks see
Alph. De Candolle in 'Bibl. Univers.' Geneva November 1862; for limes etc.
Loudon's 'Gard. Mag.' volume 11 1835 page 503.)]
GRAFT-HYBRIDS.
Before giving a summary on Bud-variation I will discuss 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 the two
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 divided into halves, one side being
bright yellow and the other purple; 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 narrow
bright yellow 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. (11/91. For analogous facts see Braun
'Rejuvenescence' in 'Ray Soc. Bot. Mem.' 1853 page 320; and 'Gardener's
Chronicle' 1842 page 397; also Braun in 'Sitzungsberichte der Ges.
naturforschender Freunde' June 1873 page 63.) The most remarkable 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 complement of seed; they certainly yield a
larger number. Two seedlings raised by Mr. Herbert from such seed (11/92.
'Journal of Hort. Soc.' volume 2 1847 page 100.) 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 exception 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 hybridised and sterile a form is an astonishing
phenomenon. The branches with purple flowers appear at first sight exactly
to resemble those of C. 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
plainly 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
singular 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
grains. Although the pollen of the reverted purple flowers was in so poor a
condition, the ovules were well formed, and the seeds, when mature,
germinated freely with me. Mr. Herbert raised plants from seeds of the
reverted purple flowers, and they differed a VERY LITTLE from the usual
state of C. purpureus. Some which I raised in the same manner did not
differ at all, either in the character of their flowers or of the whole
bush, from the pure C. purpureus. Prof. Caspary has examined the ovules of
the dingy-red and sterile flowers in several plants of C. adami on the
Continent (11/93. See 'Transact. of Hort. Congress of Amsterdam' 1865; but
I owe most of the following information to Prof. Caspary's letters.) and
finds them generally monstrous. In three plants examined by me in England,
the ovules were likewise monstrous, the nucleus varying much 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 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 observation (11/94. 'Nouvelles Archives du Museum' tome 1 page
143.) on Mirabilis, that it would be functionally effective. The fact of
the ovules of C. adami being monstrous, and the pollen apparently sound, is
all the more remarkable, because it is opposed to what usually occurs not
only with most hybrids (11/95. See on this head Naudin ibid page 141.), but
with two hybrids in the same genus, namely in C. purpureo-elongatus, and C.
alpino-laburnum. In both these hybrids, 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 elements 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 Moggridge) that with hybrid Cisti the ovarium is frequently
deformed, the ovules being in some cases quite absent, and in other cases
incapable of fertilisation.
Several theories have been propounded to account for the origin of C.
adami, and for the transformations which it undergoes. The whole case has
been attributed by some authors to 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 embryos differing in their characters may be developed within the same
seed and cohere; and it has been supposed that C. adami thus originated.
Many botanists 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 not of much value; but Reisseck, Caspary, and 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. Nevertheless, the belief that C. adami is a
spontaneously produced hybrid between these two species is supported by the
fact that such hybrids have arisen in this genus. In a bed of seedlings
from C. elongatus, which grew near to C. purpureus, and was probably
fertilised by it through the agency of insects (for these, as I know by
experiment, play an important part in the fertilisation of the laburnum),
the sterile hybrid C. purpureo-elongatus appeared. (11/96. Braun in 'Bot.
Mem. Ray. Soc.' 1853 page 23.) Thus, also, Waterer's laburnum, the C.
alpino-laburnum (11/97. This hybrid has never been described. It is exactly
intermediate in foliage, time of flowering, dark striae 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-formed
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
occasionally even four pods. Many of the pods contained no good seeds, but
generally they contained a single apparently good seed, sometimes two, and
in one case three seeds. Some of these seeds germinated, and I raised two
trees from them; one resembles the present form; the other has a remarkable
dwarf character with small leaves, but has not yet flowered.) spontaneously
appeared, as I am informed by Mr. Waterer, in a bed of seedlings.
On the other hand, we have a clear and distinct account given to Poiteau
(11/98. 'Annales de la Soc. de l'Hort. de Paris' tome 7 1830 page 93.) by
M. Adam, who raised the plant, showing that C. adami is not an ordinary
hybrid; but is what may be called a graft-hybrid, that is, one produced
from the united cellular tissue of two distinct species. 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 Poiteau after the plants had
flowered, but before they had exhibited their remarkable tendency to revert
into the two parent species. So that there was no conceivable motive for
falsification, and it is difficult to see how there could have been any
error. (11/99. An account was given in the 'Gardener's Chronicle' 1857
pages 382, 400, of a common laburnum on which grafts of C. purpureus had
been inserted, and which gradually assumed the character of C. adami; but I
have little doubt that C. adami had been sold to the purchaser, who was not
a botanist, in the place of C. purpureus. I have ascertained that this
occurred in another instance.) 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 subsequently produce a plant bearing leaves and
sterile flowers intermediate 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 reproduction.]
I will therefore give all the facts which I have been able to collect on
the formation of hybrids between distinct species or varieties, without the
intervention of the sexual organs. For if, as I am now convinced, this is
possible, it is a most important fact, which will sooner or later change
the views held by physiologists with respect to sexual reproduction. A
sufficient body of facts will afterwards be adduced, showing that the
segregation or separation of the characters of the two parent-forms by bud-
variation, as in the case of Cytisus adami, is not an unusual though a
striking phenomenon. We shall further see that a whole bud may thus revert,
or only half, or some smaller segment.
[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 describer, P. Nato (11/100. Gallesio 'Gli
Agrumi dei Giard. Bot. Agrar. di. Firenze' 1839 page 11. In his 'Traite du
Citrus' 1811 page 146, he speaks as if the compound fruit consisted in part
of a lemon, but this apparently was a mistake.), 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 cuttings, and
retains its diversified character. The so-called trifacial orange of
Alexandria and Smyrna (11/101. 'Gardener's Chron.' 1855 page 628. See also
Prof. Caspary in 'Transact. Hort. Congress of Amsterdam' 1865.) resembles
in its general nature the bizzarria, and differs only in the orange being
of the sweet kind; this and the citron are blended together in the same
fruit, or are separately produced on the same tree; nothing is known of its
origin. In regard to the bizzarria, 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 that the species in this genus often cross
spontaneously.
It is notorious that when the variegated Jessamine is budded on the common
kind, the stock sometimes produces buds bearing variegated leaves: Mr.
Rivers, as he informs me, has seen instances of this. The same thing occurs
with the Oleander. (11/102. Gartner 'Bastarderzeugung' s. 611 gives many
references on this subject.) 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 (11/103. A nearly similar account was given by Brabley in 1724 in
his 'Treatise on Husbandry' volume 1 page 199.) 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 propagated, and has preserved its character during the
last fifty years. Weeping ashes, also, were budded on the affected stocks,
and became similarly variegated. It has been repeatedly proved that several
species of Abutilon, on which the variegated A. thompsonii has been
grafted, become variegated. (11/104. Morren 'Bull. de l'Acad. R. des
Sciences de Belgique' 2de series tome 28 1869 page 434. Also Magnus
'Gesellschaft naturforschender Freunde, Berlin' February 21, 1871 page 13;
ibid June 21, 1870 and October 17, 1871. Also 'Bot. Zeitung' February 24,
1871.)
Many authors consider variegation as the result of disease; and the
foregoing cases may be looked at as the direct result of the inoculation of
a disease or some weakness. This has been almost proved to be the case by
Morren in the excellent paper just referred to, who shows that even a leaf
inserted by its footstalk into the bark of the stock is sufficient to
communicate variegation to it, though the leaf soon perishes. Even fully
formed leaves on the stock of Abutilon are sometimes affected by the graft
and become variegated. 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 disease, 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 (11/105. Loudon's 'Arboretum' volume 4 page 2595.), it sometimes
colours, as has been asserted, the leaves below the graft; although
negative evidence is not of much value, I may add that Mr. Rivers, who has
possessed hundreds of such grafted trees, has never seen an instance.
Gartner (11/106. 'Bastarderzeugung' s. 619.) 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, etc.;
and these branches produced distinct bunches of grapes of the two colours,
and other bunches with berries, 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 fruits and leaves, by
fertilising a white variety by the already variegated dark Aleppo grape.
Gartner 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 thus have varied; and H. Adorne de Tscharner
positively asserts 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' (11/107. Amsterdam 1768 page 124.) impressed me with the belief
not only of his extensive knowledge, but of his truthfulness: he says that
bulbs 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 analogous with that of the blended
colours of the grapes on the united vine branches.
In the case of roses it is supposed that several graft-hybrids have been
formed, but there is much doubt about these cases, owing to the frequency
of ordinary bud-variations. The most trustworthy instance known to me is
one, recorded by Mr. Poynter (11/108. 'Gardener's Chron.' 1860 page 672
with a woodcut.) 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 Devoniensis 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 certainly been produced by the mingling of R.
banksiae with some rose like R. devoniensis, "for while it was very greatly
increased in vigour and in size of all the parts, the leaves were half-way
between a Banksian and Tea-scented rose." It appears that rose-growers were
previously aware that the Banksian rose sometimes affects other roses. As
Mr. Poynter's new variety is intermediate in its fruit and foliage between
the stock and scion, and as it arose from the point of junction between the
two, it is very improbable that it owes its origin to mere bud-variation,
independently of the mutual influence of the stock and scion.
Lastly, with respect to potatoes. Mr. R. Trail stated in 1867 before the
Botanical Society of Edinburgh (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; 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, that is, by graft-
hybridisation.
In the 'Botanische Zeitung' (May 16, 1868), Professor Hildebrand gives an
account with a coloured figure, of his experiments on two varieties which
were found during the same season to be constant in character, namely, a
somewhat elongated rough-skinned red potato and a rounded smooth white one.
He inserted buds reciprocally into both kinds, destroying the other buds.
He thus raised two plants, and each of these produced a tuber intermediate
in character between the two parent-forms. That from the red bud grafted
into the white tuber, was at one end red and rough, as the whole tuber
ought to have been if not affected; in the middle it was smooth with red
stripes, and at the other end smooth and altogether white like that of the
stock.
Mr. Taylor, who had received several accounts of potatoes having been
grafted by wedge-shaped pieces of one variety inserted into another, though
sceptical on the subject, made twenty-four experiments which he described
in detail before the Horticultural Society. (11/109. 'Gardener's Chronicle'
1869 page 220.) He thus raised many new varieties, some like the graft or
like the stock; others having an intermediate character. Several persons
witnessed the digging up of the tubers from these graft-hybrids; and one of
them, Mr. Jameson, a large dealer in potatoes, writes thus, "They were such
a mixed lot, as I have never before or since seen. They were of all colours
and shapes, some very ugly and some very handsome." Another witness says
"some were round, some kidney, pink-eyed kidney, piebald, and mottled red
and purple, of all shapes and sizes." Some of these varieties have been
found valuable, and have been extensively propagated. Mr. Jameson took away
a large piebald potato which he cut into five sets and propagated; these
yielded round, white, red, and piebald potatoes.
Mr. Fitzpatrick followed a different plan (11/110. 'Gard. Chronicle' 1869
page 335.); he grafted together not the tubers but the young stems of
varieties producing black, white, and red potatoes. The tubers borne by
three of these twin or united plants were coloured in an extraordinary
manner; one was almost exactly half black and half white, so that some
persons on seeing it thought that two potatoes had been divided and
rejoined; other tubers were half red and half white, or curiously mottled
with red and white, or with red and black, according to the colours of the
graft and stock.
The testimony of Mr. Fenn is of much value, as he is "a well known potato-
grower" who has raised many new varieties by crossing different kinds in
the ordinary manner. He considers it "demonstrated" that new, intermediate
varieties can be produced by grafting the tubers, though he doubts whether
such will prove valuable. (11/111. 'Gardener's Chronicle' 1869 page 1018
with remarks by Dr. Masters on the adhesion of the united wedges. See also
ibid 1870 pages 1277, 1283.) He made many trials and laid the results,
exhibiting specimens, before the Horticultural Society. Not only were the
tubers affected, some being smooth and white at one end and rough and red
at the other, but the stems and leaves were modified in their manner of
growth, colour and precocity. Some of these graft-hybrids after being
propagated for three years still showed in their haulms their new
character, different from that of the kind from which the eyes had been
taken. Mr. Fenn gave twelve of the tubers of the third generation to Mr.
Alex. Dean, who grew them, and was thus converted into a believer in graft-
hybridisation, having previously been a complete sceptic. For comparison he
planted the pure parent-forms alongside the twelve tubers; and found that
many of the plants from the latter (11/112. 'Gard. Chronicle' 1871 page
837.) were intermediate between the two parent-forms in precocity, in the
tallness, uprightness, jointing, and robustness of the stems, and in the
size and colour of the leaves.
Another experimentalist, Mr. Rintoul, grafted no less than fifty-nine
tubers, which differed in shape (some being kidneys) in smoothness and
colour (11/113. 'Gardener's Chronicle' 1870 page 1506.) and many of the
plants thus raised "were intermediate in the tubers as well as in the
haulms." He describes the more striking cases.
In 1871 I received a letter from Mr. Merrick, of Boston, U.S.A., who states
that, "Mr. Fearing Burr, a very careful experimenter and author of a much
valued book, 'The Garden Vegetables of America' has succeeded in producing
distinctly mottled and most curious potatoes--evidently graft-hybrids, by
inserting eyes from blue or red potatoes into the substance of white ones,
after removing the eyes of the latter. I have seen the potatoes, and they
are very curious."
We will now turn to the experiments made in Germany, since the publication
of Prof. Hildebrand's paper. Herr Magnus relates (11/114. 'Sitzungsberichte
der Gesellschaft naturforschender Freunde zu Berlin' October 17, 1871.) the
results of numerous trials made by Herren Reuter and Lindemuth, both
attached to the Royal Gardens of Berlin. They inserted the eyes of red
potatoes into white ones, and vice versa. Many different forms partaking of
the characters of the inserted bud and of the stock were thus obtained; for
instance, some of the tubers were white with red eyes.
Herr Magnus also exhibited in the following year before the same Society
(November 19, 1872), the produce of grafts between black, white, and red
potatoes, made by Dr. Neubert. These were made by uniting not the tubers
but the young stems, as was done by Mr. Fitzpatrick. The result was
remarkable, inasmuch as all the tubers thus produced were intermediate in
character, though in a variable degree. Those between the black and the
white or the red were the most striking in appearance. Some from between
the white and red had one half of one colour and the other half of the
other colour.
At the next meeting of the society Herr Magnus communicated the results of
Dr. Heimann's experiments in grafting together the tubers of red Saxon,
blue, and elongated white potatoes. The eyes were removed by a cylindrical
instrument, and inserted into corresponding holes in other varieties. The
plants thus produced yielded a great number of tubers, which were
intermediate between the two parent-forms in shape, and in the colour both
of the flesh and skin.
Herr Reuter experimented (11/115. Ibid November 17, 1874. See also
excellent remarks by Herr Magnus.) by inserting wedges of the elongated
White Mexican potato into a Black Kidney potato. Both sorts are known to be
very constant, and differ much not only in form and colour, but in the eyes
of the Black Kidney being deeply sunk, whereas those of the White Mexican
are superficial and of a different shape. The tubers produced by these
hybrids were intermediate in colour and form; and some which resembled in
form the graft, i.e. the Mexican, had eyes deeply sunk and of the same
shape as in the stock or Black Kidney.]
Any one who will attentively consider the abstract now given, of the
experiments made by many observers in several countries, will, I think, be
convinced that by grafting two varieties of the potato together in various
ways, hybridised plants can be produced. It should be observed that several
of the experimentalists are scientific horticulturists, and some of them
potato-growers on a large scale, who, though beforehand sceptical, have
been fully convinced of the possibility, even of the ease, of making graft-
hybrids. The only way of escaping from this conclusion is to attribute all
the many recorded cases to simple bud-variation. Undoubtedly the potato, as
we have seen in this chapter, does sometimes, though not often, vary by
buds; but it should be especially noted that it is experienced potato-
growers, whose business it is to look out for new varieties, who have
expressed unbounded astonishment at the number of new forms produced by
graft-hybridisation. It may be argued that it is merely the operation of
grafting, and not the union of two kinds, which causes so extraordinary an
amount of bud-variation; but this objection is at once answered by the fact
that potatoes are habitually propagated by the tubers being cut into
pieces, and the sole difference in the case of graft-hybrids is that either
a half or a smaller segment or a cylinder is placed in close opposition
with the tissue of another variety. Moreover, in two cases, the young stems
were grafted together, and the plants thus united yielded the same results
as when the tubers were united. It is an argument of the greatest weight
that when varieties are produced by simple bud-variation, they frequently
present quite new characters; whereas in all the numerous cases above
given, as Herr Magnus likewise insists, the graft-hybrids are intermediate
in character between the two forms employed. That such a result should
follow if the one kind did not affect the other is incredible.
Characters of all kinds are affected by graft hybridisation, in whatever
way the grafting may have been effected. The plants thus raised yield
tubers which partake of the widely different colours, form, state of
surface, position and shape of the eye of the parents; and according to two
careful observers they are also intermediate in certain constitutional
peculiarities. But we should bear in mind that in all the varieties of the
potato, the tubers differ much more than any other part.
The potato affords the best evidence of the possibility of the formation of
graft-hybrids, but we must not overlook the account given of the origin of
the famous Cytisus adami by M. Adam, who had no conceivable motive for
deception, and the exactly parallel account of the origin of the Bizzarria
orange, namely by graft-hybridisation. Nor must the cases be undervalued in
which different varieties or species of vines, hyacinths and roses, have
been grafted together, and have yielded intermediate forms. It is evident
that graft-hybrids can be made much more easily with some plants, as the
potato, than with others, for instance our common fruit trees; for these
latter have been grafted by the million during many centuries, and though
the graft is often slightly affected, it is very doubtful whether this may
not be accounted for, merely by a more or less free supply of nutriment.
Nevertheless, the cases above given seem to me to prove that under certain
unknown conditions graft-hybridisation can be effected.
Herr Magnus asserts with much truth that graft-hybrids resemble in all
respects seminal hybrids, including their great diversity of character.
There is, however, a partial exception, inasmuch as the characters of the
two parent forms are not often homogeneously blended together in graft-
hybrids. They much more commonly appear in a segregated condition,--that
is, in segments either at first, or subsequently through reversion. It
would seem that the reproductive elements are not so completely blended by
grafting as by sexual generation. But segregation of this kind occurs by no
means rarely, as will be immediately shown, in seminal hybrids. Finally it
must, I think, be admitted that we learn from the foregoing cases a highly
important physiological fact, namely, that the elements that go to the
production of a new being, are not necessarily formed by the male and
female organs. They are present in the cellular tissue in such a state that
they can unite without the aid of the sexual organs, and thus give rise to
a new bud partaking of the characters of the two parent-forms.
ON THE SEGREGATION OF THE PARENTAL CHARACTERS IN SEMINAL HYBRIDS BY BUD-
VARIATION.
I will now give a sufficient number of cases to show that segregation of
this kind, namely, by buds, may occur in ordinary hybrids raised from seed.
[Hybrids were raised by Gartner between Tropaeolum minus and majus (11/116.
'Bastarderzeugung' s. 549. It is, however, doubtful whether these plants
should be ranked as species or varieties.) which at first produced flowers
intermediate in size, colour, and structure between their two parents; but
later in the season some of these plants produced flowers in all respects
like those of the mother-form, mingled with flowers still retaining the
usual intermediate condition. A hybrid Cereus between C. speciosissimus and
phyllanthus (11/117. Gartner ibid s. 550.) 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.
Kolreuter 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. (11/118. 'Journal de Physique' tome 23 1873 page 100.
'Act. Acad. St. Petersburgh' 1781 part 1 page 249.) Naudin (11/119.
'Nouvelles Archives du Museum' tome 1 page 49.) raised forty hybrids from
Datura laevis 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. laevis, the
remaining part being spinose and of larger size, like the capsule of the
pure D. stramonium: from one of these composite capsules, plants perfectly
resembling both parent-forms were raised.
Turning now to varieties. A SEEDLING apple, conjectured to be of crossed
parentage, has been described in France (11/120. L'Hermes January 14, 1837
quoted in Loudon's 'Gardener's Mag.' volume 13 page 230.) 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 as that which Gaudichaud (11/121. 'Comptes Rendus'
tome 34 1852 page 746.) 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 jaunatre: 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 foot-stalk, is green, the other
half being brown and of the nature 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 branches 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
class of graft-hybrids already given. But the branch may have sprung from
the stock, which no doubt was a seedling.
Prof. H. Lecoq, who has made a great number of crossings between the
differently coloured varieties of Mirabilis jalapa (11/122. 'Geograph. Bot.
de l'Europe' tome 3 1854 page 405; and 'De la Fecondation' 1862 page 302.)
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 (11/123. 'Traite du Citrus' 1811 page 45.) crossed
reciprocally white and red carnations, 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 following 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 fertilised the
PURPLE SWEET-PEA (Lathyrus odoratus) with pollen from the light-coloured
PAINTED LADY: seedlings raised from the same pod were not intermediate in
character, but perfectly resembled either parent. Later in the summer, the
plants which had at first borne flowers identical with those of the PAINTED
LADY, produced flowers streaked and blotched with purple; showing in these
darker marks a tendency to reversion to the mother-variety. Andrew Knight
(11/124. 'Transact. Linn. Soc.' volume 9 page 268.) 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 foot-stalk.
I will append a very curious case, not of bud-variation, but of two
cohering embryos, different in character and contained within the same
seed. A distinguished botanist, Mr. G.H. Thwaites (11/125. 'Annals and Mag.
of Nat. Hist.' March 1848.) states that a seed from Fuchsia coccinea
fertilised by F. fulgens, contained two embryos, and was "a true vegetable
twin." The two plants produced from the two embryos were "extremely
different in appearance and character," though both resembled 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. A mongrel melon described by Sageret (11/126. 'Pomologie
Physiolog.' 1830 page 126.) may perhaps have thus originated; for the two
main branches, which arose from two cotyledon-buds, produced very different
fruit,--on the one branch like that of the paternal variety, and on the
other branch like to a certain extent that of the maternal variety, the
melon of China.]
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 different 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. When,
however, the later flowers or fruit produced during the same season, or
during a succeeding year or generation, become striped or half-and-half,
etc., the segregation of the two colours is strictly a case of reversion by
bud-variation. Whether all the many recorded cases of striped flowers and
fruit are due to previous hybridisation and reversion is by no means clear,
for instance with peaches and nectarines, moss-roses, etc. 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. Finally,
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.
ON THE DIRECT OR IMMEDIATE ACTION OF THE MALE ELEMENT ON THE MOTHER FORM.
Another remarkable class of facts must be here considered, firstly, because
they have a high physiological importance, and secondly, because they have
been supposed to account for some cases of bud-variation. I refer to the
direct action of the male element, not in the ordinary way on the ovules,
but on certain parts of the female plant, or in 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 have been
affected by the pollen of a foreign variety or species, although the
development of the embryo, inside the embryonic sack, inside the ovule and
ovarium, of course, depends on the male element.
[Even as long ago as 1729 it was observed (11/127. 'Philosophical
Transact.' volume 43 1744-45 page 525.) that white and blue varieties of
the Pea, when planted near each other, mutually crossed, 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. (11/128. Mr. Goss 'Transact. Hort. Soc.' volume 5 page 234: and
Gartner 'Bastarderzeugung' 1849 ss. 81 and 499.) These statements led
Gartner, who was highly sceptical on the subject, carefully to try a long
series of experiments: he selected 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. (11/129. 'Gardener's Chronicle' 1854 page 404.)
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 (11/130. Ibid 1866 page 900.) 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. Laxton 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. A space of about two inches in length towards
the extremity of the pod, and a smaller space near the stalk, were thus
coloured. On comparing 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 immediately 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 possibly have been an accidental
circumstance, for I know not how far their thickness is a variable
character in the Tall Sugar-pea.
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 stipules, 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 themselves, 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 readily, as Gartner
doubted similar statements previously made with respect to the stock by
other observers. A well-known horticulturist, Major Trevor Clarke, informs
me (11/131. See also a paper by this observer read before the International
Hort. and Bot. Congress of London 1866.) that the seeds of the large red-
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 observed 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.
Gallesio (11/132. 'Traite du Citrus' page 40.) 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 (11/133. 'Transact. Hort. Soc.'
volume 3 page 318. See also volume 5 page 65.) fertilised a green-fleshed
melon with pollen from a scarlet-fleshed kind; in two of the fruits "a
sensible change was perceptible: and four other fruits were somewhat
altered both internally 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 Cucurbitaceae are largely cultivated,
it is the popular belief (11/134. Prof. Asa Gray 'Proc. Acad. Sc.' Boston
volume 4 1860 page 21. I have received statements to the same effect from
other persons in the United States.) that the fruit is thus directly
affected hy foreign pollen; and I have received a similar statement with
respect to the cucumber in England. It is believed 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 Teinturier; 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. (11/135. For the French case see 'Journ. Hort. Soc.' volume 1 new
series 1866 page 50. For Germany see M. Jack quoted in Henfrey's 'Botanical
Gazette' volume 1 page 277. A case in England has recently been alluded to
by the Rev. J.M. Berkeley before the Hort. Soc. of London.)
As long ago as 1751 (11/136. 'Philosophical Transactions' volume 47 1751-52
page 206.) it was observed that, when differently-coloured varieties of
maize grew near each other, they mutually affected each other's seeds, and
this is now a popular belief in the United States. Dr. Savi (11/137.
Gallesio 'Teoria della Riproduzione' 1816 page 95.) tried the experiment
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, the
differently coloured seeds being arranged irregularly or in rows. Prof.
Hildebrand has repeated the experiment (11/138. 'Bot. Zeitung' May 1868
page 326.) with the precaution of ascertaining that the mother-plant was
true. A kind bearing yellow grains was fertilised with pollen of a kind
having brown grains, and two ears produced yellow grains mingled with
others of a dirty violet tint. A third ear had only yellow grains, but one
side of the spindle was tinted of a reddish-brown; so that here we have the
important fact of the influence of the foreign pollen extending to the
axis. Mr. Arnold, in Canada, varied the experiment in an interesting
manner: "a female flower was subjected first to the action of pollen from a
yellow variety, and then to that from a white variety; the result was an
ear, each grain of which was yellow below and white above." (11/139. See
Dr. J. Stockton-Hough 'American Naturalist' January 1874 page 29.) With
other plants it has occasionally been observed that the crossed offspring
showed the influence of two kinds of pollen, but in this case the two kinds
affected the mother-plant.
Mr. Sabine states (11/140. 'Transact. Hort. Soc.' volume 5 page 69.) 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. With an allied genus, a well-known
botanist, Maximowicz, has described in detail the striking results of
reciprocally fertilising Lilium bulbiferum and davuricum with each other's
pollen. Each species produced fruit not like its own, but almost identical
with that of the pollen-bearing species; but from an accident only the
fruit of the latter species was carefully examined; the seeds were
intermediate in the development of their wings. (11/141. 'Bull. de l'Acad.
Imp. de St. Petersburg' tome 17 page 275, 1872. The author gives references
to those cases in the So1anaceae of fruit affected by foreign pollen, but
as it does not appear that the mother-plant was artificially fertilised, I
have not entered into details.)
Fritz Muller fertilised Cattleya leopoldi with pollen of Epidendron
cinnabarinum; and the capsules contained very few seeds; but these
presented a most wonderful appearance, which, from the description given,
two botanists, Hildebrand and Maximowicz, attribute to the direct action of
the pollen of the Epidendron. (11/142. 'Bot. Zeitung' September 1868 page
631. For Maximowicz's judgment see the paper last referred to.)
Mr. J. Anderson Henry (11/143. 'Journal of Horticulture' January 20, 1863
page 46.) crossed Rhododendron dalhousiae 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, measured 1 2/8 inch in length and 1 1/2 in girth; whilst three of
the pods which had been fertilised by pollen of R. nuttallii measured 1 5/8
inch in length and no less than 2 inches in girth. Here the effect of the
foreign pollen was apparently confined to increasing the size of the
ovarium; but we must be cautious in assuming, as the following case shows,
that size had been transferred 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
detailed measurements and sketches, were much larger in all their
dimensions 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 increased size
of the pods in the foregoing cases may be an analogous fact. On the other
hand, M. de Saporta informs me that an isolated female plant of Pistacia
vera is very apt to be fertilised by the pollen of neighbouring plants of
P. terebinthus, and in this case the fruits are only half their proper
size, which he attributes to the influence of the pollen of P. terebinthus.
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 (11/144. See on this head the high authority of Prof.
Decaisne in a paper translated in 'Journ. Hort. Soc.' volume 1 new series
1866 page 48.) in a metamorphosed condition, so that the effect of the
foreign pollen has extended even beyond 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' (11/145. Volume 43 1744-45 page 525 volume 45 1747-48 page
602.); in one of these a Russeting apple and an adjoining kind mutually
affected each other's fruit; and in another case a smooth apple affected a
rough-coated kind. Another instance has been given (11/146. 'Transact.
Hort. Soc.' volume 5 pages 65 and 68. See also Prof. Hildebrand with a
coloured figure in 'Bot. Zeitung' May 15, 1868 page 327. Puvis also has
collected 'De La Degeneration' 1837 page 36) several other instances; but
it is not in all cases possible to distinguish between the direct action of
foreign pollen and bud-variations.) 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, the
flowers which, from the abortion of the stamens, do not produce pollen, but
are fertilised by the girls of the neighbourhood with pollen of many kinds;
and they bear fruit, "differing from one another in size, flavour, and
colour, but resembling in character the hermaphrodite kinds by which they
have been fertilised." (11/147. T. de Clermont-Tonnerre in 'Mem. de la Soc.
Linn. de Paris' tome 3 1825 page 164.)]
I have now shown, on the authority of several excellent observers, in the
case of plants belonging to widely different orders, that the pollen of one
species or variety, when applied to the female of a distinct form,
occasionally causes the coats of the seeds, the ovarium or fruit, including
even the calyx and upper part of the peduncle of the apple, and the axis of
the ear in maize, to be modified. Sometimes the whole 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, is thus affected. It must not be
supposed that any direct 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 (11/148. 'Transact. of Hort.
Soc.' volume 5 page 68.) expressly states that he has never seen the fruit
thus affected, though he 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 directly modified by foreign pollen is itself affected, so as
afterwards 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 bud-variation in the fruit of trees,
given in the early part of this chapter can be accounted for by the action
of foreign pollen; for such fruits have commonly been propagated by budding
or grafting. It is also obvious that changes of colour in flowers, which
necessarily supervene long before they are ready for fertilisation, and
changes in the shape or colour of leaves, when due to the appearance of
modified buds, can have no relation to the action of foreign pollen.
The proofs of the action of foreign pollen on the mother-plant have been
given in considerable detail, because this action, as we shall see in a
future chapter, is of the highest theoretical importance, and because it is
in itself a remarkable 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 at the same time various parts of the mother-plant, in the same manner,
as it affects the same part in the seminal offspring from the same two
parents. We thus learn that an ovule is not indispensable for the reception
of the influence of the male element. But this direct action of the male
element is not so anomalous as it at first appears, for it comes into play
in the ordinary fertilisation of many flowers. Gartner gradually increased
the number of pollen grains until he succeeded in fertilising a Malva, and
has (11/149. Beitrage zur Kenntniss der Befruchtung' 1844 s. 347-351.)
proved that many grains are first expended in the development, or, as he
expresses it, in the satiation, of the pistil and ovarium. Again, when one
plant is fertilised 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 formed without any embryo being developed
within. Prof. Hildebrand, also, has lately shown (11/150. 'Die
Fruchtbildung der Orchideen, ein Beweis fur die doppelte Wirkung des
Pollens' 'Botanische Zeitung' No. 44 et seq. October 30, 1865; and August
4, 1865 s. 249.) that, in the normal fertilisation of several Orchideae,
the action of the plant's own pollen 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 placentae and
ovules have been formed; so that with these orchids the pollen acts
directly on the ovarium. On the other hand, we must not overrate the
efficacy of pollen in the case of hybridised plants, for an embryo may be
formed and its influence excite the surrounding tissues of the mother-
plant, and then perish at a very early age and be thus overlooked. Again,
it is well known that with many plants the ovarium may be fully developed,
though pollen be wholly excluded. Lastly, Mr. Smith, the late Curator at
Kew (as I hear through Dr. Hooker), observed with an orchid, the Bonatea
speciosa, the singular fact that the development of the ovarium could be
effected by the 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 hybridised plants which produce no seeds,--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,
independently of the intervention of the fertilised germ. Therefore, in the
previously given cases we have only to believe in the further power of
pollen, when applied to a distinct species or variety, to influence the
shape, size, colour, texture, etc., of certain parts of the mother-plant.
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 pollen
should next year produce, when self-fertilised, offspring modified by the
previous male influence. Closely analogous cases have actually occurred
with animals. In the case often quoted from Lord Morton (11/151. 'Philos.
Transact.' 1821 page 20.) a nearly purely-bred Arabian chestnut 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, are extremely rare,--I speak after having
long attended to the subject,--with horses of all kinds in Europe, and are
almost unknown in the case of Arabians. But what makes the case still more
striking is that in these colts the hair of the mane resembled that of the
quagga, being short, stiff, and upright. Hence there can be no doubt that
the quagga affected the character of the offspring subsequently begot by
the black Arabian horse. Mr. Jenner Weir informs me of a strictly parallel
case: his neighbour Mr. Lethbridge, of Blackheath, has a horse, bred by
Lord Mostyn, which had previously borne a foal by a quagga. This horse is
dun with a dark stripe down the back, faint stripes on the forehead between
the eyes, plain stripes on the inner side of the fore-legs and rather more
faint ones on the hind-legs, with no shoulder-stripe. The mane grows much
lower on the forehead than in the horse, but not so low as in the quagga or
zebra. The hoofs are proportionally longer than in the horse,--so much so
that the farrier who first shod this animal, and knew nothing of its
origin, said, "Had I not seen I was shoeing a horse, I should have thought
I was shoeing a donkey."
With respect to the varieties of our domesticated animals, many similar and
well-authenticated facts have been published (11/152. Dr. Alex. Harvey on
'A remarkable Effect of Cross-breeding' 1851. On the 'Physiology of
Breeding' by Mr. Reginald Orton 1855. 'Intermarriage' by Alex. Walker 1837.
'L'Heredite Naturelle' by Dr. Prosper Lucas tome 2 page 58. Mr. W. Sedgwick
in 'British and Foreign Medico-Chirurgical Review' 1863 July page 183.
Bronn in his 'Geschichte der Natur' 1843 b. 2 s. 127 has collected several
cases with respect to mares, sows, and dogs. Mr. W.C.L. Martin ('History of
the Dog' 1845 page 104) says he can personally vouch for the influence of
the male parent on subsequent litters by other dogs. A French poet, Jacques
Savary, who wrote in 1665 on dogs, was aware of this singular fact. Dr.
Bowerbank has given us the following striking case:--A black, hairless
Barbary bitch was first accidentally impregnated by a mongrel spaniel with
long brown hair, and she produced five puppies, three of which were
hairless and two covered with SHORT brown hair. The next time she was put
to a 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." I have
given in the text one case with pigs; an equally striking one has been
recently published in Germany, 'Illust. Landwirth. Zeitung' 1868 November
17 page 143. It is worth notice that farmers in S. Brazil (as I hear from
Fritz Muller), and at the C. of Good Hope (as I have heard from two
trustworthy persons) are convinced that mares which have once borne mules,
when subsequently put to horses, are extremely liable to produce colts,
striped like a mule. Dr. Wilckens of Pogarth gives ('Jahrbuch
Landwirthschaft' 2 1869 page 325) a striking and analogous case. A merino
ram, having two small lappets or flaps of skin on the neck, was in the
winter of 1861-62 put to several Merino ewes, all of whom bore lambs with
similar flaps on their necks. The ram was killed in the spring of 1862 and
subsequently to his death the ewes were put to other Merino rams, and in
1863 to Southdown rams, none of whom ever have neck lappets: nevertheless,
even as long afterwards as 1867, several of these ewes produced lambs
bearing these appendages.), and others have been communicated 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
instance, 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 chestnut colour; and the
"pigs produced partook in appearance of both boar and sow, but in some the
chestnut 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 chestnut
colour,--yet from this union the sow produced some young pigs which were
plainly marked with the same chestnut tint as in the first litter. Similar
cases have so frequently occurred, that careful breeders avoid putting a
choice female of any animal to an inferior male, on account of the injury
to her subsequent progeny which may be expected to follow.
Some physiologists have attempted to account for these remarkable results
from a previous impregnation, by the imagination of the mother having been
strongly affected; but it will hereafter be seen that there are very slight
grounds for any such belief. Other physiologists attribute the result to
the close attachment and freely intercommunicating blood-vessels between
the modified embryo and mother. But the analogy from the action of foreign
pollen on the ovarium, seed-coats, and other parts of the mother-plant,
strongly supports the belief that with animals the male element acts
directly on the female, and not through the crossed embryo. With birds
there is no close connection between the embryo and mother; yet a careful
observer, Dr. Chapuis, states (11/153. 'Le Pigeon Voyageur Belge' 1865 page
59.) that with pigeons the influence of a first male sometimes makes itself
perceived in the succeeding broods; but this statement 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 extraordinary modes the union of
one form with another may lead to the modification of the seminal offspring
or of the buds, afterwards produced.
There is nothing surprising in the offspring of species or varieties
crossed in the ordinary manner being modified; but the case of two plants
within the same seed, which cohere and differ from each other, is curious.
When a bud is formed after the cellular tissue of two species or two
varieties have been united, and it partakes of the characters of both
parents, the case is wonderful. But I need not here repeat what has been so
lately said on this subject. We have also seen that in the case of plants
the male element may affect in a direct manner the tissues of the mother,
and with animals may lead to the modification of her future progeny. In the
vegetable kingdom the offspring from a cross between two species or
varieties, whether effected by seminal generation or by grafting, often
revert, to a greater or less degree, in the first or in a succeeding
generation, to the two parent-forms; and this reversion may affect the
whole flower, fruit, or leaf-bud, or only the half or a smaller segment of
a single organ. In some cases, however, such segregation of character
apparently depends on an incapacity for union rather than on reversion, for
the flowers or fruit which are first produced display by segments the
characters of both parents. The various facts here given ought to be well
considered by any one who wishes to embrace under a single point of view
the many modes of reproduction by gemmation, division, and sexual union,
the reparation of lost parts, variation, inheritance, reversion, and other
such phenomena. Towards the close of the second volume I shall attempt to
connect these facts together by the hypothesis of pangenesis.
In the early half of the present chapter I have given a long list of plants
in which through bud-variation, that is, independently of reproduction by
seed, the fruit has suddenly become modified in size, colour, flavour,
hairiness, shape, and time of maturity; flowers have similarly changed in
shape, colour, in being double, 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 or in weeping; leaves have changed in
becoming variegated, in shape, period of unfolding, and in their
arrangement on the axis. Buds of all kinds, whether produced on ordinary
branches or on subterranean stems, whether simple or much modified and
supplied with a stock of nutriment, as in tubers and bulbs, are all liable
to sudden variations of the same general nature.
In the list, many of the cases are certainly due to reversion to characters
not acquired from a cross, but which were formerly present and have since
been lost for a longer or shorter time;--as when a bud on a variegated
plant produces plain leaves, or when the variously-coloured flowers of the
Chrysanthemum 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 either completely or by segments to
one of the two parent-forms. (11/154. It may be worth while to call
attention to the several means by which flowers and fruit become striped or
mottled. Firstly, by the direct action of the pollen of another variety or
species, as in the cases given of oranges and maize. Secondly, in crosses
of the first generation, when the colours of the two parents do not readily
unite, as with Mirabilis and Dianthus. Thirdly, in crossed plants of a
subsequent generation by reversion, through either bud or seminal
generation. Fourthly, by reversion to a character not originally gained by
a cross, but which had long been lost, as with white-flowered varieties,
which we shall hereafter see often become striped with some other colour.
Lastly, there are cases, as when peaches are produced with a half or
quarter of the fruit like a nectarine, in which the change is apparently
due to mere variation, through either bud or seminal generation.)
We may suspect that the strong tendency in the Chrysanthemum to produce by
bud-variation differently-coloured flowers, results from the varieties
having been at some time intentionally or accidentally crossed; and this is
certainly the case with some kinds of Pelargonium. So it may be to a large
extent with the bud-varieties of the Dahlia, and with the "broken colours
"of Tulips. When, however, a plant reverts by bud-variation to its two
parent forms, or to one of them, it sometimes does not revert perfectly,
but assumes a somewhat new character,--of which fact, instances have been
given, and Carriere gives (11/155. 'Production des Varietes' page 37.)
another in the cherry.
Many cases of bud-variation, however, cannot be attributed to reversion,
but to so-called spontaneous variability, as is so common with cultivated
plants raised from seed. As a single variety of the Chrysanthemum has
produced by buds six other varieties, and as one variety of the gooseberry
has borne at the same time four distinct kinds of fruit, it is scarcely
possible to believe that all these variations are due to reversion. 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
rose which bears opposite leaves, in that of the Imatophyllum, etc., there
is no known natural species or variety from which the characters in
question could have been derived by a cross. We must attribute all such
cases to the appearance of absolutely new characters in the buds. The
varieties 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.
The plants which have varied by buds belong to so many orders that we may
infer that almost every plant would be liable to 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 list are perennials, and have been largely
propagated in many soils, under different climates, by cuttings, offsets,
bulbs, tubers, and especially by budding or grafting. The instances of
annuals varying by buds, or producing on the same plant differently
coloured flowers, are comparatively rare: Hopkirk (11/156. 'Flora Anomala'
page 164.) has seen this with Convolvulus tricolor; and it is not uncommon
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. Domingo, are eminently liable to bud-variation. I am
informed by Mr. Sedgwick that moss-roses which have often been taken to
Calcutta always there lose their mossiness; but change of climate is by no
means a necessary contingent, as we see with the gooseberry, currant, and
in many other cases. Plants living under their natural conditions are very
rarely subject to bud-variation. Variegated leaves have, however, been
observed under such circumstances; and I have given an instance of
variation by buds on an ash-tree planted in ornamental grounds, but it is
doubtful whether such a tree can be considered as living under strictly
natural conditions. Gartner has seen white and dark-red flowers produced
from the same root of the wild Achillea millefolium; and Prof. Caspary has
seen a completely wild Viola lutea bearing flowers of two different colours
and sizes. (11/157. 'Schriften der physisch-okon. Gesell. zu Konigsberg' b.
6 February 3, 1865 s. 4.)
As wild plants are so rarely liable to bud-variation, whilst highly
cultivated plants long propagated by artificial means have yielded many
varieties by this form of reproduction, 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 purple plum-tree suddenly
becoming yellow,--all the fruit on a double-flowered almond suddenly
becoming peach like,--all the buds on grafted trees being in a 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 colour, size,
and shape,--we are led by such a series to look at every case of bud-
variation as the direct result of the conditions of life to which the plant
has been exposed. On the other hand, plants of the same variety may be
cultivated in two adjoining beds, apparently under exactly the same
conditions, and those in the one bed, as Carriere insists (11/158.
'Production des Varietes' pages 58, 70.) will produce many bud-variations,
and those in the other not a single one. Again, if we look 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 millions of buds, all of which have apparently been exposed to
precisely the same conditions, yet at last suddenly produces a single bud
with its whole character greatly transformed, we are driven to the
conclusion 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 appearance that they cannot be distinguished.
Just as certain species and groups of species, when propagated by seed, are
more variable than other species or 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 Rollisson's Unique Pelargonium
four distinct varieties; moss-roses have also produced several other moss-
roses. The Rosaceae 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 same group, the peach has often varied by
buds, whilst the apple and pear, both grafted trees extensively cultivated,
have afforded, as far as I can ascertain, extremely few instances of bud-
variation.
The law of analogous variation holds good with varieties 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 commonly 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 seedling varieties,
some transmit their character much more truly by seed than others; so with
bud-varieties, some retain 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 and pelargoniums.
Notwithstanding the sudden production of bud-varieties, the characters thus
acquired are sometimes capable of transmission by seminal reproduction: Mr.
Rivers has found that moss-roses generally reproduce themselves by seed;
and the mossy character has been transferred 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. On the other hand,
seedlings from some bud-variations have proved variable to an extreme
degree. (11/159. Carriere 'Production des Varietes' page 39.) We have also
heard, on the authority of Mr. Salter, that seeds taken from a branch with
leaves variegated through bud-variation, transmit this character very
feebly; whilst many plants, which were variegated as seedlings, transmit
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 very many more cases, yet their total
number is as nothing in comparison with that of seminal varieties. With
seedlings raised from the more variable cultivated plants, the variations
are almost infinitely numerous, 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 inexplicable 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 were overlooked or not recorded from being of no value.
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 nectarines, 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 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 irregular 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 all their functions--in their power of
inheritance with occasional reversion,--and in their capacity for variation
of the same general nature, in obedience to the same laws. This
resemblance, or rather identity of character, is shown in the most striking
manner by the fact 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. We have seen that variability does not depend on sexual
generation, though much more frequently its concomitant than of bud
reproduction. We have seen that bud-variability is not solely dependent on
reversion or atavism to long-lost characters, or to those formerly acquired
from a cross, but appears often to be 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 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
combustible matter.
CHAPTER 1.XII.
INHERITANCE.
WONDERFUL NATURE OF INHERITANCE.
PEDIGREES OF OUR DOMESTICATED ANIMALS.
INHERITANCE NOT DUE TO CHANCE.
TRIFLING CHARACTERS INHERITED.
DISEASES INHERITED.
PECULIARITIES IN THE EYE INHERITED.
DISEASES IN THE HORSE.
LONGEVITY AND VIGOUR.
ASYMMETRICAL DEVIATIONS OF STRUCTURE.
POLYDACTYLISM AND REGROWTH OF SUPERNUMERARY DIGITS AFTER AMPUTATION.
CASES OF SEVERAL CHILDREN SIMILARLY AFFECTED FROM NON-AFFECTED PARENTS.
WEAK AND FLUCTUATING INHERITANCE: IN WEEPING TREES, IN DWARFNESS, COLOUR OF
FRUIT AND FLOWERS.
COLOUR OF HORSES.
NON-INHERITANCE IN CERTAIN CASES.
INHERITANCE OF STRUCTURE AND HABITS OVERBORNE BY HOSTILE CONDITIONS OF
LIFE, BY INCESSANTLY RECURRING VARIABILITY, AND BY REVERSION.
CONCLUSION.
The subject of inheritance is an immense one, and has been treated by many
authors. One work alone, 'De l'Heredite Naturelle' by Dr. Prosper Lucas,
runs to the length of 1562 pages. We must confine ourselves to certain
points which have an important bearing on the general subject of variation,
both with domestic and natural productions. It is obvious that a variation
which is not inherited throws no light on the derivation of species, nor is
of any service to man, except in the case of perennial plants, which can be
propagated by buds.
If animals and plants had never been domesticated, and wild ones alone had
been observed, we should probably never have heard the saying, that "like
begets like." The proposition would have been as self-evident as that all
the buds on the same tree are alike, though neither proposition is strictly
true. For, as has often been remarked, probably no two individuals are
identically the same. All wild animals recognise each other, which shows
that there is some difference between them; and when the eye is well
practised, the shepherd knows each sheep, and man can distinguish a fellow-
man out of millions on millions of other men. Some authors have gone so far
as to maintain that the production of slight differences is as much a
necessary function of the powers of generation, as the production of
offspring like their parents. This view, as we shall see in a future
chapter, is not theoretically probable, though practically it holds good.
The saying that "like begets like" has, in fact, arisen from the perfect
confidence felt by breeders, that a superior or inferior animal will
generally reproduce its kind; but this very superiority or inferiority
shows that the individual in question has departed slightly from its type.
The whole subject of inheritance is wonderful. When a new character arises,
whatever its nature may be, it generally tends to be inherited, at least in
a temporary and sometimes in a most persistent manner. What can be more
wonderful than that some trifling peculiarity, not primordially attached to
the species, should be transmitted through the male or female sexual cells,
which are so minute as not to be visible to the naked eye, and afterwards
through the incessant changes of a long course of development, undergone
either in the womb or in the egg, and ultimately appear in the offspring
when mature, or even when quite old, as in the case of certain diseases? Or
again, what can be more wonderful than the well-ascertained fact that the
minute ovule of a good milking cow will produce a male, from whom a cell,
in union with an ovule, will produce a female, and she, when mature, will
have large mammary glands, yielding an abundant supply of milk, and even
milk of a particular quality? Nevertheless, the real subject of surprise
is, as Sir H. Holland has well remarked (12/1. 'Medical Notes and
Reflections' 3rd edition 1855 page 267.) not that a character should be
inherited, but that any should ever fail to be inherited. In a future
chapter, devoted to an hypothesis which I have termed pangenesis, an
attempt will be made to show the means by which characters of all kinds are
transmitted from generation to generation.
Some writers (12/2. Mr. Buckle in his 'History of Civilisation' expresses
doubts on the subject, owing to the want of statistics. See also Mr. Bowen,
Professor of Moral Philosophy, in 'Proc. American Acad. of Sciences' volume
5 page 102.), who have not attended to natural history, have attempted to
show that the force of inheritance has been much exaggerated. The breeders
of animals would smile at such simplicity; and if they condescended to make
any answer, might ask what would be the chance of winning a prize if two
inferior animals were paired together? They might ask whether the half-wild
Arabs were led by theoretical notions to keep pedigrees of their horses?
Why have pedigrees been scrupulously kept and published of the Shorthorn
cattle, and more recently of the Hereford breed? Is it an illusion that
these recently improved animals safely transmit their excellent qualities
even when crossed with other breeds? have the Shorthorns, without good
reason, been purchased at immense prices and exported to almost every
quarter of the globe, a thousand guineas having been given for a bull? With
greyhounds pedigrees have likewise been kept, and the names of such dogs,
as Snowball, Major, etc., are as well known to coursers as those of Eclipse
and Herod on the turf. Even with the Gamecock, pedigrees of famous strains
were formerly kept, and extended back for a century. With pigs, the
Yorkshire and Cumberland breeders "preserve and print pedigrees;" and to
show how such highly-bred animals are valued, I may mention that Mr. Brown,
who won all the first prizes for small breeds at Birmingham in 1850, sold a
young sow and boar of his breed to Lord Ducie for 43 guineas; the sow alone
was afterwards sold to the Rev. F. Thursby for 65 guineas; who writes, "She
paid me very well, having sold her produce for 300 pounds, and having now
four breeding sows from her." (12/3. For greyhounds see Low 'Domestic
Animals of the British Islands' 1845 page 721. For game-fowls see 'The
Poultry Book' by Mr. Tegetmeier 1866 page 123. For pigs see Mr. Sidney's
edition of 'Youatt on the Pig' 1860 pages 11, 22.) Hard cash paid down,
over and over again, is an excellent test of inherited superiority. In
fact, the whole art of breeding, from which such great results have been
attained during the present century, depends on the inheritance of each
small detail of structure. But inheritance is not certain; for if it were,
the breeder's art (12/4. 'The Stud Farm' by Cecil page 39.) would be
reduced to a certainty, and there would be little scope left for that
wonderful skill and perseverance shown by the men who have left an enduring
monument of their success in the present state of our domesticated animals.
It is hardly possible, within a moderate compass, to impress on the mind of
those who have not attended to the subject, the full conviction of the
force of inheritance which is slowly acquired by rearing animals, by
studying the many treatises which have been published on the various
domestic animals, and by conversing with breeders. I will select a few
facts of the kind, which, as far as I can judge, have most influenced my
own mind. With man and the domestic animals, certain peculiarities have
appeared in an individual, at rare intervals, or only once or twice in the
history of the world, but have reappeared in several of the children and
grandchildren. Thus Lambert, "the porcupine-man," whose skin was thickly
covered with warty projections, which were periodically moulted, had all
his six children and two grandsons similarly affected. (12/5.
'Philosophical Transactions' 1755 page 23. I have seen only second-hand
accounts of the two grandsons. Mr. Sedgwick, in a paper to which I shall
hereafter often refer, states that FOUR generations were affected, and in
each the males alone.) The face and body being covered with long hair,
accompanied by deficient teeth (to which I shall hereafter refer), occurred
in three successive generations in a Siamese family; but this case is not
unique, as a woman (12/6. Barbara Van Beck, figured, as I am informed by
the Rev. W.D. Fox, in Woodburn's 'Gallery of Rare Portraits' 1816 volume 2)
with a completely hairy face who was exhibited in London in 1663, and
another instance has recently occurred. Colonel Hallam (12/7. 'Proc.
Zoolog. Soc.' 1833 page 16.) has described a race of two-legged pigs, "the
hinder extremities being entirely wanting;" and this deficiency was
transmitted through three generations. In fact, all races presenting any
remarkable peculiarity, such as solid-hoofed swine, Mauchamp sheep, niata
cattle, etc., are instances of the long-continued inheritance of rare
deviations of structure.
When we reflect that certain extraordinary peculiarities have thus appeared
in a single individual out of many millions, all exposed in the same
country to the same general conditions of life, and, again, that the same
extraordinary peculiarity has sometimes appeared in individuals living
under widely different conditions of life, we are driven to conclude that
such peculiarities are not directly due to the action of the surrounding
conditions, but to unknown laws acting on the organisation or constitution
of the individual;--that their production stands in hardly closer relation
to the conditions of life than does life itself. If this be so, and the
occurrence of the same unusual character in the child and parent cannot be
attributed to both having been exposed to the same unusual conditions, then
the following problem is worth consideration, as showing that the result
cannot be due, as some authors have supposed, to mere coincidence, but must
be consequent on the members of the same family inheriting something in
common in their constitution. Let it be assumed that, in a large
population, a particular affection occurs on an average in one out of a
million, so that the a priori chance that an individual taken at random
will be so affected is only one in a million. Let the population consist of
sixty millions, composed, we will assume, of ten million families, each
containing six members. On these data, Professor Stokes has calculated for
me that the odds will be no less than 8333 millions to 1 that in the ten
million families there will not be even a single family in which one parent
and two children will be affected by the peculiarity in question. But
numerous instances could be given, in which several children have been
affected by the same rare peculiarity with one of their parents; and in
this case, more especially if the grandchildren be included in the
calculation, the odds against mere coincidence become something prodigious,
almost beyond enumeration.
In some respects the evidence of inheritance is more striking when we
consider the reappearance of trifling peculiarities. Dr. Hodgkin formerly
told me of an English family in which, for many generations, some members
had a single lock differently coloured from the rest of the hair. I knew an
Irish gentleman, who, on the right side of his head, had a small white lock
in the midst of his dark hair: he assured me that his grandmother had a
similar lock on the same side, and his mother on the opposite side. But it
is superfluous to give instances; every shade of expression, which may
often be seen alike in parents and children, tells the same story. On what
a curious combination of corporeal structure, mental character, and
training, handwriting depends! yet every one must have noted the occasional
close similarity of the handwriting in father and son, although the father
had not taught his son. A great collector of autographs assured me that in
his collection there were several signatures of father and son hardly
distinguishable except by their dates. Hofacker, in Germany, remarks on the
inheritance of handwriting; and it has even been asserted that English boys
when taught to write in France naturally cling to their English manner of
writing; but for so extraordinary a statement more evidence is requisite.
(12/8. Hofacker 'Ueber die Eigenschaften' etc. 1828 s. 34. With respect to
France, Report by Pariset in 'Comptes Rendus' 1847 page 592.) Gait,
gestures, voice, and general bearing are all inherited, as the illustrious
Hunter and Sir A. Carlisle have insisted. (12/9. Hunter as quoted in
Harlan's 'Med. Researches' page 530. Sir A. Carlisle 'Phil. Transact.' 1814
page 94.) My father communicated to me some striking instances, in one of
which a man died during the early infancy of his son, and my father, who
did not see this son until grown up and out of health, declared that it
seemed to him as if his old friend had risen from the grave, with all his
highly peculiar habits and manners. Peculiar manners pass into tricks, and
several instances could be given of their inheritance; as in the case,
often quoted, of the father who generally slept on his back, with his right
leg crossed over the left, and whose daughter, whilst an infant in the
cradle, followed exactly the same habit, though an attempt was made to cure
her. (12/10. Girou de Buzareingues 'De la Generation' page 282. I have
given an analogous case in my book on 'The Expression of the Emotions.') I
will give one instance which has fallen under my own observation, and which
is curious from being a trick associated with a peculiar state of mind,
namely, pleasureable emotion. A boy had the singular habit, when pleased,
of rapidly moving his fingers parallel to each other, and, when much
excited, of raising both hands, with the fingers still moving, to the sides
of his face on a level with the eyes; when this boy was almost an old man,
he could still hardly resist this trick when much pleased, but from its
absurdity concealed it. He had eight children. Of these, a girl, when
pleased, at the age of four and a half years, moved her fingers in exactly
the same way, and what is still odder, when much excited, she raised both
her hands, with her fingers still moving, to the sides of her face, in
exactly the same manner as her father had done, and sometimes even still
continued to do so when alone. I never heard of any one, excepting this one
man and his little daughter, who had this strange habit; and certainly
imitation was in this instance out of the question.
Some writers have doubted whether those complex mental attributes, on which
genius and talent depend, are inherited, even when both parents are thus
endowed. But he who will study Mr. Galton's able work on 'Hereditary
Genius' will have its doubts allayed.
Unfortunately it matters not, as far as inheritance is concerned, how
injurious a quality or structure may be if compatible with life. No one can
read the many treatises (12/11. The works which I have read and found most
useful are Dr. Prosper Lucas's great work 'Traite de l'Heredite Naturelle'
1847; Mr. W. Sedgwick in 'British and Foreign Medico-Chirurg. Review' April
and July 1861 and April and July 1863: Dr. Garrod on Gout is quoted in
these articles. Sir Henry Holland 'Medical Notes and Reflections' 3rd
edition 1855. Piorry 'De l'Heredite dans les Maladies' 1840. Adams 'A
Philosophical Treatise on Hereditary Peculiarities' 2nd edition 1815. Essay
on 'Hereditary Diseases' by Dr. J. Steinan 1843. See Paget in 'Medical
Times' 1857 page 192 on the Inheritance of Cancer; Dr. Gould in 'Proc. of
American Acad. of Sciences' November 8, 1853 gives a curious case of
hereditary bleeding in four generations. Harlan 'Medical Researches' page
593.) on hereditary disease and doubt this. The ancients were strongly of
this opinion, or, as Ranchin expresses it, Omnes Graeci, Arabes, et Latini
in eo consentiunt. A long catalogue could be given of all sorts of
inherited malformations and of predisposition to various diseases. With
gout, fifty per cent of the cases observed in hospital practice are,
according to Dr. Garrod, inherited, and a greater percentage in private
practice. Every one knows how often insanity runs in families, and some of
the cases given by Mr. Sedgwick are awful,--as of a surgeon, whose brother,
father, and four paternal uncles were all insane, the latter dying by
suicide; of a Jew, whose father, mother, and six brothers and sisters were
all mad; and in some other cases several members of the same family, during
three or four successive generations, have committed suicide. Striking
instances have been recorded of epilepsy, consumption, asthma, stone in the
bladder, cancer, profuse bleeding from the slightest injuries, of the
mother not giving milk, and of bad parturition being inherited. In this
latter respect I may mention an odd case given by a good observer (12/12.
Marshall quoted by Youatt in his work on 'Cattle' page 284.), in which the
fault lay in the offspring, and not in the mother: in a part of Yorkshire
the farmers continued to select cattle with large hind-quarters, until they
made a strain called "Dutch-buttocked," and "the monstrous size of the
buttocks of the calf was frequently fatal to the cow, and numbers of cows
were annually lost in calving."
[Instead of giving numerous details on various inherited malformations and
diseases, I will confine myself to one organ, that which is the most
complex, delicate, and probably best-known in the human frame, namely, the
eye, with its accessory parts. (12/13. Almost any other organ might have
been selected. For instance Mr. J. Tomes 'System of Dental Surgery' 2nd
edition 1873 page 114 gives many instances with teeth, and others have been
communicated to me.) To begin with the latter: I have received an account
of a family in which one parent and the children are affected by drooping
eyelids, in so peculiar a manner, that they cannot see without throwing
their heads backwards. Mr. Wade, of Wakefield, has given me an analogous
case of a man who had not his eyelids thus affected at birth, nor owed
their state, as far as was known, to inheritance, but they began to droop
whilst he was an infant after suffering from fits, and he has transmitted
the affection to two out of his three children, as was evident in the
photographs of the whole family sent to me together with this account. Sir
A. Carlisle (12/14. 'Philosoph. Transact.' 1814 page 94.) specifies a
pendulous fold to the eyelids, as inherited. "In a family," says Sir H.
Holland (12/15. 'Medical Notes and Reflections' 3rd edition page 33.)
"where the father had a singular elongation of the upper eyelid, seven or
eight children were born with the same deformity; two or three other
children having it not." Many persons, as I hear from Sir J. Paget, have
two or three hairs in their eyebrows much longer than the others; and even
so trifling a peculiarity as this certainly runs in families.
With respect to the eye itself, the highest authority in England, Mr.
Bowman, has been so kind as to give me the following remarks on certain
inherited imperfections. First, hypermetropia, or morbidly long sight: in
this affection, the organ, instead of being spherical, is too flat from
front to back, and is often altogether too small, so that the retina is
brought too forward for the focus of the humours; consequently a convex
glass is required for clear vision of near objects, and frequently even of
distant ones. This state occurs congenitally, or at a very early age, often
in several children of the same family, where one of the parents has
presented it. (12/16. This affection, as I hear from Mr. Bowman, has been
ably described and spoken of as hereditary by Dr. Donders of Utrecht, whose
work was published in English by the Sydenham Society in 1864.) Secondly,
myopia, or short-sight, in which the eye is egg-shaped and too long from
front to back; the retina in this case lies behind the focus, and is
therefore fitted to see distinctly only very near objects. This condition
is not commonly congenital, but comes on in youth, the liability to it
being well known to be transmissible from parent to child. The change from
the spherical to the ovoidal shape seems the immediate consequence of
something like inflammation of the coats, under which they yield, and there
is ground for believing that it may often originate in causes acting on the
individual affected (12/17. M. Giraud-Teulon has recently collected
abundant statistical evidence, 'Revue des Cours Scientifiques' September
1870 page 625, showing that short sight is due to the habit of viewing
objects from a short distance, c'est le travail assidu, de pres.) and may
thenceforward become transmissible. When both parents are myopic Mr. Bowman
has observed the hereditary tendency in this direction to be heightened,
and some of the children to be myopic at an earlier age or in a higher
degree than their parents. Thirdly, squinting is a familiar example of
hereditary transmission: it is frequently a result of such optical defects
as have been above mentioned; but the more primary and uncomplicated forms
of it are also sometimes in a marked degree transmitted in a family.
Fourthly, CATARACT, or opacity of the crystalline lens, is commonly
observed in persons whose parents have been similarly affected, and often
at an earlier age in the children than in the parents. Occasionally more
than one child in a family is thus afflicted, one of whose parents or other
relations, presents the senile form of the complaint. When cataract affects
several members of a family in the same generation, it is often seen to
commence at about the same age in each: e.g., in one family several infants
or young persons may suffer from it; in another, several persons of middle
age. Mr. Bowman also informs me that he has occasionally seen, in several
members of the same family, various defects in either the right or left
eye; and Mr. White Cooper has often seen peculiarities of vision confined
to one eye reappearing in the same eye in the offspring. (12/18. Quoted by
Mr. Herbert Spencer 'Principles of Biology' volume 1 page 244.)
The following cases are taken from an able paper by Mr. W. Sedgwick, and
from Dr. Prosper Lucas. (12/19. 'British and Foreign Medico-Chirurg.
Review' April 1861 pages 482-6; 'L'Hered. Nat.' tome 1 pages 391-408.)
Amaurosis, either congenital or coming on late in life, and causing total
blindness, is often inherited; it has been observed in three successive
generations. Congenital absence of the iris has likewise been transmitted
for three generations, a cleft-iris for four generations, being limited in
this latter case to the males of the family. Opacity of the cornea and
congenital smallness of the eyes have been inherited. Portal records a
curious case, in which a father and two sons were rendered blind, whenever
the head was bent downwards, apparently owing to the crystalline lens, with
its capsule, slipping through an unusually large pupil into the anterior
chamber of the eye. Day-blindness, or imperfect vision under a bright
light, is inherited, as is night-blindness, or an incapacity to see except
under a strong light: a case has been recorded, by M. Cunier, of this
latter defect having affected eighty-five members of the same family during
six generations. The singular incapacity of distinguishing colours, which
has been called DALTONISM, is notoriously hereditary, and has been traced
through five generations, in which it was confined to the female sex.
With respect to the colour of the iris: deficiency of colouring matter is
well known to be hereditary in albinoes. The iris of one eye being of
different colour from that of the other, and the iris being spotted, are
cases which have been inherited. Mr. Sedgwick gives, in addition, on the
authority of Dr. Osborne (12/20. Dr. Osborne, Pres. of Royal College of
Phys. in Ireland, published this case in the 'Dublin Medical Journal' for
1835.), the following curious instance of strong inheritance: a family of
sixteen sons and five daughters all had eyes "resembling in miniature the
markings on the back of a tortoiseshell cat." The mother of this large
family had three sisters and a brother all similarly marked, and they
derived this peculiarity from their mother, who belonged to a family
notorious for transmitting it to their posterity.
Finally, Dr. Lucas emphatically remarks that there is not one single
faculty of the eye which is not subject to anomalies; and not one which is
not subjected to the principle of inheritance. Mr. Bowman agrees with the
general truth of this proposition; which of course does not imply that all
malformations are necessarily inherited; this would not even follow if both
parents were affected by an anomaly which in most cases was transmissible.]
Even if no single fact had been known with respect to the inheritance of
disease and malformations by man, the evidence would have been ample in the
case of the horse. And this might have been expected, as horses breed much
quicker than man, are matched with care, and are highly valued. I have
consulted many works, and the unanimity of the belief by veterinaries of
all nations in the transmission of various morbid tendencies is surprising.
Authors who have had wide experience give in detail many singular cases,
and assert that contracted feet, with the numerous contingent evils, of
ring-bones, curbs, splints, spavin, founder and weakness of the front legs,
roaring or broken and thick wind, melanosis, specific ophthalmia, and
blindness (the great French veterinary Huzard going so far as to say that a
blind race could soon be formed), crib-biting, jibbing and ill-temper, are
all plainly hereditary. Youatt sums up by saying "there is scarcely a
malady to which the horse is subject which is not hereditary;" and M.
Bernard adds that the doctrine "that there is scarcely a disease which does
not run in the stock, is gaining new advocates every day." (12/21. These
various statements are taken from the following works and papers:--Youatt
on 'The Horse' pages 35, 220. Lawrence 'The Horse' page 30. Karkeek in an
excellent paper in 'Gard. Chronicle' 1853 page 92. Mr. Burke in 'Journal of
R. Agricul. Soc. of England' volume 5 page 511. 'Encyclop. of Rural Sports'
page 279. Girou de Buzareingues 'Philosoph. Phys.' page 215. See following
papers in 'The Veterinary;' Roberts in volume 2 page 144; M. Marrimpoey
volume 2 page 387; Mr. Karkeek volume 4 page 5; Youatt on Goitre in 'Dogs'
volume 5 page 483: Youatt in volume 6 pages 66, 348, 412; M. Bernard volume
11 page 539; Dr. Samesreuther on 'Cattle' in volume 12 page 181; Percivall
in volume 13 page 47. With respect to blindness in horses see also a whole
row of authorities in Dr. P. Lucas's great work, tome 1 page 399. Mr. Baker
in 'The Veterinary' volume 13 page 721, gives a strong case of hereditary
imperfect vision and of jibbing.) So it is in regard to cattle, with
consumption, good and bad teeth, fine skin, etc. etc. But enough, and more
than enough, has been said on disease. Andrew Knight, from his own
experience, asserts that disease is hereditary with plants; and this
assertion is endorsed by Lindley. (12/22. Knight on 'The Culture of the
Apple and Pear' page 34. Lindley's 'Horticulture' page 180.)
Seeing how hereditary evil qualities are, it is fortunate that good health,
vigour, and longevity are equally inherited. It was formerly a well-known
practice, when annuities were purchased to be received during the life-time
of a nominee, to search out a person belonging to a family of which many
members had lived to extreme old age. As to the inheritance of vigour and
endurance, the English race-horse offers an excellent instance. Eclipse
begot 334, and King Herod 497 winners. A "cock-tail" is a horse not purely
bred, but with only one-eighth, or one-sixteenth impure blood in his veins,
yet very few instances have ever occurred of such horses having won a great
race. They are sometimes as fleet for short distances as thoroughbreds, but
as Mr. Robson, the great trainer, asserts, they are deficient in wind, and
cannot keep up the pace. Mr. Lawrence also remarks, "perhaps no instance
has ever occurred of a three-part-bred horse saving his 'DISTANCE' in
running two miles with thoroughbred racers." It has been stated by Cecil,
that when unknown horses, whose parents were not celebrated, have
unexpectedly won great races, as in the case of Priam, they can always be
proved to be descended, on both sides, through many generations, from
first-rate ancestors. On the Continent, Baron Cameronn challenges, in a
German veterinary periodical, the opponents of the English race-horse to
name one good horse on the Continent, which has not some English race-blood
in his veins. (12/23. These statements are taken from the following works
in order:--Youatt on 'The Horse' page 48; Mr. Darvill in 'The Veterinary'
volume 8 page 50. With respect to Robson see 'The Veterinary' volume 3 page
580; Mr. Lawrence on 'The Horse' 1829 page 9; 'The Stud Farm' by Cecil
1851; Baron Cameronn quoted in 'The Veterinary' volume 10 page 500.)
With respect to the transmission of the many slight, but infinitely
diversified characters, by which the domestic races of animals and plants
are distinguished, nothing need be said; for the very existence of
persistent races proclaims the power of inheritance.
A few special cases, however, deserve some consideration. It might have
been anticipated, that deviations from the law of symmetry would not have
been inherited. But Anderson (12/24. 'Recreations in Agriculture and Nat.
Hist.' volume 1 page 68.) states that a rabbit produced in a litter a young
animal having only one ear; and from this animal a breed was formed which
steadily produced one-eared rabbits. He also mentions a bitch with a single
leg deficient, and she produced several puppies with the same deficiency.
From Hofacker's account (12/25. 'Ueber die Eigenschaften' etc. 1828 s.
107.) it appears that a one-horned stag was seen in 1781 in a forest in
Germany, in 1788 two, and afterwards, from year to year, many were observed
with only one horn on the right side of the head. A cow lost a horn by
suppuration (12/26. Bronn 'Geschichte der Natur' b. 2 s. 132.), and she
produced three calves which had on the same side of the head, instead of a
horn, a small bony lump attached merely to the skin; but we here encroach
on the subject of inherited mutilations. A man who is left-handed, and a
shell in which the spire turns in the wrong directions, are departures from
the normal asymmetrical condition, and they are well-known to be inherited.
[POLYDACTYLISM.
Supernumerary fingers and toes are eminently liable, as various authors
have insisted, to be inherited. Polydactylism graduates (12/27. Vrolik has
discussed this point at full length in a work published in Dutch, from
which Sir J. Paget has kindly translated for me passages. See, also,
Isidore Geoffroy St. Hilaire 'Hist. des Anomalies' 1832 tome 1 page 684.)
by multifarious steps from a mere cutaneous appendage, not including any
bone, to a double hand. But an additional digit, supported on a metacarpal
bone, and furnished with all the proper muscles, nerves, and vessels, is
sometimes so perfect, that it escapes detection, unless the fingers are
actually counted. Occasionally there are several supernumerary digits; but
usually only one, making the total number six. This one may be attached to
the inner or outer margin of the hand, representing either a thumb or
little finger, the latter being the more frequent. Generally, through the
law of correlation, both hands and both feet are similarly affected. Dr.
Burt Wilder has tabulated (12/28. 'Massachusetts Medical Society' volume 2
No. 3; and 'Proc. Boston Soc. of Nat. Hist.' volume 14 1871 page 154.) a
large number of cases, and finds that supernumerary digits are more common
on the hands than on the feet, and that men are affected oftener than
women. Both these facts can be explained on two principles which seem
generally to hold good; firstly, that of two parts, the more specialised
one is the more variable, and the arm is more highly specialised than the
leg; and secondly that male animals are more variable than females.
The presence of a greater number of digits than five is a great anomaly,
for this number is not normally exceeded by any existing mammal, bird, or
reptile. Nevertheless, supernumerary digits are strongly inherited; they
have been transmitted through five generations; and in some cases, after
disappearing for one, two, or even three generations, have reappeared
through reversion. These facts are rendered, as Professor Huxley has
observed, more remarkable from its being known in most cases that the
affected person has not married one similarly affected. In such cases the
child of the fifth generation would have only 1-32nd part of the blood of
his first sedigitated ancestor. Other cases are rendered remarkable by the
affection gathering force, as Dr. Struthers has shown, in each generation,
though in each the affected person married one not affected; moreover, such
additional digits are often amputated soon after birth, and can seldom have
been strengthened by use. Dr. Struthers gives the following instance: in
the first generation an additional digit appeared on one hand; in the
second, on both hands; in the third, three brothers had both hands, and one
of the brothers a foot affected; and in the fourth generation all four
limbs were affected. Yet we must not over-estimate the force of
inheritance. Dr. Struthers asserts that cases of non-inheritance and of the
first appearance of additional digits in unaffected families are much more
frequent than cases of inheritance. Many other deviations of structure, of
a nature almost as anomalous as supernumerary digits, such as deficient
phalanges (12/29. Dr. J.W. Ogle gives a case of the inheritance of
deficient phalanges during four generations. He adds references to various
recent papers on inheritance 'Brit. and For. Med.-Chirurg. Review' April
1872.), thickened joints, crooked fingers, etc., are, in like manner,
strongly inherited, and are equally subject to intermission, together with
reversion, though in such cases there is no reason to suppose that both
parents had been similarly affected. (12/30. For these several statements
see Dr. Struthers 'Edinburgh New Phil. Journal' July 1863 especially on
intermissions in the line of descent. Prof. Huxley 'Lectures on our
Knowledge of Organic Nature' 1863 page 97. With respect to inheritance, see
Dr. Prosper Lucas 'L'Heredite Nat.' tome 1 page 325. Isid. Geoffroy 'Anom.'
tome 1 page 701. Sir A. Carlisle in 'Phil. Transact.' 1814 page 94. A.
Walker on 'Intermarriage' 1838 page 140 gives a case of five generations;
as does Mr. Sedgwick in 'Brit. and Foreign Medico-Chirurg. Review' April
1863 page 462. On the inheritance of other anomalies in the extremities see
Dr. H. Dobell in volume 46 of Medico-Chirurg. Transactions 1863; also Mr.
Sedgwick in op. cit. April 1863 page 460. With respect to additional digits
in the negro see Prichard 'Physical History of Mankind.' Dr. Dieffenbach
'Jour. Royal Geograph. Soc.' 1841 page 208 says this anomaly is not
uncommon with the Polynesians of the Chatham Islands; and I have heard of
several cases with Hindus and Arabs.)
Additional digits have been observed in negroes as well as in other races
of man, and in several of the lower animals, and have been inherited. Six
toes have been described on the hind feet of the newt (Salamandra
cristata), and are said to have occurred with the frog. It deserves notice,
that the six-toed newt, though adult, preserved some of its larval
characters; for part of the hyoidal apparatus, which is properly absorbed
during the act of metamorphosis, was retained. It is also remarkable that
in the case of man various structures in an embryonic or arrested state of
development, such as a cleft-palate, bifid uterus, etc., are often
accompanied by polydactylism. (12/31. Meckel and Isid G. St. Hilaire insist
on this fact. See also M. A. Roujou 'Sur quelques Analogies du Type Humain'
page 61 published, I believe, in the 'Journal of the Anthropolog. Soc. of
Paris' January 1872.) Six toes on the hinder feet are known to have been
inherited for three generations of cats. In several breeds of the fowl the
hinder toe is double, and is generally transmitted truly, as is well shown
when Dorkings are crossed with common four-toed breeds (12/32. 'The Poultry
Chronicle' 1854 page 559.) With animals which have properly less than five
digits, the number is sometimes increased to five, especially on the front
legs, though rarely carried beyond that number; but this is due to the
development of a digit already existing in a more or less rudimentary
state. Thus, the dog has properly four toes behind, but in the larger
breeds a fifth toe is commonly, though not perfectly, developed. Horses,
which properly have one toe alone fully developed with rudiments of the
others, have been described with each foot bearing two or three small
separate hoofs: analogous facts have been noticed with cows, sheep, goats,
and pigs. (12/33. The statements in this paragraph are taken from Isidore
Geoffroy St. Hilaire 'Hist. des Anomalies' tome 1 pages 688-693. Mr.
Goodman gives, 'Phil. Soc. of Cambridge' November 25, 1872 the case of a
cow with three well developed toes on each hind limb, besides the ordinary
rudiments; and her calf by an ordinary bull had extra digits. This calf
also bore two calves having extra digits.)
There is a famous case described by Mr. White of a child, three years old,
with a thumb double from the first joint. He removed the lesser thumb,
which was furnished with a nail; but to his astonishment it grew again and
reproduced a nail. The child was then taken to an eminent London surgeon,
and the newly-grown thumb was removed by its socket-joint, but again it
grew and reproduced a nail. Dr. Struthers mentions a case of the partial
regrowth of an additional thumb, amputated when a child was three months
old; and the late Dr. Falconer communicated to me an analogous instance. In
the last edition of this work I also gave a case of the regrowth of a
supernumerary little-finger after amputation; but having been informed by
Dr. Bachmaier that several eminent surgeons expressed, at a meeting of the
Anthropological Society of Munich, great doubt about my statements, I have
made more particular inquiries. The full information thus gained, together
with a tracing of the hand in its present state, has been laid before Sir
J. Paget, and he has come to the conclusion that the degree of regrowth in
this case is not greater than sometimes occurs with normal bones,
especially with the humerus, when amputated at an early age. He further
does not feel fully satisfied about the facts recorded by Mr. White. This
being so, it is necessary for me to withdraw the view which I formerly
advanced, with much hesitation, chiefly on the ground of the supposed
regrowth of additional digits, namely, that their occasional development in
man is a case of reversion to a lowly, organised progenitor provided with
more than five digits.]
I may here allude to a class of facts closely allied to, but somewhat
different from, ordinary cases of inheritance. Sir H. Holland (12/34.
'Medical Notes and Reflections' 1839 pages 24, 34. See also Dr. P. Lucas
'L'Hered. Nat.' tome 2 page 33.) states that brothers and sisters of the
same family are frequently affected, often at about the same age, by the
same peculiar disease, not known to have previously occurred in the family.
He specifies the occurrence of diabetes in three brothers under ten years
old; he also remarks that children of the same family often exhibit in
common infantile diseases, the same peculiar symptoms. My father mentioned
to me the case of four brothers who died between the ages of sixty and
seventy, in the same highly peculiar comatose state. An instance has
already been given of supernumerary digits appearing in four children out
of six in a previously unaffected family. Dr. Devay states (12/35. 'Du
Danger des Mariages Consanguins' 2nd edition 1862 page 103.) that two
brothers married two sisters, their first-cousins, none of the four nor any
relation being an albino; but the seven children produced from this double
marriage were all perfect albinoes. Some of these cases, as Mr. Sedgwick
(12/36. 'British and Foreign Medico-Chirurg. Review' July 1863 pages 183,
189.) has shown, are probably the result of reversion to a remote ancestor,
of whom no record had been preserved; and all these cases are so far
directly connected with inheritance that no doubt the children inherited a
similar constitution from their parents, and, from being exposed to nearly
similar conditions of life, it is not surprising that they should be
affected in the same manner and at the same period of life.
Most of the facts hitherto given have served to illustrate the force of
inheritance, but we must now consider cases grouped as well as the subject
allows into classes, showing how feeble, capricious, or deficient the power
of inheritance sometimes is. When a new peculiarity first appears, we can
never predict whether it will be inherited. If both parents from their
birth present the same peculiarity, the probability is strong that it will
be transmitted to at least some of their offspring. We have seen that
variegation is transmitted much more feebly by seed, taken from a branch
which had become variegated through bud-variation, than from plants which
were variegated as seedlings. With most plants the power of transmission
notoriously depends on some innate capacity in the individual: thus
Vilmorin (12/37. Verlot 'La Product. des Varietes' 1865 page 32.) raised
from a peculiarly coloured balsam some seedlings, which all resembled their
parent; but of these seedlings some failed to transmit the new character,
whilst others transmitted it to all their descendants during several
successive generations. So again with a variety of the rose, two plants
alone out of six were found by Vilmorin to be capable of transmitting the
desired character; numerous analogous cases could be given.
[The weeping or pendulous growth of trees is strongly inherited in some
cases, and, without any assignable reason, feebly in other cases. I have
selected this character as an instance of capricious inheritance, because
it is certainly not proper to the parent-species, and because, both sexes
being borne on the same tree, both tend to transmit the same character.
Even supposing that there may have been in some instances crossing with
adjoining trees of the same species, it is not probable that all the
seedlings would have been thus affected. At Moccas Court there is a famous
weeping oak; many of its branches "are 30 feet long, and no thicker in any
part of this length than a common rope:" this tree transmits its weeping
character, in a greater or less degree, to all its seedlings; some of the
young oaks being so flexible that they have to be supported by props;
others not showing the weeping tendency till about twenty years old.
(12/38. Loudon's 'Gardener's Mag.' volume 12 1836 page 368.) Mr. Rivers
fertilised, as he informs me, the flowers of a new Belgian weeping thorn
(Crataegus oxyacantha) with pollen from a crimson not-weeping variety, and
three young trees, "now six or seven years old, show a decided tendency to
be pendulous, but as yet are not so much so as the mother-plant." According
to Mr. MacNab (12/39. Verlot 'La Product. des Varietes' 1865 page 94.),
seedlings from a magnificent weeping birch (Betula alba), in the Botanic
Garden at Edinburgh, grew for the first ten or fifteen years upright, but
then all became weepers like their parent. A peach with pendulous branches,
like those of the weeping willow, has been found capable of propagation by
seed. (12/40. Bronn 'Geschichte der Natur' b. 2 s. 121. Mr. Meehan makes a
similar statement in 'Proc. Nat. of Philadelphia' 1872 page 235.) Lastly, a
weeping or rather a prostrate yew (Taxus baccata) was found in a hedge in
Shropshire; it was a male, but one branch bore female flowers, and produced
berries; these, being sown, produced seventeen trees all of which had
exactly the same peculiar habit with the parent-tree. (12/41. Rev. W.A.
Leighton 'Flora of Shropshire' page 497; and Charlesworth 'Mag. of Nat.
Hist.' volume 1 1837 page 30. I possess prostrate trees produced from these
seeds.)
These facts, it might have been thought, would have been sufficient to
render it probable that a pendulous habit would in all cases be strictly
inherited. But let us look to the other side. Mr. MacNab (12/42. Verlot op.
cit. page 93.) sowed seeds of the weeping beech (Fagus sylvatica), but
succeeded in raising only common beeches. Mr. Rivers, at my request, raised
a number of seedlings from three distinct varieties of weeping elm; and at
least one of the parent-trees was so situated that it could not have been
crossed by any other elm; but none of the young trees, now about a foot or
two in height, show the least signs of weeping. Mr. Rivers formerly sowed
above twenty thousand seeds of the weeping ash (Fraxinus excelsior), and
not a single seedling was in the least degree pendulous: in Germany, M.
Borchmeyer raised a thousand seedlings, with the same result. Nevertheless,
Mr. Anderson, of the Chelsea Botanic Garden, by sowing seed from a weeping
ash, which was found before the year 1780, in Cambridgeshire, raised
several pendulous trees. (12/43. For these several statements see Loudon's
'Gard. Magazine' volume 10 1834 pages 408, 180; and volume 9 1833 page
597.) Professor Henslow also informs me that some seedlings from a female
weeping ash in the Botanic Garden at Cambridge were at first a little
pendulous, but afterwards became quite upright: it is probable that this
latter tree, which transmits to a certain extent its pendulous habit, was
derived by a bud from the same original Cambridgeshire stock; whilst other
weeping ashes may have had a distinct origin. But the crowning case,
communicated to me by Mr. Rivers, which shows how capricious is the
inheritance of a pendulous habit, is that a variety of another species of
ash (F. lentiscifolia), now about twenty years old, which was formerly
pendulous, "has long lost this habit, every shoot being remarkably erect;
but seedlings formerly raised from it were perfectly prostrate, the stems
not rising more than two inches above the ground." Thus the weeping variety
of the common ash, which has been extensively propagated by buds during a
long period, did not with Mr. Rivers, transmit its character to one
seedling out of above twenty thousand; whereas the weeping variety of a
second species of ash, which could not, whilst grown in the same garden,
retain its own weeping character, transmitted to its character the
pendulous habit in excess!
Many analogous facts could be given, showing how apparently capricious is
the principle of inheritance. All the seedlings from a variety of the
Barberry (B. vulgaris) with red leaves inherited the same character; only
about one-third of the seedlings of the copper Beech (Fagus sylvatica) had
purple leaves. Not one out of a hundred seedlings of a variety of the
Cerasus padus, with yellow fruit, bore yellow fruit: one-twelfth of the
seedlings of the variety of Cornus mascula, with yellow fruit, came true
(12/44. These statements are taken from Alph. De Candolle 'Bot. Geograph.'
page 1083.): and lastly, all the trees raised by my father from a yellow-
berried holly (Ilex aquifolium), found wild, produced yellow berries.
Vilmorin (12/45. Verlot op. cit. page 38.) observed in a bed of Saponaria
calabrica an extremely dwarf variety, and raised from it a large number of
seedlings; some of these partially resembled their parent, and he selected
their seed; but the grandchildren were not in the least dwarfed: on the
other hand, he observed a stunted and bushy variety of Tagetes signata
growing in the midst of the common varieties by which it was probably
crossed; for most of the seedlings raised from this plant were intermediate
in character, only two perfectly resembling their parent; but seed saved
from these two plants reproduced the new variety so truly, that hardly any
selection has since been necessary.
Flowers transmit their colour truly, or most capriciously. Many annuals
come true: thus I purchased German seeds of thirty-four named sub-varieties
of one RACE of ten-week stocks (Matthiola annua), and raised a hundred and
forty plants, all of which, with the exception of a single plant, came
true. In saying this, however, it must be understood that I could
distinguish only twenty kinds out of the thirty-four named sub-varieties;
nor did the colour of the flower always correspond with the name affixed to
the packet; but I say that they came true, because in each of the thirty-
six short rows every plant was absolutely alike, with the one single
exception. Again, I procured packets of German seed of twenty-five named
varieties of common and quilled asters, and raised a hundred and twenty-
four plants; of these, all except ten were true in the above limited sense;
and I considered even a wrong shade of colour as false.
It is a singular circumstance that white varieties generally transmit their
colour much more truly than any other variety. This fact probably stands in
close relation with one observed by Verlot (12/46. Op. cit. page 59.),
namely, that flowers which are normally white rarely vary into any other
colour. I have found that the white varieties of Delphinium consolida and
of the Stock are the truest. It is, indeed, sufficient to look through a
nurseryman's seed-list, to see the large number of white varieties which
can be propagated by seed. The several coloured varieties of the sweet-pea
(Lathyrus odoratus) are very true; but I hear from Mr. Masters, of
Canterbury, who has particularly attended to this plant, that the white
variety is the truest. The hyacinth, when propagated by seed, is extremely
inconstant in colour, but "white hyacinths almost always give by seed
white-flowered plants" (12/47. Alph. De Candolle 'Geograph. Bot.' page
1082.); and Mr. Masters informs me that the yellow varieties also reproduce
their colour, but of different shades. On the other hand, pink and blue
varieties, the latter being the natural colour, are not nearly so true:
hence, as Mr. Masters has remarked to me, "we see that a garden variety may
acquire a more permanent habit than a natural species;" but it should have
been added, that this occurs under cultivation, and therefore under changed
conditions.
With many flowers, especially perennials, nothing can be more fluctuating
than the colour of the seedlings, as is notoriously the case with verbenas,
carnations, dahlias, cinerarias, and others. (12/48. See 'Cottage Gardener'
April 10, 1860 page 18 and September 10, 1861 page 456; 'Gardener's
Chronicle' 1845 page 102.) I sowed seed of twelve named varieties of
Snapdragon (Antirrhinum majus), and utter confusion was the result. In most
cases the extremely fluctuating colour of seedling plants is probably in
chief part due to crosses between differently-coloured varieties during
previous generations. It is almost certain that this is the case with the
polyanthus and coloured primrose (Primula veris and vulgaris), from their
reciprocally dimorphic structure (12/49. Darwin in 'Journal of Proc. Linn.
Soc. Bot.' 1862 page 94.); and these are plants which florists speak of as
never coming true by seed: but if care be taken to prevent crossing,
neither species is by any means very inconstant, in colour; thus I raised
twenty-three plants from a purple primrose, fertilised by Mr. J. Scott with
its pollen, and eighteen came up purple of different shades, and only five
reverted to the ordinary yellow colour: again, I raised twenty plants from
a bright-red cowslip, similarly treated by Mr. Scott, and every one
perfectly resembled its parent in colour, as likewise did, with the
exception of a single plant, 72 grandchildren. Even with the most variable
flowers, it is probable that each delicate shade of colour might be
permanently fixed so as to be transmitted by seed, by cultivation in the
same soil, by long-continued selection, and especially by the prevention of
crosses. I infer this from certain annual larkspurs (Delphinium consolida
and ajacis), of which common seedlings present a greater diversity of
colour than any other plant known to me; yet on procuring seed of five
named German varieties of D. consolida, only nine plants out of ninety-four
were false; and the seedlings of six varieties of D. ajacis were true in
the same manner and degree as with the stocks above described. A
distinguished botanist maintains that the annual species of Delphinium are
always self-fertilised; therefore I may mention that thirty-two flowers on
a branch of D. consolida, enclosed in a net, yielded twenty-seven capsules,
with an average of 17.2 seed in each; whilst five flowers, under the same
net, which were artificially fertilised, in the same manner as must be
effected by bees during their incessant visits, yielded five capsules with
an average of 35.2 fine seed; and this shows that the agency of insects is
necessary for the full fertility of this plant. Analogous facts could be
given with respect to the crossing of many other flowers, such as
carnations, etc., of which the varieties fluctuate much in colour.
As with flowers, so with our domesticated animals, no character is more
variable than colour, and probably in no animal more so than with the
horse. Yet, with a little care in breeding, it appears that races of any
colour might soon be formed. Hofacker gives the result of matching two
hundred and sixteen mares of four different colours with like-coloured
stallions, without regard to the colour of their ancestors; and of the two
hundred and sixteen colts born, eleven alone failed to inherit the colour
of their parents: Autenrieth and Ammon assert that, after two generations,
colts of a uniform colour are produced with certainty. (12/50. Hofacker
'Ueber die Eigenschaften' etc. s. 10.)]
In a few rare cases peculiarities fail to be inherited, apparently from the
force of inheritance being too strong. I have been assured by breeders of
the canary-bird that to get a good jonquil-coloured bird it does not answer
to pair two jonquils, as the colour then comes out too strong, or is even
brown; but this statement is disputed by other breeders. So again, if two
crested canaries are paired, the young birds rarely inherit this character
(12/51. Bechstein 'Naturgesch. Deutschlands' b. 4 s. 462. Mr. Brent, a
great breeder of canaries, informs me that he believes that these
statements are correct.): for in crested birds a narrow space of bare skin
is left on the back of the head, where the feathers are up-turned to form
the crest, and, when both parents are thus characterised, the bareness
becomes excessive, and the crest itself fails to be developed. Mr. Hewitt,
speaking of Laced Sebright Bantams, says (12/52. 'The Poultry Book' by W.B.
Tegetmeier 1866 page 245.) that, "why this should be so I know not, but I
am confident that those that are best laced frequently produce offspring
very far from perfect in their markings, whilst those exhibited by myself,
which have so often proved successful, were bred from the union of heavily-
laced birds with those that were scarcely sufficiently laced."
It is a singular fact that, although several deaf-mutes often occur in the
same family, and though their cousins and other relations are often in the
same condition, yet their parents are rarely deaf-mutes. To give a single
instance: not one scholar out of 148, who were at the same time in the
London Institution, was the child of parents similarly affected. So again,
when a male or female deaf-mute marries a sound person, their children are
most rarely affected: in Ireland, out of 203 children thus produced one
alone was mute. Even when both parents have been deaf-mutes, as in the case
of forty-one marriages in the United States and of six in Ireland, only two
deaf and dumb children were produced. Mr. Sedgwick (12/53. 'British and
Foreign Med.-Chirurg. Review' July 1861 pages 200-204. Mr. Sedgwick has
given such full details on this subject, with ample references, that I need
refer to no other authorities), in commenting on this remarkable and
fortunate failure in the power of transmission in the direct line, remarks
that it may possibly be owing to "excess having reversed the action of some
natural law in development." But it is safer in the present state of our
knowledge to look at the whole case as simply unintelligible.
Although many congenital monstrosities are inherited, of which examples
have already been given, and to which may be added the lately recorded case
of the transmission during a century of hare-lip with a cleft-palate in the
writer's own family (12/54. Mr. Sproule in 'British Medical Journal' April
18, 1863.) yet other malformations are rarely or never inherited. Of these
latter cases, many are probably due to injuries in the womb or egg, and
would come under the head of non-inherited injuries or mutilations. With
plants, a long catalogue of inherited monstrosities of the most serious and
diversified nature could easily be given; and with plants, there is no
reason to suppose that monstrosities are caused by direct injuries to the
seed or embryo.
With respect to the inheritance of structures mutilated by injuries or
altered by disease, it was until lately difficult to come to any definite
conclusion. Some mutilations have been practised for a vast number of
generations without any inherited result. Godron remarks (12/55. 'De
l'Espece' tome 2 1859 page 299.) that different races of man have from time
immemorial knocked out their upper incisors, cut off joints of their
fingers, made holes of immense size through the lobes of their ears or
through their nostrils, tatooed themselves, made deep gashes in various
parts of their bodies, and there is no reason to suppose that these
mutilations have ever been inherited. (12/56. Nevertheless Mr. Wetherell
states, 'Nature' December 1870 page 168, that when he visited fifteen years
ago the Sioux Indians, he was informed "by a physician, who has passed much
of his time with these tribes, that some times a child was born with these
marks. This was confirmed by the U.S. Government Indian Agent.") Adhesions
due to inflammation and pits from the small-pox (and formerly many
consecutive generations must have been thus pitted) are not inherited. With
respect to Jews, I have been assured by three medical men of the Jewish
faith that circumcision, which has been practised for so many ages, has
produced no inherited effect. Blumenbach, however, asserts (12/57.
'Philosoph. Mag.' volume 4 1799 page 5.) that Jews are often born in
Germany in a condition rendering circumcision difficult, so that a name is
given them signifying "born circumcised;" and Professor Preyer informs me
that this is the case in Bonn, such children being considered the special
favourites of Jehovah. I have also heard from Dr. A. Newman, of Guy's
Hospital, of the grandson of a circumcised Jew, the father not having been
circumcised, in a similar condition. But it is possible that all these
cases may be accidental coincidence, for Sir J. Paget has seen five sons of
a lady and one son of her sister with adherent prepuces; and one of these
boys was affected in a manner "which might be considered like that commonly
produced by circumcision;" yet there was no suspicion of Jewish blood in
the family of these two sisters. Circumcision is practised by Mahomedans,
but at a much later age than by Jews; and Dr. Riedel, Assistant Resident in
North Celebes, writes to me that the boys there go naked until from six to
ten years old; and he has observed that many of them, though not all, have
their prepuces much reduced in length, and this he attributes to the
inherited effects of the operation. In the vegetable kingdom oaks and other
trees have borne galls from primeval times, yet they do not produce
inherited excrescences; and many other such facts could be adduced.
Notwithstanding the above several negative cases, we now possess conclusive
evidence that the effects of operations are sometimes inherited. Dr. Brown-
Sequard (12/58. 'Proc. Royal Soc.' volume 10 page 297. 'Communication to
the Brit. Assoc.' 1870. 'The Lancet' January 1875 page 7. The extracts are
from this last paper. It appears that Obersteiner 'Stricker's Med.
Jahrbucher' 1875 No. 2 has confirmed Brown-Sequard's observations.) gives
the following summary of his observations on guinea-pigs; and this summary
is so important that I will quote the whole:--
["1st. Appearance of epilepsy in animals born of parents having been
rendered epileptic by an injury to the spinal cord.
"2nd. Appearance of epilepsy also in animals born of parents having been
rendered epileptic by the section of the sciatic nerve.
"3rd. A change in the shape of the ear in animals born of parents in which
such a change was the effect of a division of the cervical sympathetic
nerve.
"4th. Partial closure of the eyelids in animals born of parents in which
that state of the eyelids had been caused either by the section of the
cervical sympathetic nerve or the removal of the superior cervical
ganglion.
"5th. Exophthalmia in animals born of parents in which an injury to the
restiform body had produced that protrusion of the eyeball. This
interesting fact I have witnessed a good many times, and I have seen the
transmission of the morbid state of the eye continue through four
generations. In these animals, modified by heredity, the two eyes generally
protruded, although in the parents usually only one showed exophthalmia,
the lesion having been made in most cases only on one of the corpora
restiformia.
"6th. Haematoma and dry gangrene of the ears in animals born of parents in
which these ear-alterations had been caused by an injury to the restiform
body near the nib of the calamus.
"7th. Absence of two toes out of the three of the hind leg, and sometimes
of the three, in animals whose parents had eaten up their hind-leg toes
which had become anaesthetic from a section of the sciatic nerve alone, or
of that nerve and also of the crural. Sometimes, instead of complete
absence of the toes, only a part of one or two or three was missing in the
young, although in the parent not only the toes but the whole foot was
absent (partly eaten off, partly destroyed by inflammation, ulceration, or
gangrene).
"8th. Appearance of various morbid states of the skin and hair of the neck
and face in animals born of parents having had similar alterations in the
same parts, as effects of an injury to the sciatic nerve."]
It should be especially observed that Brown-Sequard has bred during thirty
years many thousand guinea-pigs from animals which had not been operated
upon, and not one of these manifested the epileptic tendency. Nor has he
ever seen a guinea-pig born without toes, which was not the offspring of
parents which had gnawed off their own toes owing to the sciatic nerve
having been divided. Of this latter fact thirteen instances were carefully
recorded, and a greater number were seen; yet Brown-Sequard speaks of such
cases as one of the rarer forms of inheritance. It is a still more
interesting fact--
["That the sciatic nerve in the congenitally toeless animal has inherited
the power of passing through all the different morbid states which have
occurred in one of its parents from the time of the division till after its
reunion with the peripheric end. It is not therefore simply the power of
performing an action which is inherited, but the power of performing a
whole series of actions, in a certain order."]
In most of the cases of inheritance recorded by Brown-Sequard only one of
the two parents had been operated upon and was affected. He concludes by
expressing his belief that "what is transmitted is the morbid state of the
nervous system," due to the operation performed on the parents.
With the lower animals Dr. Prosper Lucas has collected a long list of
inherited injuries. A few instances will suffice. A cow lost a horn from an
accident with consequent suppuration, and she produced three calves which
were hornless on the same side of the head. With the horse, there seems
hardly a doubt that exostoses on the legs, caused by too much travelling on
hard roads, are inherited. Blumenbach records the case of a man who had his
little finger on the right hand almost cut off, and which in consequence
grew crooked, and his sons had the same finger on the same hand similarly
crooked. A soldier, fifteen years before his marriage, lost his left eye
from purulent ophthalmia, and his two sons were microphthalmic on the same
side. (12/59. This last case is quoted by Mr. Sedgwick in 'British and
Foreign Medico-Chirurg. Review' April, 1861 page 484. For Blumenbach see
above-cited paper. See also Dr. P. Lucas 'Traite de l'Hered. Nat.' tome 2
page 492. Also 'Transact. Linn. Soc.' volume 9 page 323. Some curious cases
are given by Mr. Baker in the 'Veterinary' volume 13 page 723. Another
curious case is given in the 'Annales des Scienc. Nat.' 1st series, tome 11
page 324.) In all cases in which a parent has had an organ injured on one
side, and two or more of the offspring are born with the same organ
affected on the same side, the chances against mere coincidence are almost
infinitely great. Even when only a single child is born having exactly the
same part of the body affected as that of his injured parent, the chances
against coincidence are great; and Professor Rolleston has given me two
such cases which have fallen under his own observation,--namely of two men,
one of whom had his knee and the other his cheek severely cut, and both had
children born with exactly the same spot marked or scarred. Many instances
have been recorded of cats, dogs, and horses, which have had their tails,
legs, etc., amputated or injured, producing offspring with the same parts
ill-formed; but as it is not very rare for similar malformations to appear
spontaneously, all such cases may be due to coincidence. It is, however, an
argument on the other side that "under the old excise laws the shepherd-dog
was only exempt from tax when without a tail, and for this reason it was
always removed" (12/60. 'The Dog' by Stonehenge 1867 page 118.); and there
still exist breeds of the shepherd-dog which are always born destitute of a
tail. Finally, it must be admitted, more especially since the publication
of Brown-Sequard's observations, that the effects of injuries, especially
when followed by disease, or perhaps exclusively when thus followed, are
occasionally inherited. (12/61. The Mot-mot habitually bites the barbs off
the middle part of the two central tail-feathers, and as the barbs are
congenitally somewhat reduced on the same part of these feathers, it seems
extremely probable, as Mr. Salvin remarks, 'Proc. Zoolog. Soc.' 1873 page
429, that this is due to the inherited effects of long-continued
mutilation.)
CAUSES OF NON-INHERITANCE.
A large number of cases of non-inheritance are intelligible on the
principle, that a strong tendency to inheritance does exist, but that it is
overborne by hostile or unfavourable conditions of life. No one would
expect that our improved pigs, if forced during several generations to
travel about and root in the ground for their own subsistence, would
transmit, as truly as they now do their short muzzles and legs, and their
tendency to fatten. Dray-horses assuredly would not long transmit their
great size and massive limbs, if compelled to live on a cold, damp
mountainous region; we have indeed evidence of such deterioration in the
horses which have run wild on the Falkland Islands. European dogs in India
often fail to transmit their true character. Our sheep in tropical
countries lose their wool in a few generations. There seems also to be a
close relation between certain peculiar pastures and the inheritance of an
enlarged tail in fat-tailed sheep, which form one of the most ancient
breeds in the world. With plants, we have seen that tropical varieties of
maize lose their proper character in the course of two or three
generations, when cultivated in Europe; and conversely so it is with
European varieties cultivated in Brazil. Our cabbages, which here come so
true by seed, cannot form heads in hot countries. According to Carriere
(12/62. 'Production et Fixation des Varietes' 1865 page 72.) the purple-
leafed beech and barberry transmit their character by seed far less truly
in certain districts than in others. Under changed circumstances,
periodical habits of life soon fail to be transmitted, as the period of
maturity in summer and winter wheat, barley, and vetches. So it is with
animals: for instance, a person, whose statement I can trust, procured eggs
of Aylesbury ducks from that town, where they are kept in houses and are
reared as early as possible for the London market; the ducks bred from
these eggs in a distant part of England, hatched their first brood on
January 24th, whilst common ducks, kept in the same yard and treated in the
same manner, did not hatch till the end of March; and this shows that the
period of hatching was inherited. But the grandchildren of these Aylesbury
ducks completely lost their habit of early incubation, and hatched their
eggs at the same time with the common ducks of the same place.
Many cases of non-inheritance apparently result from the conditions of life
continually inducing fresh variability. We have seen that when the seeds of
pears, plums, apples, etc., are sown, the seedlings generally inherit some
degree of family likeness. Mingled with these seedlings, a few, and
sometimes many, worthless, wild-looking plants commonly appear, and their
appearance may be attributed to the principle of reversion. But scarcely a
single seedling will be found perfectly to resemble the parent-form; and
thus may be accounted for by constantly recurring variability induced by
the conditions of life. I believe in this, because it has been observed
that certain fruit-trees truly propagate their kind whilst growing on their
own roots; but when grafted on other stocks, and by this process their
natural state is manifestly affected, they produce seedlings which vary
greatly, departing from the parental type in many characters. (12/63.
Downing 'Fruits of America' page 5: Sageret 'Pom. Phys.' pages 43, 72.)
Metzger, as stated in the ninth chapter, found that certain kinds of wheat
brought from Spain and cultivated in Germany, failed during many years to
reproduce themselves truly; but at last, when accustomed to their new
conditions, they ceased to be variable,--that is, they became amenable to
the power of inheritance. Nearly all the plants which cannot be propagated
with any approach to certainty by seed, are kinds which have been long
propagated by buds, cuttings, offsets, tubers, etc., and have in
consequence been frequently exposed during what may be called their
individual lives to widely diversified conditions of life. Plants thus
propagated become so variable, that they are subject, as we have seen in
the last chapter, even to bud-variation. Our domesticated animals, on the
other hand, are not commonly exposed during the life of the individual to
such extremely diversified conditions, and are not liable to such extreme
variability; therefore they do not lose the power of transmitting most of
their characteristic features. In the foregoing remarks on non-inheritance,
crossed breeds are of course excluded, as their diversity mainly depends on
the unequal development of character derived from either parent or their
ancestors.
CONCLUSION.
It has been shown in the early part of this chapter how commonly new
characters of the most diversified nature, whether normal or abnormal,
injurious or beneficial, whether affecting organs of the highest or most
trifling importance, are inherited. It is often sufficient for the
inheritance of some peculiar character, that one parent alone should
possess it, as in most cases in which the rarer anomalies have been
transmitted. But the power of transmission is extremely variable. In a
number of individuals descended from the same parents, and treated in the
same manner, some display this power in a perfect manner, and in some it is
quite deficient; and for this difference no reason can be assigned. The
effects of injuries or mutilations are occasionally inherited; and we shall
see in a future chapter that the long-continued use and disuse of parts
produces an inherited effect. Even those characters which are considered
the most fluctuating, such as colour, are with rare exceptions transmitted
much more forcibly than is generally supposed. The wonder, indeed, in all
cases is not that any character should be transmitted, but that the power
of inheritance should ever fail. The checks to inheritance, as far as we
know them, are, firstly, circumstances hostile to the particular character
in question; secondly, conditions of life incessantly inducing fresh
variability; and lastly, the crossing of distinct varieties during some
previous generation, together with reversion or atavism-that is, the
tendency in the child to resemble its grand-parents or more remote
ancestors instead of its immediate parents. This latter subject will be
discussed in the following chapter.
END OF VOLUME I.
End of this Project Gutenberg Etext of The Variation of Animals and Plants
under Domestication Volume I by Charles Darwin
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