OH if THE VARIATION a tee : “ANIMALS AND PLANTS DOMESTICATION. By CHARLES DARWIN, MA, F.RS., &. SECOND EDITION, REVISED. FOURTH THOUSAND. IN TWO VOLUMES.—Vo.. I. WITH ILLUSTRATIONS. ee 2 OVULI y,f NEW YORK: Dp APPLETON AND COMPANY, Ps, any 5 BOND STREET, 1894. PREFACE TO THE SECOND EDITION. —_+*«> —- Dourixe the seven years which have elapsed since the pub- lication 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 addi- tional facts, chiefly through the kindness of many corre- spondents. 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 dis- covery 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. PRR ODWCTION s c.50 1s tee. we ee ees ee ee) Paes 4 CHAPTER LI. 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 VARIABI- LITY Se ae aida Se aes, CO Siena tba (ho) ieee oe Re am eee Ee) CHAPTER II. HORSES AND ASSES. BORSE.—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 V1 CONTENTS. THE HOKSE—DAPPLING—DARK STRIPES ON THE SPINE, LEGS, SHOULDERS, AND FOREHEAD—DUN-COLOURED HORSES MOST FRE- QUENTLY 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. Pages 51-67 CHAPTER III. PIGS—CATTLE—SHEEP— GOATS. PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICUS— TORFSCHWEIN—JAPAN PIGS—FERTILITY OF CROSSED PIGS—CHANGES IN THE SKULL OF THE HIGHLY CULTIVATED RACES—CONVERGENCE OF CHARACTER—GESTATION—SOLID-HOOFED SWINE—CURIOUS AP- PENDAGES TO THE JAWS—DECREASE IN SIZE OF THE TUSKS— YOUNG PIGS LONGITUDINALLY STRIPED—FERAL PIGS—CROSSED BREEDS. CATTLE.—zeEsu 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 ~! Vee SS ee CHAPTER IY. 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— YERAL RABBITS IN JAMAICA AND THE FALKLAND ISLANDS—PORTO SANTO FERAL RABBITS — OSTEOLOGICAL CHARACTERS -— SKUiI-- CONTENTS. Vli SKULL OF HALF-LOP RABBITS — VARIATIONS IN THE SKULL ANALAGOUS TO DIFFERENCES IN DIFFERENT SPECIES OF HARES— VERTEBR4&—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... .. Pages 107-136 CHAPTER V. DOMESTIC PIGEONS. ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS—INDIVIDUAL VARIABILITY—VARIATIONS OF A REMARKABLE NATURE—OSTEO- LOGICAL CHARACTERS: SKULL, LOWER JAW, NUMBER OF VERTEBR& —CORRELATION 0! 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 CORRE- LATION WITH LENGTH OF BEAK—LENGTH OF STERNUM, SCAPULA, AND FURCULUM—LENGTH OF WINGS—SUMMARY ON THE POINTS OF DIFFERENCE IN THE SEVERAL BREEDS .. . « 137-188 CHAPTER VI. PIGEONS—continued. THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES —HABITS OF LIFE—WILD RACES OF THE ROCK-PIGEON—DOVECOT- PIGEONS—PROOFS OF THE DESCENT OF THE SEVERAL RACES FROM COLUMBA LIVIA—FERTILITY OF THE RACES WHEN CROSSED—RBE- VERSION TO THE PLUMAGE OF THE WILD ROCK-PIGEON—CIRCUM- STANCES FAVOURABLE TO THE FORMATION OF THE RACES—AN- TIQUITY AND HISTORY OF THE PRINCIPAL RACES—-MANNER OF THEIR FORMATION — SELECTION — UNCONSCIOUS SELECTION — SARE TAKEN BY FANCIERS IN SELECTING THEIR BIRDS—SLIGHTLY DIF- FERENT STRAINS GRADUALLY CHANGE INTO WELL-MARKED BREEDS —EXTINCTION OF INTERMEDIATE FORMS—CERTAIN BREEDS REMAIN PERMANENT, WHILST OTHERS CHANGE—SUMMARY.. .. 189-28h Vill CONTENTS, CHAPTER Vt. FOWLS. BRIEF DESCRIPTIONS OF THE CHIEF BREEDS—ARGUMENTS IN FAVOUB 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 DIF- FERENCES BETWEEN THE SEVERAL BREEDS — EGGS — CHICKENS— SECONDARY SEXUAL CHARACTERS—WING- AND TAIL- FEATHERS, VOICE, DISPOSITION, ETC.—OSTEOLOGICAL DIFFERENCES IN THE SKULL, VERTEBRZ, ETC.—EFFECTS OF USE AND DISUSE ON CERTAIN PARTS—CORRELATION OF GROWTH -- « e Pages 2386-289 CHAPTER VIII. DUCK—GOOSE—PEACOCK—TURKEY—GUINEA-FOW L— 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 DIF- FERENT BREEDS—OSTEOLOGICAL DIFFERENCES—EFFECTS OF USE AND DISUSE CN THE LIMB-BONES. GOOSE, ANCIENTLY DOMESTICATED—LITTLE VARIATION OF-—SEBAS- TOPOL 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, sPpecIES AND BREEDS OF—ANCIENTLY DOMESTICATED —CARE IN THEIR SELECTION—DIFFERENCES IN THE DiFFERENT RACES—IN THE EGG, CATERPILLAR, AND COCOON STATES—INHERIT- ANCE OF CHARACTERS—IMPERFECT WINGS —LOST INSTINCTS—CORRE- Paw CHARACTERS, “see>! 23° cue “yao! ye eee. oe ee ee CONTENTS. 1X CHAPTER IX. CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS. PRELIMINARY REMARKS on THE NUMBER AND PARENTAGE OF CULTIVATED PLANTS—FIRST STEPS IN CULTIVATION—GEOGRAPHICAL DISTRIBUTION OF CULTIVATED PLANTS. CEREALIA.—pouztTs 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 CON- STANT, SOME HIGHLY VARIABLE—DO NOT INTERCROSS.——BEANS. ——POTATOES : NUMEROUS VARIETIES OF—DIFFER LITTLE EXCEPT IN THE TUBERS—CHARACTERS INHERITED .. .. Pages 322-351 CHAPTER X. PLANTS continued—FRUITS—ORNAMENTAL TREES— FLOWERS, FRUITS—cGRAPES—VARY IN ODD AND TRIFLING PARTICULARS.—— MULBERRY—THE ORANGE GROUP—SINGULAR RESULTS FROM CROSS- ING.——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.——CUCURBIT'A- CEOUS PLANTS—WONDERFUL VARIATION OF. ORNAMENTAL TREES—tTHEIR VARIATION IN DEGREE AND KIND — ASH-TREE—SCOTCH-FIR—HAWTHORN. FLOW ERS—maUvLTIPLE ORIGIN OF MANY KINDS—VARIATION IN CON- STITUTIONAL PECULIARITIES —KIND OF VARIATION. ROSES — SEVERAL SPECIES CULTIVATED.——PANSY.——DAHLIA. HYA- PinTH HISTORY AND VARIATION OF ~ 3, 46° 2. 3... 302-396 4 CONTENTS. CHAPTER 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 CHARAC- TERS IN SEMINAL HYBRIDS BY BUD-VARIATION—ON THE DIRECT OR IMMEDIATE ACTION OF FOREIGN POLLEN ON THE MOTHER- PLANTI—ON THE EFFECTS OF A PREVIOUS IMPREGNATION ON THE SUBSEQUENT OFFSPRING OF FEMALE ANIMALS—CONCLUSION AND SUMMARY .. .. Soe. Vowel ee eee cd eon eRe CHAPTER XII. INHERITANCE. WONDERFUL NATURE OF INHERITANCE—PEDIGREES OF OUR DOMESTI- CATED 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—POLYDAC- TYLISM AND REGROWTH OF SUPERNUMERARY DIGITS AFTER AM- PUTATION—CASES OF SEVERAL CHILDREN SIMILARLY AFFECTED FROM NON-AFFECTED PARENTS—-WEAK AND FLUCTUATING IN- HERITANCE: 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 VARIA- BILITY, AND BY REVERSION—CONCLUSION .. ce «2 445-473 Fie. atin Gon Cotes oe LIST OF ILLUSTRATIONS. Dun DEVONSHIRE PoNY, WITH SHOULDER, SPINAL, AND LEG SPSS OR al Oe i nc ae aOR eee HEAD oF JAPAN OR MASKED Pia Bh Secs Loretta ara Heap oF Wiip Boar, anp or “GoLDEN Days,” A PIG gel THE: PORKSHIRE LARGE: BREED: 30-25 0+ aes es Oup IrnisH PIG WITH JAW-APPENDAGES oe eee a RPCEC TOP OEABEED teh ste ofr at | aha Lee we ee Eee ie SKULL OF WILD Rappit CUR Seo ome ane Meare nee SKULL OF TARGE: LOP-EARED RABBIT. «7 6 et) > Part oF ZYGOMATIC ARCH, SHOWING THE OF THE PROJECTING END MALAR BONE OF THE AUDITORY MEATUS, OF DP TSEEET Tt re tt aah, ae AN POSTERIOR END OF SKULL, SHOWING THE INTER-PARIETAL BONE, OF RABBITS Sen Beet Sean haa Se ren eee GeerrrraT, -BORAMEN OF FABBITS=*, 205,°° 25 iioty Sie SKULL oF HALF-LoP Raspir pith Sie RC ne ean AtGAS VERDEBR A OF RABBITS: <2" > tose Be See THirpD CERVICAL VERTEBRZ OF RABBITS .. .. .. DorsaAL VERTEBR&, FROM SIXTH TO TENTH INCLUSIVE, OF PUMP BUEN pay inet esas Sty OMe Mes. pra seaee tomas rae TERMINAL BoNE OF STERNUM OF RABBITS... .. .. .. ACROMION, OF SCAPULA OF RABBITS. fp) 66.02.58 ae Gee THE Rock-PicEon, orn CoLtumBa Livia Beas al Geer Uc RON GEIST EO PERM er, | \c)e% Sesh teem Leal hl Mon MotaT he Tap Mice PNG ES OAR RUDR Ge acpn caren kite a aaa a aaa Tees UPSETS SIE dT Derg tok ee ae Ie wa Pan One gran PNGE ASH DAN DAT ain ai-o ls sei aoe Few eRe ges, preg arta eae J ES AV ANAS 55 Coe ie ee RE Ra SEA Oe SR eRe Ra SHORT-FACED -ENGHISH. TUMPLER ..0 92. 056 wee oes FAGB b2. LIST OF ILLUSTRATIONS. SKULLS OF PIGEONS, VIEWED LATERALLY .. .. «+ ee LowER JAWS OF PIGEONS, SEEN FROM ABOVE .- «2 ee SKULL OF RUNT, SEEN FEOM ABOVE .. 3. <2 oe) es GATERAT, VIEW OF JAWS OF PIGEONS .. © 22 Je.) e= eee DEAPUIZ OF) PIGEONS © ..0 $c) oe ae eee EDRCULA OF PIGEONS .. 9 3i- 4. 4 See NEANISH “POW cs .s- 3.) ac be) ee (Se aAMBUReGH HOWL 2. 6.25. «sa eb eel ) eee ROMISH OWI c.9 ("sk yao Geos se ees, ne OcciPITAL FoRAMEN OF THE SKULLS OF FOWLS Ppa SKULLS OF FowLs, VIEWED FROM AROVE, A LITTLE OB- PAQUET Ve sca sic = Ses) ge Jlase! staat. a) 2 aa ee ee LONGITUDINAL SECTIONS OF SKULLS OF FOWLS, VIEWED PAUP UAT NOD wee Wee tae hn eae “sso! -Ge)) ecru SKULL OF HorNED FowL, VIEWED FROM ABOVE, A LITTLE OBIAGUELY (ye (te "Saki eeee oe: tea,” Gael ee ee StxTH CERVICAL VERTEBRZ OF FowLs, VIEWED LATERALLY EXTREMITY OF THE FurRCULA OF FOWLS, VIEWED LATER- AGEN Sean Rees nay spn) es. 00. - weg. eee ee SKULLS OF DUCKS, VIEWED LATERALLY, RELUCED TO TWO- THIRDS OF THE NATUBAL SIZE ..° , so) “subse Mee nee CERVICAL VERTEBRZ OF DUCKS, OF NATURAL SIZE .. .. Pons or wHE ‘Common’ PEA...) 623: a. Ges te PEACH AND ALMOND STONES, OF NATURAL SIZE, VIEWED EDGEWAYS ee ee ee ee ee os oe ee ee PLUM STONES, OF NATURAL SIZE, VIEWED LATERALLY .. (a 3 TABLE OF PRINCIPAL ADDITIONS AND CORRECTIONS. First Second | Edition. | Edition. ——— Vor. 1. | Vot. 1 Page Page a ; 34 350 | Dr. Burt Wilder’s observations on the brains of different breeds of the Dog. 38 40 | Degeneracy of Dogs imported into Guinea. dl 54 | Difference in the number of the lumbar vertebre in the races or species of the Horse. 102 | 106 | Hairy appendages to the throats of Goats. 162 | 170 | Sexual differences in colour in the domestic Pigeon. Y17. | 228 | Movements like those of the Tumbler-pigeon, | caused by injury to the brain. 990 | 806 ! Additional facts with respect to the Black-shouldered Peacock. 996 | 312 | Ancient selection of Gold-fish in China. 314 | 332 | Major Hallett’s ‘Pedigree Wheat.’ 326 | 3845 | The common radish descended from Raphanus raph- | anistrum. 274 | 898 | Several additional cases of bud-variation given. 396 | 420 | An abstract of all the cases recently published of graft-hybrids in the potato, together with a gene- ral summary on graft-hybridisation. 399 | 429 | An erroneous statement with respect to the pollen of the date-palm affecting the fruit of the Cham- gerops omitted. 400 | 480 | New cases of the direct action of pollen on the mother-plant. 404 | 485 | Additional and remarkable instances of the action | | of the male parent on the future progeny of the female. An erroneous statement corrected, with respect to the regrowth of supernumerary digits after am- putation. | ( xiv) TABLE OF PRINCIPAL ADDITIONS AND CORRECTIONS. ’ First | Second Edition.| Edition. VOL. al Vot. I. Page Page 23 23 | 800 | 94 | 469 | Vor. IL} AD Ue 72 48 105 83 120 99 123 | 103 135 117 to to 141 | 122 149 | ok 152 | 134 IA) iS 273 | 262 317 | 309 324 | 316 to | to O28 | 327 | 309 | 83 357 | 349 to to 404 | 399 ( Continued.) i | ! Additional facts with respect to the inherited effects | of circumcision. | Dr. Brown-Séquard on the inherited effects of opera- | tions on the Guinea-pig. | Other cases of inherited mutilations. | An additional case of reversion due to a cross. Inheritance as limited by sex: Two varieties of maize which cannot be neers. Some additional facts on the advantages of cross- breeding in animals. Discussion on the effects of close interbreeding in | the case of man. | Additional cases of plants sterile with pollen from | the same plant. | Mr. Sclater on the infertility of animals under con- finement. The Aperea a distinct species from the Guinea-pig. Prof. Jiger on hawks killing light-coloured pigeons. | Prof. Weismann on the effects of isolation in the | development of species. | The direct action of the conditions of life in causing variation. Mr. Romanes on rudimentary parts. Some additional cases of correlated variability. On Geoffroy St. Hilaire’s law of “ soi pour sot.” | The chapter on Pangenesis has been largeiy altered and re-modelled; but the essential principles re- | main 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 col- lect 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 varia- tion. 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 itis nota little difficult to decide how they can be bestarranged. 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. When- ever I have found it necessary to give numerous details, in support of any proposition or conclusion, small type has been 2 INTRODUCTION. used. The reader will, I think, find this plan a convenience. for, if he does not doubt the conclusion or care about the details, he can easily pass them over; vet Imay 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 nething about Natural Selection, if I here give a brief sketch of the whole subject and of its bearing on the origin of species.! This is the more desirable, as it is impossible in the present werk 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 aman 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.” He unintentionally exposes his animals and plants to various conditions of hfe, and varia- bility 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 conse- quently 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 tinued ill-health. 2 M. Pouchet has recently (‘ Plural- ity of Races,’ Eng. Translat., 1864, p. 83, &c.) insisted that variation under 1 To any one who has attentively read my ‘Origin of Species’ this lutro- duction 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 con- domestication throws no light on the natural modification of species. J cannot perceive the force of his a1zu- ments, or, to speak more accurately, of his assertions to this effect. NATURAL SELECTION. 5) alluvial flat; and lastly,it has been exposed to changes in its conditions, being grown sometimes in one district and some- times 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 hfe, 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 ap- parently 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, 4 INTRODUCTION. 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 uncon- sciously and unintentionally. Man may select and preserve each successive variation, with the distinct intention of im- proving and altering a breed, in accordance with a precon- ceived idea; and by thus adding up variations, often so slight as to be imperceptible by an uneducated eye, he has eifected 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 scme varieties,—that their organization is NATURAL SELECTION. i in some slight degree plastic; granting that many aninals 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 in- herited races; granting all this, how, it may be asked, have species arisen in a state of nature? The differences between natural varieties are sight; whereas the differences are con- siderable 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 | have called them incipient species, become converted into true and well-defined species ? How has each new species been adapted to the surrounding physical con- ditions, and to the other forms of life on which it in any way depends? We see on every side of us innumerable adapta- tions ana contrivances, which have justly excited the highest admiration of every observer. There is, for instance, a fly (Cecidomyia)? 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 (Miso- campus) 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 itis, 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 character- istic 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 ot 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, ? Léon Dufour in ‘ Annales des Scienc. Nat.’ (3rd series, Zoolog.), tom. v. p. 6. 6 INTRODUCTION. 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 nbundant 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 occur- rence 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 preserva- tion of varieties and species ina 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 attrac- tion of gravity as ruling the movements of the planets, or as agriculturists speak cf man making domestic races by his NATURAL SELECTION. q 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 | 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. lt 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 inhabi- tants. 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 character- istic of species, and, by the extermination of the older inter- mediate 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 organization occasionally varies in some slight degree, and as natural selection acts exclusively by the preservation of variations which are advantageous under the excessively complex conditions to which each being is exposed, no limit 8 INTRODUCTION. exists to the number, singularity, and perfection of the contrivances and co-adaptations which may thus be pro- duced. An animal or a pliant 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 organiza- tion 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 ad- vancement in the scale of organization. We are almost compelled to look at the specialization or differentiation of parts or organs for different functions as the best or even sole standard of advancement; for by such division of labour each function of body and mind is better performed. And as natural selection acts exclusively through the preservation of profitable modications of structure, and as the conditions of life in each area generally become more and more complex from the increasing number of different forms which inhabit it and from most of these forms acquiring a more and more perfect structure, we may confidently believe, that, on the whole, organization advances. Nevertheless a very simple form fitted for very simple conditions of life might remain for indefinite ages unaltered or unimproved ; for what would it profit an infusorial animalcule, for instance, or an intestinal worm, to become highly organized? Members of a high group might even become, and this apparently has often occurred, fitted for simpler conditions of life; and in this case natural selection would tend to simplify or degrade the organization, for complicated mechanism for simple actions would be useless or even disadvantageous. 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 NATURAL SELECTION, 9 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 typothesis, 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 hypo- thetical, 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 Beers Arehipelago, situated in the Pacific Ocean about 10 _ INTRODUCTION. 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 realized this idea, I saw, on reading Malthus on Population, that Natural Selection was the inevitable result of the rapid increase of all organic beings ; for I was prepared to appreciate the struggle for existence by having long studied the habits of animals. Before visiting the Galapagos I had collected many animals whilst travelling from north to south on both sides of America, and everywhere, under conditions of life as different as it is possible to conceive. American forms were met with—species replacing species of the same peculiar genera. ‘Thus it was when the Cordilleras were ascended, or the thick tropical forests penetrated, or the fresh waters of America searched. Subsequently I visited other countries, which in all thew NATURAL SELECTION. 11 conditions of life were incomparably more lke parts of South America, than the diferent 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 succes- sion of allied forms had been previously observed in Australia. Here then we see the prevalence, as if by descent, in time as in space, of the same types in the same areas ; and in neither case does the similarity of the conditions by any means seem sufficient to account for the similarity of the forms of life. It is notorious that the fossil remains of closely consecutive formations are closely allied in structure, and we can at once understand the fact if they are 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 modifiation 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 ruleg ee 12 INTRODUCTION. 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 mean 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 erouped 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; thongh this more simple NATURAL SELECTION, 13 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 froma single ancestor. A number of organic beings are included in the same class, because they present, independently of their habits of life, the same funda- mental 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 pro- genitor. 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, &¢e.—on which we are confessedly ignorant; nor do we know how ignorant we are. If we are much more ignorant thin 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 14 INTRODUCTION, the theory, because it collects, under one point of view, and gives a rational explanation of, many apparently independez:t classes of facts.* * In treating the several subjects included in the present and my other works I have continually been led to ask for information from many zoolo- gists, botanists, geologists, breeders of animals, and horticulturists, and I have invariably received from them the most generous assistance. With- out 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 re- siding 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. Ouar. 1. DOGS: THEIR PARENTAGE. 15 CHAPTER I. DOMESTIC DOGS AND CATS. ANCIENT VARIETIES OF THE DOG—RESEMBLANCE OF DOMESTIC DOGS IN VARIOUS COUNTRIES TO NATIVE CANINE SPECIES—ANIMALS NOT ACQUAINTED WITH MAN AT FIRST FEARLESS—DOGS RESEMBLING WOLVES AND JACKALS—HABIT OF BARKING ACQUIRED AND LOST—FERAL DOGS—TAN-COLOURED EYE-SPOTS —PERIOD OF GESTATION—OFFENSIVE ODOUR—FERTILITY OF THE RACES WHEN (ROSSED—DIFFERENCES IN THE SEVERAL RACES IN PART DUE TO DESCENT FROM DISTINCT SPECIES—DIFFERENCES IN THE SKULL AND TEETH—DIFFER- ENCES 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—EXTINOTION 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 VAR{ABILITY. Tae 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. Paleontology! does not throw much light on the question, owing, on the one hand, to the close similarity of the skulls of extinct as well as living wolves and jackals, and owing, on the other hand, to 1 Owen, ‘ British Fossil Mammals,’ pp- 123 to 133. Pictet’s ‘ Traité de Pal.,’ 1853, tom. i. p. 202, De Blain- ville, in his ‘Ostéographie, Canide,’ habits. See also Boyd Dawkins, ‘Cave Hunting,’ 1874, p. 131, &c., and his other publications. Jeitteles has discussed in great detail the p- 142, has largely discussed the whole subject, and concludes that the extinct parent ofall domesticated dogs came nearest to the wolf in organization, and to the jackal in character of the breeds of pre-historic dogs: ‘Die vorgeschichtlichen Alter- thiimer der Stadt Olmiitz,’ 1. Theii, 1872, p. 44 to ead. 16 DOGS. Cuap. I. 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 ex- tinct 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 ne cessarily 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.?. 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 ditferences between the several races of various domesticated animals which cer- tainly 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. force than the late James Wilson, of Edinburgh, in various papers read before the Highland Agricultural and ? Pallas, I believe, originated this doctrine in ‘Act. Acad. St. Peters- burgh,’ 1780, Part ii. Ehrenberg has advocated it, as may be seen in De Blainville’s ‘O-téographie,’ p. 79. It has been carried to an extreme extent by Col. Hamilton Smith in the ‘Naturalist Library,’ vols. ix. and x. Mr. W. C.. Martin adopts it in his excellent ‘ History of the Dog, 1845; as does Dr. Morton, as well as Nott and Gliddon, in the United States. Prof. Low, in his ‘ Domesticated Animals,’ 1845, p. 656, comes to this same conclusion. No one has argued on this side with more clearness and Wernerian Societies. Isidore Geoffroy Saint-Hilaire (* Hist. Nat. Gén.,’ 1860, tom. iil p. 107), though he believes that most dogs have descended from the jackal, yet inclines to the belief that some are descended from the wolf. Prof. Gervais (‘Hist. Nat. Mamm.’ 1855, tom. ii. p. 69, referrin z to the view that all the domest-< races are the modified descendants of a single species, after a long discussion, says, “Cette opinion est, suivant nous du moins, la moins protable.” Cuap, L. THEIR PARENTAGE. Livi The materials are remarkably deficient between the four- teenth century and the Roman classical period.? At this latter period various breeds, namely hounds, house-dogs, lap- dogs, &c., existed; but, as Dr. Walther has remarked, it is eens to recognise the greater number with any cer- tainty. Youatt, however, gives a drawing of a beautiful sculpture of two greyhound puppies from the Villa of An- toninus. On an Assyrian monument, about 640 B.c., an | enormous mastiff* is figured; and according to Sir H. Rawlinson (as I was informed at the British Museum), similar dogs are still imported into this same country. I have looked through the magnificent works of Lepsius and Rosellini, and on the Egyptian monuments from the fourth to the twelfth dynasties (7.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® 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 from the tomb of the son of Esar Haddor, and clay models in the British Museum. Nott and Gliddon, in their ‘Types of Mankind,’ 1854, p. 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 Kultur- geschichte,’ Leipzig, 1852, s. 115 ‘Youatt on the Dog,’ 1845, p.6. Quoted by Col. Ham. Smith in © Nat. Lib.,’ vol. x. p. 79. 6° De Blainville, ‘Ostéographie, Canidae,’ p. 134. F. Cuvier, ‘ Annales du Muséum.’ tom. xviii. p. 342. In regard to mastitts, see Col. H. Smith, eNaAt. lib, vol. x. p. 218) For the Thibet mastiff, see Mr. Hodgson in ‘Journal of As. Soc, of Bengal,’ vol. i., 1832, p. 342. 87 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.” ¢ Pfahlbauten,’ s. 163, et passim. 7 Stan. Julien, quoted by de Blain- 6 SeeJ.W.Schiitz’ interesting essay, ville, ‘Ostéographie,’ p. 163. Zur Kenntniss des Torfschweins,’ 8 Richardson, ‘Pigs, their Origin,’ 1868. This author believes that the M&c., p. 26. Torfschwein is descended from a 9 ‘Die Racen des Schweines,’ s. 47, distinct species, the S. sennariensis of 64. Central Atrica. 72 DOMESTIC PIGS. Cuap. ITT, 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 (Schweine- schadel, s. 138), that the infusion of the 34nd, or even of the oth, 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, &c., 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. 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, oreat fleshy ears, and deeply furrowed skin. The following woodcut is copied from that given by Mr. Bartlett!° Not only is the face furrowed, but thick folds of skin, which are harder than the other parts, almost lke 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." Dr. Gray !* 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 domesti- cated variety of S. indicus: if this really be the case, it isa 10 «Proc. Zoolog. Soc.,’ 1861, p. 263. in a very interesting essay, ‘ Der 1l Sclater, in ‘Proc. Zoolog. Scc.,’ Schidel des Maskenschweines,’ 1870. Feb. 26th, 1861. He confirms the conclusion of yen 12 ¢ Proc. Zoolog. Soc.,’ 1862, p. 13. | Nathusius on the relationship of this The skull has since been described kind of pig. much more fully by Professor Lucae Cuap. III. THEIR VARIATION, Ge 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 !* as of small size, hump- backed, with a disproportionately long head, with short ears Wz zt Zz ; i 4 he { bull HN Fig. 2.—Head of Japan or Masked Pig. (Copied from Mr. Bartlett’s paper in ‘ Proc. Zoolog. Soc.’ 1861, p. 263.) 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 13 ¢ Journal of Voyages and Travels from 1821 to 1829,’ vol. i. p. 300. 74 DOMESTIC PIGS. Cuap. IIL expected, seera 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.” 1* 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 un- impaired fertility. One great breeder who had used pure Chinese pigs assured me that the fertility of the half-breeds inter se and of their recrossed progeny was actually increased ; and this is the general belief of agriculturists. Again, the Japan pig or S. pliciceps of Gray is so distinct in appearance from all common pigs, that it stretches one’s belief to the utmost to admit that it is simply a domestic variety; yet this breed has been found perfectly fertile with the Berkshire breed ; and Mr. Eyton informs me that he paired a half-bred brother and sister and found them quite fertile together. The 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 1# Rey. G. Low, ‘ Fauna Oreadensis,’ p. 19. See also Dr. Hibbert’s account of the pig of the Shetland Islands Cuap. III. THEIR VARIATION. res) and various other modifications, as Nathusius observes, can hardly be considered as monstrosities, for they are not in- jurious, and are strictly inherited. ‘The whole head is much shortened ; thus, whilst in common breeds its length to that of the body is as | to 6, in the “cultur-racen ” the proportion is as 1 to 9, and even recently as 1 to11.% The fol- lowing woodcut! of the head of a wild boar and of a sow from a photo- graph 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 ¥iz.3—Head of Wild Boar, and of “Gold-n Days,” a 7 U7 TN, a pig of the York hire Large Breed; the latter from a ty eZ Nathusius states pho'ograph.. (Copied from Sidney’s edit. of ‘The positively (s. 99, 103), Piss’ by Youa.t.) ‘NK i \ ANI \ MANN ik li Mi as the result of common experience and of his experiments, = ‘ Die Racen des Schweines,’ s. 70 excellent edition of ‘The Pig,’ by These woodcuts are copied from Youatt, 1860. See pp. 1, 16, 19. engravings given in Mr. S. Sidney’s 17 «Schweineschadel,’ s. 74, 135. 76 DOMESTIC PIGS. Cuap. IT. 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 miata 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. Miller has shown, this seems caused by an abnormal state of the pri- mordial cartilage. We may, however, readily admit that abundant and rich food supplied during many generations would give an inherited tendency to increased size of body, and that, from disuse, the limbs would become finer and shorter.1* 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 in- teresting 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 18 Nathusius, ‘ Vie Racen des Schweines,’ s. 71. Quap. LIL. THEIR VARIATION. Tie partly to man breeding the pig for one sole purpose, namely, for the greatest amount of flesh and fat; so that selection has always tended towards one and the same end. With most domestic animals the result of selection has been divergence of character, here it has been convergence.!® The nature of the food supplied during many generations has apparently affected the length of the intestines; for, according to Cuvier,”® tkeir 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 tol. 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 mamme vary, as does the period of gestation. ‘The latest authority says?! that “the period averages from 17 to 20 veeks,” but I think there must be some error in this state- ment: 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 eaily to ma- turity ; 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,?? 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 vertebre and ribs in different kinds of pigs, as observed by Mr. Eyton,” and as given in the following table, has often been quoted. The African sow probably belongs to the S. scrofa type; and Mr. 19 «Tie Racen des Schweines,’s. 47. ‘ Schweineschadel,’ s. 104. Compare, also, the figures of the old Irish and the improved Irish breeds in Richard- son on ‘ The Pig,’ 1847. 20 Quoted by Isid. Geoffroy, ‘ Hist. Nat. Gén.,’ tom. iii. p 441. 21 S. Sidney, ‘The Pig,’ p. 61. 22 ¢Schweineschadel,’ s. 2, 20. 23 “Proc. Zoolog. Soc.,’ 1837, p. 23. I have not given the caudal vertebre, as Mr. Eyton says some mizht possibly have been lost. Ihave added together the dorsal and lumbar vertebre, owing to Prof. Owen’s remarks (‘ Journal Linn. Soc. vol. ii. p. 28) on the difter- ence between dorsal and lumbar vertebra depending only on the Jevelopment of the ribs. Nevertheless the difference in the number ot the ribs in pigs deserves notice. M. Sanson gives the number of lumbar vertebre in various pigs; ‘Comptes Rendus,’ Ixiii. p. 843. 78 DOMESTIC PIGS. Cura. IIT. 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. | English | ,-: 3) French Long-legg oa African Chinese Wild Boar | Domestic Male. | Female. Male. | from Cuvier. | Boar, from Pt ee at So Cuvier. Dorsal vertebre . 15 eee | 14 | 1t Lumbar.. 16 | te Dorsal and lta) | : together .. i 9 19 | 19 | 19 Sacral 5) 5 + | 4 | “ Total number of 26 °4 93 OC 93 | 23 vertebrze 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** 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. 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 cylin- 4 ¢Edinburgh New Philosoph. Blainville’s ‘ Ostéographie,’ p. 128, for Journal,’ April, 1863. See also De various authorities on this subject. Cuap. IIL THEIR VARIATION. 79 drical, 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 longi- tudinal 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 Nathu- sius states that they occasionally appear im all the long eared Fig. 4.—Oid Irish Pig, with jaw-appendag s. (Copied from H. D. Richardson on Pigs.’ races, but are not strictly inherited, for they occur or fail im animals of the same litter.27 As no wild pigs are known te have analogous appendages, we have at present no reason to suppose that their appearance is due to reversion; and if this be so, we aie forced to admit that a somewhat com- plex, 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), mImay we not venture to surmise that the reduction of the tusks 25 * Eudes- Deslongchamps, ‘Mé- ‘Pigs, their Origin, &.,’ 1847, p. 30 moires de la Soc. Linn.de Normandie, | Nathusius, ‘Die Racen des Schweines,’ vol. vii., 1842, p. 41. Richardson, 1863, s. 54. 80 DOMESTIC PIGS. Cuap. LIL 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 lable to much variation. It is a well-known fact that the young of wild European and Indian pigs,”° 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 ;” 7" whether these latter pigs belong to the same curly-haired race as the Turkish swine, Ido 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.”® 26 D. Johnson’s ‘ Sketches of Indian Field Sports,’ p. 272. Mr. Crawfurd informs me that the same fact holds good with the wild pigs of the Malay peninsula. 27 For Turkish pigs, see Desmarest, “Mammalogie,’ 1820, p. 391. For those of Westphalia, see Richardson’s ‘Pigs, their Origin, &c.,’ 1847, p. 41. 28 With respect to the several fore- going and following statements on feral pigs, see Roulin, in ‘Mém. preé- sentés par divers Savans a ]’Acad.,’ &c., Paris, tom. vi. 1835, p. 326. It should be observed that his account does not apply to truly feral! pigs; but to pigs loug introduced into the country and living in a_half-wild state. For the truly feral pigs of Jamaica, see Gosse’s ‘Sojourn in Jamaica,’ 1851, p. 586; and Ccl Hamilton Smith, in ‘Nat. Library, vol. ix. p. 95. With respect to Africa see Livingstone’s ‘ Expedition to the Zambesi,’ 1865, p. 153. The most precise statement with respect to the tusks of the West Indian feral boars is by P. Labat (quoted by Roulin); out 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, ‘ Sdugethiere,’ s. 331) have not reverted to the wild type. De Blainville (‘Ostéographie,’ p. 132) refers to two skulls of domestic pigs Cuap. ITI. THEIR CHARACTER WHEN FERAL. §1 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.” Jn 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.2? With respect to colour, feral pigs generally revert to that of the wild boar; but in certain parts of 8. 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, ard this colour has likewise occasionally been observed in eat from Patagonia by Al. d’Orbigny, and he states that they have the accipital elevation of the wild European boar, but that the head altogether is “plus courte et plus ramassée.” He refers, also, to the skin of a feral] pig from North America, and says, “il ressemble tout a fait 4 un petit sang- 5 lier, mais il est presque tout noir, et peut-€tre un peu plus ramassé dans ses formes.” 29 Gosse’s ‘ Jamaica,’ p. 386, with a quotation from Williamson’s ‘ Oriental Field Sports.” Also Col. Hamilton Smith, in ‘ Naturalist Library,’ vol ix. p. 94. 0424 3 a &2 CATTLE. Cuap. IIL. 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 kecping 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.*® In breeds thus formed by complex crosses, the most careful and unre- mitting selection during many generations has heen found to be indispensable. Chiefly in consequence of so much crossing, some well-known breeds have undergone rapid changes ; thus, according to Nathusius,*! 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 inhabit- ing 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 39 S. Sidney’s edition of ‘ Youatt on 31 * Schweineschiidel,” s 140. the Pig,’ 1860, pp. 7, 26, 27, 29, 30. “AG Cuap. IIT, THEIR PARENTAGE. So degree, according to Riitimeyer,*? than do the fossil and prehistoric European species, namely, Bos primigenius and longifrons, from each other. They differ, also, as Mr. Blyth,** who has particularly attended to this subject, remarks, in general configuration, in the shape of their ears, in the point where the dewlap commences, in the typical curvature of their horns, in their manner of carrying their heads when at rest, in their ordinary variations of colour, 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 Robilcund, and can maintain themselves in a region infested by tigers. They have given rise to many races differing greatly in size, in the presence of one or two humps, in length of horns, and other respects. Mr. Blyth sums up emphatically that the humped and hump- less cattle must be considered as distinct species. When we consider the number of points in external structure and habits, independently of 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 32 ¢Die Fauna der Pfahlbauten,’ 1861, s. 109, 149, 222. Sce also Geoffroy Saint-Hilaire, in ‘Mém. du Mus. d’Hist. Nat.,’? tom. x. p. 172; and his son Isidore, in ‘ Hist. Nat. Gen.,’ tom. iii. p. 69. Vasey, in his “Delineations of the Ox Tribe,’ 1851, p. 127, says the zebu has four, and common ox five, sacral vertebre. Mr. Hodgson found the ribs either thirteen or fourteen in number; see a note in ‘Indian Field,’ 1858, p. 62. 33 “The Indian Field,’ 1858, p. 74, where Mr. Blyth gives his authorities with respect to the feral humped cattle. Pickering, also, in his ‘ Races of Man,’ 1850, p. 274, notices the peculiar grunt-like character of the voice of the humped cattie. Curr. IIL 84 CATTLE. possess their own sub-breeds;** and these again differ from the cattle of the othér British islands, such as Anglesea, and the western isles of Scotland. Desmarest, who paid attention fo 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: *° the Podolian cattle also are remarkable from the height of their fore-quarters. In the most recent work on Cattle,*° 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 Jong-civilized countries, for we shall presently see that several kinds are kept by the savages of Southern Africa. With respect to the parentage of the several European breeds, we already know much from Nilsson’s Memoir,’ and more especially from Riitimeyer’s works and those of Boyd Dawkins. ‘wo 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 Riitimeyer, we have :— Bos primigenius.— This magnificent, well known species was domesticated in Switzerland during the Neolithic period; even at this early period it varied a little, having apparently been crossed with other races. Some of the larger races on the Continent, as the Friesland, &c., and the Pembroke race in England, closely resemble in essential structure B. primigenivs, and no doubt are its descen- dants. This is likewise the opinion of Nilsson. Bos primigentus existed as a wild animal in Cesar’s time, and is now semi-wild, though much degenerated in size, in the park of Chillmgham; for I am informed by Professor Riitimeyer, to whom Lord Tankerville sent a skull, that the Chillingham cattle are less altered from the true primigenius type than any other known breed.* 34 Mr. H. E. Marquand, in ‘The Times,’ June 25rd, 1856. sance Gen. du Beuf,’ Paris, 1860. Fig. 82 is that of the Podolian breed. 35 Vasey, ‘ Delineations of the Ox- Tribe, p. 124. Brace’s ‘ Hungary,’ 1851, p. 94. The Hungarian cattle descend, according to Rutimeyer (‘ Zahmen Europ. Kindes,’ 1866, s. 13 from Bos primigznius. 36 Moll and Gayot, ‘La Connais- s7 A translation appeared in three parts in the ‘ Annals and Mag. of Nat. Hist.,’? 2nd series, vol. iv., 1849. 38 See, also, Riitimeyer’s ‘ Beitrage pal. Gesch. der Wiederkauer Basel, 1865, s. 54. Cesp. II, THEIR PARENTAGE. 85 Bos trochoceros.—This form is not included in the three species above mentioned, for it is now considered by Riitimeyer to be the female of an early domesticated form of B. primigenius, and as the progenitor of his frontosus race. Imay add that specific names have been given to four other fossil oxen, now believed to be identical with B. primigenius.** 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 *° it was introduced as a domesticated animal into Britain at a very early period, and supplied food to the Roman legionaries.*t Some remains have been found in Ireland in certain erannoges, of which the dates are believed to be from 843-9833 a.p.# It was also the commonest form in a domesticated condition in Switzerland during the earliest part of the Neolithic period. Pro- fessor Owen * thinks it probable that the Welsh and Highland cattle are descended from this form; as likewise is the case, according to Riitimeyer, 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,* and both have been found in the Irish crannoges.*? 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*® 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 39 Pictet’s ‘ Paléontologie,’ tom i. p. 365 (2nd edit.). With respect to B. trochoceros, see Riitimeyer’s ‘Zahmen Europ. Rindes,’ 1866, s. 26. #0 W. Boyd Dawkins on the British Fossil Oxen,’ ‘Journal of the Geolog. Soc.,’ Aug. 1867, p.182. Also ‘ Proc: Phil. Soc. of Manchester,’ Nov. 14, 1871, and ‘Cave Hunting,’ 1875, p. 27, 138. 41 “British Pleistocene Mammalia,’ by W. B. Dawkins and W. A. Sandford, 1866, p. xv. #2 W. R. Wilde, ‘An Essay on the Animal Remains, &c. Royal Irish Academy,’ 1860, p. 29. Also ‘ Proc. of R. Irish Academy,’ 1858, p. 48. #3 “Lecture: Royal Institution of G. Britain,’ May 2nd, 1856, p. 4. ‘British Fossil Mammals,’ p. 513. ** Nilsson, in ‘Annals and Mag. of Nat. Hist.,’ 1849, vol. iv. p. 354. * See W. R. Wilde, ut supra; and Mr. Blythe, in ‘ Proc. Irish Academy,’ March 5th, 1864, 46 Laing’s ‘Tour in Norway,’ p 110. _- 86 CATTLE. Cuap. IL 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 *” (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.*® 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 magni- ficent 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.” #9 Bos primigenus and longifrons have been ranked by nearly all paleontologists 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 steri- lity 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 47 Tsid. Geoffroy Saint-Hilaire, *© Cuar. III. THEIR VARIATION. 91 who has given a table of the average period of their denti- ‘ion, 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.” °° With respect to the period of conception, it seems certain that Alderney and Zetland cows often become pregnant earlier than other breeds.°? Lastly, as four fully developed mamme is a generic character in the genus Bos,°° it is worth notice that with our domestic cows the two rudi- mentary mammee often become fairly well developed and yield milk. As numerous breeds are generally found only in long- civilized countries, 1t 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 commu- nication, several distinct breeds of cattle now exist or for- merly existed. At the Cape of Good Hope Leguat observed, in the year 1720, three kinds.®! At the present day various travellers have noticed the differences in the breeds in Southern Africa. Sir Andrew Smith several years ago remarked to me that the cattle possessed by the different tribes of Caffres, though living near each other under the same latitude and in the same kind of country, yet differed, and he expressed much surprise at the fact. Mr. Andersson has described ®* 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 58 ¢Ann. Agricult. France,’ April, des Cours Scientifiques,’ Feb. 12, 1688, 1857, as quoted in ‘The Veterinary,’ p- 657), that the cattle of Piacentino vol. xii. p. 725. I quote Tessier’s obser- vations from Youatt on Cattle, p. 527. 59 The Veterinary,’ vol. viii. p. 681, and vol. x. p. 268. Low’s ‘Domest. Animals, &c.,’ p. 297. 60 Mr. Ogleby, in ‘Proc. Zoolog. Soc.,’? 1836, p. 138, and 1840, p. 4. Quatrefages quotes Philippi (‘ Revue have thirteen dorsal vertebre and ribs in the place of the ordinary number of twelve, $1 Leguat’s Voyage, quoted by Vasey in his ‘ Delineations of the Ox- tribe,’ p. 132. ° «Travels in South Africa,’ pp. 317, 336. 92 CATTLE. Cuap. ITI. the case with the cattle of Benguela. The Namaqua cattle in size and shape nearly resemble European catile, 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 extra- ordinarily large. The Bechuana cattle have even larger horns, and there is now a skull in London with the two horns 8 ft. 8} in. long, as measured in a straight line from tip to tip, and no Jess than 13 ft. 5 in. as measured along their curva- ture! 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 ® 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 hke European cattle, the other different, with remarkable horns. In Para- guay, Azara describes a breed which certainly originated in §. 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 smali 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 63 ¢ Mém. de |’Institut présent. par June 15,1846. See Azara, ‘Quadru- divers Savans,’ tom. vi., 1835, p.333. | pédes du Paraguay, tom. ii. pp. 359 For Brazil, see ‘Comptes Rendus, 361. Cuap. III. THEIR VARIATION. 93 other dogs, or as improved pigs, according to H. von Nathusius, do to common pigs.®* Riitimeyer believes that these cattle belong to the primigenius type.® The forehead is very short and broad, with the nasal end of the skull, together with the whole plane of the upper molar-teeth, curved upwards. The lower jaw projects beyond the upper, and has a corre- sponding upward curvature. It is an interesting fact that an almost similar confirmation characterizes, as lam 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 isusual. 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: °° “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 con- nexion of some of the bones is changed. Other differences might be added: thus the plane of the condyles is somewhat 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. 65 Ueber Art des zahmen Europ. Rindes, 1866, s. 28. 66 * Descriptive Cat. of Ost. Collect. of College of Surgeons,’ 1853, p. 624. Vasey, in his ‘ Delineations of the Ox- tribe,’ has given a figure of this skull; and I sent a photograph of it to Prof Rutimeyer. 94 CATTLE. Cuap. IIL 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 ecllected 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 ori- ginated 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 civilized nature in being fiercer than common cattle, and in the cow, if visited too often, easily deserting her first calf. The breed is very true, anda 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 oftspring 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 acommon 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 lps 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 ave able to judge from the ordinary habits of an animal, on what circumstances, occurring only at long intervals of time, its rarity or extinc- tion 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. Curap. IDL CAUSES OF VARIATION. 95 Having described the semi-monstrous niata breed, I may allude to a white bull, said to have been brought from Africa, which was exhibited in London in 1829, and which has been well figured by Mr. Harvey.®’ It had a hump, and was fur- nished 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 trans- versely 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 ques- tion 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.°* That climate directly affects the thickness of the skin and the hair is likewise certain: thus Roulin asserts °° that the hides of the feral cattle on the hot Llanos “ are always much less heavy than those of the cattle raised on the high plat- form of Bogota; and that these hides yield in weight and in thickness of hair to those of the cattle which have run wild on the lofty Paramos,’ ‘The same difference has been observed 67 Loudons Magazine of Nat. 68 Low, ‘Domesticated Animals of Hist.,’ vol. i., 1829, p.113. Separate the British Isles,’ p. 264. figures are given of the animal, its 69 “Mém. de l'Institut présent. par hoofs, eye, and dewlap. divers Savans,’ tom. vi., 1835, p. 332 36 CATTLE. *. Grap- Fee in the hides of the cattle reared on the bleak Falkland Islands and on the temperate Pampas. Low has remarked “° 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 tha 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. Itis probable that some breeds, such as the semi- monstrous niata cattle, and some peculiarities, such as being hornless, &c., 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 characterized must be at least partially separated from others. This degree of care, however, has sometimes been taken even in little-civilized districts, where we should least have expected it, as in the case of the niata, chivo, and horn- less cattle in 8. America. That methodical selection has done wonders within a recent period in modifying our cattle, no one doubts. During the process of methodical selection it has occasionally happened that deviations of structure, more strongly pronounced than mere individual differences, yet by no means deserving to be called monstrosities, have been taken advantage of: thus the famous Long-horn Bull, Shakespeare, though of the pure Canley stock, “scarcely inherited a single point of the long-horned breed, his horns excepted ;71 yet in the hands of Mr. Fowler, this bull greatly improved his race. We have also reason to believe that selection, carried on so far unconsciously that there was at no one time any distinct intention to improve or change the breed, has in the course of time modified most of-our cattle ; for by this process, aided by more abundant food, all the lowland British breeds have increased greatly in size and in early maturity since the reign of Henry VII. It should never be forgotten that many animals have to be annually 70 Idem, pp. 304, 368, &e. 72 Youatt on Cattle, p. 116. Lord 71 Youatt on Cattle,p.195. Afull Srencer has written on this same account of this bull is taken from subject Marshall. . Cap. Til. SHEEP: THEIR VARIATION. 97 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 Brita, I conclude that, although slight differences in the nature of the climate, food, &c., as well as changed habits of life, aided by correlation of growth, and the occasional appearance from unknown causes of con- siderable 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,’* but that our domestic sheep form a distinct genus, now completely extinct. A German 73 Blyth, on the genus Ovis, in Mr. Blyth’s excellent articles in ‘ Land “Annals and Mag. of Nat. History,’ and Water,’ 1867, pp. 134, 156. vol, vii., 1841, p. 261. With respect Gervais, ‘Hist. Nat. des Mammitéres, fo the parentage of the breeds, see 1855, tom. ii. p. 191. 98 SHEEP: Cuap. IIL naturalist’* believes that our sheep descend from ten aborigi- nally distinct species, of which only one is still hving in a wild state! Another ingenious observer,’® though nota naturalist, with a bold defiance of everything known on geographical dis- tribution, 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. Riitimeyer 7° found in the Swiss lake-dwellings the remains of a small breed, with thin tal! 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 vertebre, 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 ereat masses of fat on the rump, with the tail in a rudimen- tary condition. ‘The Angola variety of the long-tailed race has curious masses of fat on the back of the head and beneath the jaws.’7 Mr. Hodgson in an admirable paper ‘® on the sheep of the Himalaya infers from the distribution of the several races, “ that this caudal augmentation in most of its phases 1s 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. 7” Dr. L. Fitzinger, ‘Ueber die vol. ii. p. 264. Racen des Zahmen Schafes,’ 1860, s. 76 ¢Pfahlbauten,’ s. 127, 193. 86. 77 Youatt on Sheep, p. 120. 75 J, Anderson, ‘Recreations in 78 ¢ Journal of the Asiatic Soe. of Agricuiture and Natural History,’ Bengal,’ vol. xvi. pp. 1007, 1016. Cnap. Il THEIR VARIATION. 99 It is remarkable that multiplicity of horns “is generally accompanied by great length and coarseness of the fleece.” ‘9 This correlation, however, is far from being general; for instance, Iam 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 mammez 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.” *° 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 like- wise been considered as a generic distinction in sheep; but Isidore Geoffroy *! 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.” *2 Mr. Hodgson states that the ex- traordinarily 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.*? 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 79 Youatt on Sheep, pp. 142-169.’ = 435. 89 ¢ Journal Asiat. Soc. of Bengal, 82 Youatt on Sheep, p. 138. tol. xvi., 1847, p. 1015. 83 ‘Journal Asiat. Soc. of Bengal,’ $1 «Hist. Nat. Gen. tom. iii. p. vol. xvi., 1847, pp. 1015, 1016. LOO SHEEP: Cuap IIL than in the female; and Fitzinger ** 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.” °° Marshall relates °° 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.” §7 There is very good evidence that Englsh breeds of sheep will not succeed in France.*? Even in certain parts of England it has been found im- 84 $ a2 s *1> = eae by Godron, ‘De l’Espéce,’ p. 364),a qu’on éléve en domesticité. Cuap. 1YV. FERAL RABBITS. 119 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 Zoo- logical 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 de- scended from a domesticated breed. I was so much surprised at it, that Irequested Mr. Haywood to make inquiries on the spot, whether they were much hunted by the inhabitants, or per- secuted 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 w.th them. 120 DOMESTIC RABBITS: Cuar. IV. 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 cha- racter, 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 appled 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 propor- sional length of the limbs in comparison with that of the body. As a standard of comparison, 1 prepared skeletons of two wild rabbits from Kent, one from the Shetland Islands, and one from Cuar. LV. DIFFERENCES IN THEIR SKELETONS. 12) 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. Ina wild rabbit the length was 3°15 inches, in a large fancy rabbit 43; 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 elon- gated. ‘The lop-eared rabbits which I examined were, though not fat, more than twice as heavy as the wild specimens; but the skull was very far from being twice as long. Even if we take the fairer 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 aniversal 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 explana- tion seems to lie in the circumstance that during a number of gene- rations the artificial races have been closely confined, and have had little occasion to exert either their senses, or intellect, or voluntary muscles; consequently the brain, as we shall presently more fully see, has not increased relatively with the size of body. As the brain has not increased, the bony case enclosing it has not increased, and this has evidently affected through correlation the breadth of the entire skull from end to end. In all the skulls of the large lop-eared rabbits, the supra-orbital plates or processes of the frontal bones are much broader than in the wild rabbit, and they generally project more upwards. In the zygomatic arch the posterior or projecting point of the malar-bone 122 DOMESTIC RABBITS: Cuap. IV. is broader and blunter; and in the specimen, fig. 8, it is so in a remarkable degree. This point approaches nearer to the auditory meatus than in the wild rabbit, as may be best seen in fig. 8; but this circumstance mainly depends on the changed direction of the TN ray Ky Sa Te ms we a 5 SS SNe =“ ete ae rs = — ; ME FS eee 54 ae " gz Ke = = Son — See ay Se Fig. 6.—Skull of Wild Rabbit, of natural size. Fig. 7.—Skull of large Lop-eared Rabbit, of natura. s1Ze. meatus. The inter-parietal bone (see fig. 9) differs much in shape in the several skulls; generally it is more oval, that is more ex- tended in the line of the longitudinal axis of the skull, than in the wild rabbit. The posterior margin of “the square raised plat- Cuap. IV. DIFFERENCES IN THEIR SKELETONS. form’ of the occiput, instead of being truncated, or projecting slightly as in the wild rabbit, is in most Jop-eared rabbits pointed, as in fie.9,C. The paramastoids rela- tively to the size of the skull are generally much thicker than in the wild rabbit. The occipital foramen (fig. 10) presents some remarkable differ- ences: in the wild rabbit, the lower edge between the condyles is considerably and almost angu- larly 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 ex- ceeds 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 Fig 8.— Part of Zygomatic Arch, showing the projecting end of the malar bene of the auditory meatus: of natural size. Upper figure, Wild Rabbit. Jower figure, Lop- eared, hare-coloured Rabbit. square notch, in three there was a trace of the notch, and in two alone it was well developed. These differences in the shape of the foramen are remarkable, considering that it gives passage to so Important a structure as the spinal marrow, though apparently the outline of the latter is not affected by the shape of the passage. In all the skulls of the large lop-eared rabbits, the A B Cc ath i 0 LT} 7 Fig. 9.—Posterior end of skull, of natural size, showing the inter-parietal bone. A. Wild Rabbit. Rabbit from island of P, Santo, near Madeira. C. Large Lop-eared Rabbit. B. Feral bony auditory meatus is conspicuously larger than in the wild rabbit. Ina skull 4°8 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 Wey Fig. 10.—Occipital Foramen, of natural size, in— A. Wild Rabbit; B. Large Lop-eared Rabbit. its margin on the side nearest the skull stands up higher than 5 Waterhouse, ‘Nat. Hist. Mammalia,’ vol. ii. p. 36. 124 DOMESTIC RABBITS: Cuap. IV. 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 con- tinued selection of indi- viduals having larger and larger ears. The influence iy of the external ear on the 3 bony meatus is well shown ‘i “\ in the skulls (I have ex- mn ye i apt 1 pie 5 5 \ Ee” LS a Ue -// “4A * 3 =: Vi; Mg | 5 j Wh ‘ | \ vA ka My ez, 1h 43 ‘ ly if i fi, amined three) of half-lops (see fig. 5), im 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 struc- ture of the whole skull. I here give a drawing (fig. 11) of the skull of a halt-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 Fi = Sknll, of natural size, of Half-lop Rabbit night one; both hens Sorcae the ‘different Tteiion of the aie and anterior Margins of the meatus on the two sides, and the consequent left zygomatic arch on the general distortion we the skull. ate a side of the lopping ear stand sbaveuimel Cor might side of ‘fgure). lopped. little- in advaaeenaenee 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 tyar. IV. DIFFERENCES IN THEIR SKELETONS. £25 an animal during many generations under confinement, and so leading tc 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 smal! 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 inter- mediate in general proportions, and in most other characters, between those of the largest lop-eared and wild rabbits. It presents only one singular character: though considerably longer than the skull of the wild rabbit, the breadth measured within the posterior supra- orbital fissures is nearly a third less than in the wild. The skulls of the séver-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. % ae * 126 DOMESTIC RABBITS: Cuap. IV- Consequently J examined skulls of the hare, but no light could thus be thrown onthe 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 pomts in which domestic rabbits vary,—namely, in general proportions, in the form’ and size of the subra-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. Vertebre.—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 vertebree. This is remarkable, as Gervais gives seven as the number for the whole genus Lepus. The caudal vertebre apparently differ by two or three, but I did not attend to them, and they are difficult to count with certainty. In the first cervical vertebra, or atlas, the anterior margin of the neural arch varies a little in wild specimens, being either nearly smooth, or furnished with a small supra-median atlantoid process; I have figured a specimen with the - largest process (a) which I have seen ; but it will be observed how inferior this is in size and different in shape to that in a large lop-eared rabbit. In the latter, the infra-median pro- cess (b) is also proportionally much ~ thicker and longer. The ale are a little squarer in outline. Third cervical vertebra.—In the wild rabbit (fig. 13, a a) this ver- tebra, 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 Fig. 12.—Atlas Vertebre, ofnaturalsize; large lop-eared rabbits this process Upper figure, Wild Rabbit. lower (B@ ) is forked in the third vertebra, See enoUrd, Loree, Bhi as in the fourth of the wild rabbit. Seats ESE Lae eae nae =. But the third cervical vertebre of the wild and Jop-eared (A J, B Db) rabbits differ more conspicnously when their anterior articular Cuap. IV. DIFFERENCES IN THEIR SKELETONS. 1 A7¢ surfaces are compared; for the extremities of the antero-dorsal pro- cesses 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 (8 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 atten- tion. First dorsal vertebra.— Its neural spine varies in length in the wild rabbit 5 EF g- 13.—Third Cervical Vertebre, of natural size, Pee ry ee 2 ae laut eee short, but generally more anterior articular surfaces. Scat than half as long as that of the second dorsal; but I have seen it in two large lop-eared rabbits three-fourths of the length of that of the second dorsal vertebra. Ninth and tenth dorsal vertebre.—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 vertebree. In the large lop- eared rabbits the neural spines of the tenth, ninth, and eighth vertebre, 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. Sothat 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 vertebree in some species of hares. In the Angora, Chinchilla, and Hima- layan rabbits, the neural spines of the eighth and ninth vertebrze 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 cha- racters 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 vertebre were not at all larger than those of the several anterior vertebree. In this same Porto Santo specimen there was no trace in the ninth vertebra of the anterior lateral processes (see woodcut 14), which are plainly deve- loped in all British wild rabbits, and still more plainly developed in the large lop-eared rabbits. In a half-wild rabbit from Sandon Park,” a hemal spine was moderately well developed on the under \) (iP hy *\ HS it = 2 f vi AF -4 Ala NN § 76 These rabbits have run wild for and in cther places in Staffordshire a considerable time in Sandon Park, and Shropshire. They originated, as 128 DOMESTIC RABBITS: Cuap. IV, side of the twelfth dorsal vertebra, and I have seen this in no other specimen. Fig. 14.—Dorsal Vertebra, from sixth to tenth inclusive, of natural size, viewed laterally. A. Wild Rabbit. B. Large, Hare-culoured, so cailed Spanish Rabbit. Lumbar Vertebroe.—I have stated that in two cases there were Fig. 15.—Terminal bone of Sternum, of natural size. A. Wild Rabbit. B. Hare- coloured, Lop-eared Rabbit. C. Hare-coloured. Spanish Rabbit. (N.B. The left- hand angle of the upper articular extremity of B was broken, and has been accidentally thus _ repre- sented.) eight instead of seven lumbar vertebre. The third lumbar vertebre in one skeleton of a wild British rabbit, and in one of the Porto Santo feral rabbits, had a heemal spine; whilst in four skeletons of large lop-eared rabbits, and in the Himalayan rabbit, this same vertebra had a well developed hzmal spine. felvis—In four wild specimens this bone was almost absolutely identical in shape; but in several domesticated breeds shades of differences could be distinguished. In the large lop-eared rabbits, the whole upper part of the ilium is straighter, or less splayed out- wards, than in the wild rabbit; and the tuberosity on the inner lip of the anterior and upper part of the ilium is proportionally more prominent. Sternum.—The posterior end of the pos- terior sternal bone in the wild rabbit (fig. 15, A) is thin and slightly enlarged; in some of the large lop-eared rabbits (8) it is much more enlarged towards I have been informed by the game- being white with a streak along the keeper, from variously-coloured do- spine, and with the ears and certain mestic rabbits which had been turned marks about the head of a blackish- out. They vary in colour; but grey tint. They have rather longer many are symmetrically coloured, bodies than common rabbits. Cuar.1V. DIFFERENCES IN THEIR SKELETONS, 129 the extremity; whilst in other specimens (©) it keeps nearly of the same breadth from end to end, but is much thicker at the extremity. Scapula.—-The acromion sends out a rectangular bar, ending inan oblique knob, which latter in the wild rabbit (tig. 16, A) varies a little in shape and size, as does the apex of the acromion in sharpuess, 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 (8) the oblique terminal knob is de- veloped 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 Cc D figs. BC; and D. Fig. 16.—Acromion of Scapula, of natural size. Limbs.—In these I could as Rabbit. B, C, D, Large, Lop-eared detect no variation; but the bones of the feet were too troublesome to compare with much care. I have now described ail 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 import- ance become modified under domestication. Noone will say, for instance, that the oc ipital foramen, or the atlas, or the third cervical vertebra is a partof slightimportance. Ifthe several vertebre of the wild and lop-eared rabbits, of which figures have been given, had been found fossil, paleeontologists 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 ae and hind legs compared with each other, have 130 DOMESTIC RABBITS: Cuar. IV. remained nearly the same as in the wild rabbit; but in weight, the bones of the hind legs apparently have not increased in due pro- portion 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 ereat 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; con- sequently 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 mecreased 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 te 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 mere interesting point, to which I was led to attend by finding, as previously stated, that with all domesticated rabbits the length of the skull relatively to its breadth has greatly increased in comparison with that of the wild rabbits. If we had possessed a large number of domesticated rabbits of nearly the same size with the wild rabbits, it would have been a simple task to have measured and compared the capacities of their skulls. But this is not the case: almost all the domestic breeds have larger bodies than wild rabbits, and the lop-eared kinds are more than double their weight. As a small animal has to exert its senses, intellect, and instincts equally with a large animal, we ought not by any means to expect an animal twice or thrice as large as another to have a brain of double or treble the size.27 Now, after weighing the bodies of four wild rabbits, and of four large but not fattened lop-eared rabbits, I find that on an average the wild are to the lop-eared in weight as 1 to 2°17; in average length of body as 1 to 1-41; whilst in capacity of skull they are as 1 to 1:15. Hence we see that the capacity of the skull, and consequently the size of the brain, has increased but little, relatively to the increased size of the body; and this fact explains the narrowness of the skull relatively to its length in all domestic rabbits. 27 See Prof. Owen’sremarks on this 1852: with respect to Birds, see subject in his paper on the ‘Zoological ‘ Proc. Zoolog. Soc.,’ Jan. 11th, 1848, Significance of the Brain, &c.,of Man, pp. 8. &c.,’ read before Brit. Association, Cuap. IV EFFECTS OF USE AND DISUSE. 131 In the upper half of the following table I have given the measure- ments of the skull of ten wild rabbits; and in the lower half, of eleven thoroughly domesticated kinds. As these rabbits differ so ereatly 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 auy other part affords a perfect standard. In the first column of figures the extreme length of the skull is given in inches and decimals. J am aware that these measurements pretend to greater accuracy than is possible; but I have found it the least trouble to record the exact leneth 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'[6 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 vil), 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 wildrabbit. Had I taken the Irish rabbit, No. 3, as the standard, the following results would have been somewhat more striking. Turing to the table: the first four wild rabbits have skulls of the same length, and these differ but little in capacity. The Sandon rabbit (No. 4) is interesting, as, though now wild, it is known to be descended from a domesticated breed, as is still shown by its pecu- liar colouring and longer body; nevertheless the skull has recovered its normal length and full capacity. The next three rabbits are wild, 132 DOMESTIC RABBITS: Cuar JV 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 J 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 ease. Looking to the lower half of the Table, which gives the measure- ments of domesticated rabbits, we see that in all the capacity of the skull is less, but in very various degrees, than might have been anticipated according to the length of their skulls, relatively to that of the wild rabbit No.1. In line 22 the average measurements of seven large lop-eared rabbits are given. Now the question arises, has the average capacity of the skull in these seven large rabbits increased as much as might have been expected from their greatly increased size of body. We may endeavour to answer this question in two ways: in the upper half of the Table we have measurements of the skulls of six small wild rabbits (Nos. 5 to 10), and we find that on an average the skulls are ‘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 11386 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. 23 This standard is apparently con- Zoolog. Soc.,’ 1861, p. 86) gives 210 siderably too low, for Dr.Crisp(‘Proc. grains as the actual weight of the 135 EFFECTS OF USE AND DISUSE. ry i . Cuar. IV 1, 2. 3. 4, o. 6. ie 8. 9. 10. Ge 12. 15. 14. 15. 16. 17. 1x, 19. 20. fails 22, 23. 24, Name of Breed. WILD AND SEMI-WILD RABBITS, Wild Rabbit, Kent .. 5 ShetlandIslands .. .. .. - tclamdae tease nc: se us Domestic rabbit, run wild, Sandon che Wild, common variety, small specimen, Kent Wild, fawn-coloured variety, Scotland . Silver-grey, small specimen, ‘Thetford warren Feral rabbit, Porto Santo : 99 99 ee . oe ee eo oe oo 99 Average of the three Porto Santo rabbits 99 ee ° Domestic Rasits. UMA AV AM cnet e sada iceny ute Cae ion DIOS CO Wariner int healer man timer enn Ss See PAOLA ve oes, oo aety wremntoaT? Fas ie shoe Pape koe (GHouliprol WDE ae rectea. on], Wace Scene re Large lop-eared PTO Rp Sirasatal ie, Nees Large hare-coloured.. .. «2 «) - Aver Tage gi above seven large lop-eared rabbits Hare i vinudus) Wnglish specimen .. .. oe ee German specimen .... Length of Skull. . Hr mer Cz Or bo a Cr . =~ — =I Ree OD OO SE — ° = 455 ~~ Jt II. Length of Body from Incisors to Anus. Ilr Weight of whole Body. inches. Wee: 18°5 17°0 15°5 IV. Capacity of Skull mea- sured by Small Shot. grains. 972 979 992 977 879 918 938 893 756 835 828 968 803 697 995 1065 1158 1037 1208 1232 112+ 1ISl 1136 J315 1455 V. Cupacity cal ulated according to Length of Skull rela- tively to that ot No. 1. grains. 913 950 910 873 879 910 888 1080 1002 1080 1126 1265 1265 1255 1265 1826 1311 119 1268 VI. VII. : Showing vow much per cent Difference ‘ : iatareen 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 actual and calculated ape ities of Skulls. grains. [2 per cent. too heavy in comparison with No. 1.] \ 38 4 per cent. too light. 32 ee 1) 28 3. ,, £«2+too heavy 20 2 ”) 9 123 16 ,, &too light, 79 Deca ” 60 7 9? 9? 117 {ees Fe * 199 rh ea 0 38: ara s rs 13 AB aaah oe 200 1S ivessis as 112 ear, : 218 Dllisen ies an o7 fos y O4 RES 30 187 1 Syaen ae ie 6 ° oe) ”? 182 VOL Bere ‘it 154 IT 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, 15 of Table) are only a little larger in body and have skulls only a little Ienger, 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 eould 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, oiters the most remark- able 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 Angera 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 ali these standards the brain appears to be much too small, though ina less degree when the standard of the limb-bones was used ; and this latter cireum- stance 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 remark- able amount of reduction. DOMESTIC RABBITS: Cuap. IV. 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 brain of a hare which weighed 7 lbs., and 125 grains as the weight of the brain of a rabbit which weighed 3 lbs. 5 oz., that is, the same weight as the rabbit No. 1 in my list. Now the contents of the skull of rabbit No. 1 in shot is in my 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, Cuap. IV. EFFECTS OF USE AND DISUSE, 1 Bas 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, not- withstanding 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 faras 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 hmbs 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 as- sumed characters proper to the fourth cervical vertebra; and the eighth and ninth dorsal vertebre have similarly assumed cha- racters proper to the tenth and posterior vertebre. The skull in the larger breeds has increased in length, but not in due pro- portion 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 ccrrelation has affected the bones of the face 136 DOMESTIC RABBITS. Cuap. IV. 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 selec- tion; 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 shght degree of almost every bone in the upper part of the skull, and even the position of the condyles of the lower jaw. Cuar. V. PIGEONS : DESCRIPTION OF BREEDS. aw CHAPTER V. DOMESTIC PIGEONS. ENUMERATION AND DESCRIPTION OF THE SEVERAL BREEDS—INDIVIDUAL VARIABILITY—VARIATIONS OF A REMARKABLE NATURE—OSTEOLOGICAL CHARACTERS: SKULL, LOWER JAW, NUMBER OF VERTEBRA:—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 evi- dence 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. LT have kept alive all the most distinct breeds, which I could procure in England or from the Continent; and have pre- pared skeletons of all. I have received skins from Persia, and a large number from India and. other quarters of the 138 DOMESTIC PIGEONS: Cuar. V. world.! Since my almission into two of the London pigeon- clubs, I have received the kindest assistance from many of the most eminent amateurs.’ The races of the Pigeon which can be distinguished, and which breed true, are very numerous. MM. Boitard and Corbié * 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 ereater 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 1 The Hon. C. Murray has sent me and gave me specimens. I had access 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 infor- mation 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. 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 co_-ection of Pouters, 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 speci- mens 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 1 beg permission to return my sincere and cordial thanks. 3 ‘Les Pigeons de Voliére et de Colombier,’ Paris, 1824. During forty- five years the sole occupation of M. Corbié 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 ecolombi Domestici. Padova, 1873. Cuap. V. DESCRIPTION OF BREEDS. 139 collection of the Columbide in the British Museum, and, with the exception of a few forms (such as the Didunculus, Calznas, Goura, &c.), Ido 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 * 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 ex- panded upraised tail like that of the fantail; or for an ceso- phagus like that of the pouter. Ido not fora moment pretend that the domestic races differ from each other in their whole organisation as much as the more distinct natural genera. 1] 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 follow- ing 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-charac- terized 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 * “Coup d’Oeil sur JVOrdre des Paris, 1855. This author makes 288 Pigeons,’ par Prince C. L. Bonaparte, species, runked under 85 genera. 140 DOMESTIC PIGEONS: Cuap. V- 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 “aber- rant” 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 ofequalvalue. 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 higha 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.?® I will now give a brief description of all the principal breeds. The diagram on the following page may aid the reader in learning their names and seeing their affinities. The rock-pigeon, or Columba livia ((ncluding under this name 5 As Iso often refer to the sizeof tween the measurements of two wild the C. livia, or rock-pigeon, it may birds, kindly sent me by Dr. Edmond- be convenient to give the mean be- stone from the Shetland Islands. Inches Length from feathered base of beak toend oftail.. .. .. .. .. 14°25 ~ to oil-gland. .. 2. <.. “2. eee us ree tip of beak to eat of tail <5.) es eee S 2) ee - of tail-feathers .. =) ope ta Se Vine ewe ©” cteedy 2a eae ies » from tip to tip of wing PM eer e3 of folded wing . me "ONO “MOPQUuen T ° peg & TL h paov sy "10 "LOLLY "LOOT 4 * BS ‘unqoowe unepuyt — ~plousy UnoyisT a LULUG fl qog nung ysrpugy you OC 8 Ss : 88 = § 8 QUT, te ae, as & 2d WoC] 8 3 8 § ES s 2 : eudAc uoSur RaweRAS pep gy uoostg Lg ory, BAL? : : ‘q uvuiey OJUOLT, : a wooLopuvog a ay 05 : gy CR SOS Tew eg en ame a ne mma gee RECO Sew asenesen : tH ; : py “Udcossn gy iS) i : m roy wun Y, Q : ; uLyory = @ BSSVAN 3 o) ' R ' : LoTqQum T, ‘ : Mia es is, : ; UVISIOT ‘IV T-UCM : Pee | i seanoue ‘gan0up ‘IL ‘OT «HAS G 8 v4 ‘9 ¢ "F '€ % “as S| eT SS SS —————EE : nN — “AT dnowo TM dowd TE dQOwo T dlhowo “i OT SR a3 - = et s) a - va ‘NOWUDId-MOOU XO VIATT VaWoOTOO Cap. VY. DESCRIPTION OF BREEDS. 143 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. 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 ieee the most distinct of all domesticated pigeons. Race I.—Pourer Picrons. (Kropftauben, German. Grosses- gorges, or boulans, French.) (sophagus of great size, barely separated from the crop, often inflated. Body and legs elongated. Beak of moderate dimen- sions. Sub-race ].—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 cesophagus. The diameter of the upper part of the cesophagus is immense, even close up to the head. The beak in one bird which I possessed was almost completely buried when the cesophagus 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 144 DOMESTIC PIGEONS: Cuap. Y. often strike the backs of their wings togethcr, 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 Fig. 18.—English Pouter. eenerally broader and the vertebree 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 ielation to the length of body, Guar, V. DESCRIPTION OF BREEDS. 145 this is not the case. The beak likewise appears longer, but it is in fact a little shorter (about -03 of an inch), proportionally with the size of the body, and relatively to the beak of the rock-pigeon. The Pouter, though not bulky, is a large bird; I measured one which was 343 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+ and 143. There are many sub-varieties of the Pouter of different colours, but these I pass over. Sub-race II. Dutch Pouter—This seems to be the parent-form of our improved English Pouters. I kept a pair, but I suspect that they were not pure birds. They are smaller than English pouters, and less well developed in all their characters. Neumeister’ says that the wings are crossed over the tail, and do not reach to its extremity. Sub-race IIT. The Lille Pouter—I know this breed only from description. It approaches in general form the Dutch Pcuter, but the inflated cesophagus 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 erown of the head. The middle toe alone is feathered. A variety ef this sub-race, called the claquant, is described by MM. Boitard and Corbié; it pouts but little, and is characterised by the habit of violently hitting its wings together over its back,—a habit which the English Pouter has in a slight degree. Sub-race IV. Common German Pouter—I know this bird only from the figures and description given by the accurate Neumeister, one of the few writers on pigeons who, as I have found, may always be trusted. This sub-race seems considerably different. The upper part of the esophagus is much less distended. The bird stands less upright. The feet are not feathered, and the legs and beak are shorter. In these respects there is an approach in form to the common rock-pigeon. The tail-feathers are very long, yet the tips of the closed wings extend beyond the end of the tail; and the length of the wings, from tip to tip, and of the body, is greater than in the English Pouter. Group II. This group includes three Races, namely, Carriers, Runts, and Barbs, which are manifestly allied to each other. Indeed, certain carriers and runts pass into each other by such in- sensible gradations that an arbitrary line has to be diawn between them. Carriers also graduate through foreign bieeds into the rock-pigeon. Yet, if well-characterised Carriers and 7 “Das Ganze der Tanbenzucht :’ 8 Boitard and Corhié, ‘ Les Pigeons, Weimar, 1837, pl. 11 and 12. &c., p. 177, pl. 6. re REE ak : 146 DOMESTIC. PIGEONS: Cuap. V. Barbs (see figs. 19 and 20) had existed as wild species, no ornithologist would have placed them in the same genus with each other or with the rock-pigeon. This group may, as a general rule, be recognised by the beak being long, with the skin over the nostrils swollen and often carunculated or wattled, and with that round the eyes bare and likewise carunculated. The mouth is very wide, and the feet are large. Nevertheless the Barb, which must be classed in this same group, has a very short beak, and some runts have very little bare skin round their eyes. Race I1.—Carriers. (Tiirkische Tauben; pigeons tures, dragons. ) Beak elongated, narrow, pointed; eyes surrounded by much naked, generally carunculated, skin ; neck and body elongated. Sub-race I, The English Carrier.—This is a fine bird, of large size, close feathered, generally dark-coleured, 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; there- fore 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 com- parison; 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 313 inches from tip to tip of wing. Birds of this sub-race are too valuable to be flown as carriers. Sub-race II. Dragons; Persian Carriers—The English Dragon differs from the improved English Carrier in being smaller in all its dimensions, and in having less wattle round the eyes and over 147 Sir W. Elliot sent DESCRIPTION OF BREEDS. Cxuap. Y. mandible. ls, and none on the lower me from Madras a Bagdad Carrier (someti Ti the nost mes called khandési), the it would be considered name of which shows its Persian origin: ‘IOPIIVD YSTSuy—'GT “Sly oy ee orr Sy S| aie a See cuts WG. : ‘ moO of =a = <5 : FSS = : V0 °S113M7 acs "OS MAYS 7 HLeloMes -. = or a) i= is aw © (2) da} () . nM Ras aa 42h on rob) cb) ooo et o°0 4q ge fas} E ge: Hi bo Oo Haye) {SI pale 2» £3 ao oO) aod 3S us 2 Ass S ere Gey eb) ee R Eiaica w = 3S e ae o oog & oor Oo. re) SF as 148 DOMESTIC PIGEONS: Cuap. V. wattled, whilst that over the nostrils was fairly wattled. The Hon. C Murray, also, sent me two Carriers direct from Persia; these had nearly the same character as the Madras bird, being about as - large as the rock-pigeon, but the beak in one specimen was as much as 1:15 in length; the skin over the nostrils was only moderately, and that round the eyes scarcely at all wattled. Sub-race III. Bagadotten-Tauben of Neumeister (Pavdotten- or Tocker-Tauben).—I owe to the kindness of Mr. Baily, jun., a dead specimen of this singular breed imported from Germany. It is certainly allied to the Runts; nevertheless, from its close affinity with Carriers, it will be convenient here to describe it. The beak is long, and is hooked or bowed downwards in a highly remarkable manner, as will be seen in the woodcut to be hereafter given when I treat of the skeleton. The eyes are surrounded by a wide space of bright red skin, which, as well as that over the nostrils, is modes rately wattled. The breast-bone is remarkably protuberant, being abruptly bowed outwards. The feet and tarsi are of great length, larger than in first-rate English Carriers. The whole bird is of large size, but in proportion to the size of the body the feathers of the wing and tail are short; a wild rock-pigeon, of considerably less size, had tail-feathers 4°6 inches in length, whereas in the large Bagadotten these feathers were scarcely over 4:1 inches in length. Riedel® remarks that it is a very silent bird. Sub-race IV. Bussorah Carrier—Two specimens were sent me by Sir W. Elliot from Madras, one in spirits and the other skinned. The name shows its Persian origin. It is much valued in India, and is considered as a distinct breed from the Bagdad Carrier, which forms my second sub-race. At first I suspected that these two sub:races might have been recently formed by crosses with other breeds, though the estimation in which they are held renders this improbable; but in a Persian treatise,° 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 cme 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 ® ‘Die Taubenzucht,’ Ulm, 1824,s. in 1770: I owe to the great kindness 42. of Sir W. Elliot a translation of this 1@ This treatise was written by curious tseatise. Sayzid Mohammed Musari, who died Cuar. Y. DESCRIPTION OF BREEDS. 149 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 Kalé 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 IIJ.—Ronts. (Scanderoons: die Florentiner Tauben and Hinkeltauben of Neumeister; pigeon bagadais, pigeon romain. ) Beak long, massive; body of great size. Tnextricable 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-ruce 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 [ have felt myself compelled to place the Bagadotten in Race II., or that of the Carriers, and the present bird in Race ITJ., or that of 150 DOMESTIC PIGEONS: Cup. V. the Rants. The Scanderoon has a very short, narrow, ard elevated tail; wings extremely short, so that the first primary feathers were not longer than those of a smail 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, ofien 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 42 inches in length. The Hinkel- or Florentiner Taube of Neumeister (Table Kite fig. 1) agrees with the above description in all the specified charac- ters (for the beak is not mentioned), except that Neumeister expressly says that the neck is short, whereas in my Scanderoon it was remarkably long aud bowed; so that the Hinkel forms a well-marked variety. Sub-race Il, Pigeon eygre and Pigeon bagqadais of Bottard and Corbié (Scanderoon of French writers).—l 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) 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+ inches in lencth, and therefore 2; 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 Il], Spanish and Roman Runts.—I am not sure that I am right in placing these Runts in a distinct sub-race; yet, if we take well-characterized birds, there can be no doubt of the propriety of the separation. They are heavy, massive birds, with shorter. necks, legs, and beaks than in the foregoing races. The skin over the nostrils is swollen, but not carunculated ; the naked skin round the eyes is not very wide, and only slightly carunculated; and I have seen a fine so-called Spanish Runt with hardly any naked skin round the eyes. Of the two varieties to be seen in England, one, which is the rarer, has very long wings and tail, and agrees pretty closely with the last sub-race ; “the other, with shorter wings and tail, is apparently the Pigeon romain ordinaire of Boitard and Corbié. nap, V DESCRIPTION OF BREEDS. 151 These Runts are apt to tremble like Fantails. They are bad flyers. A few years ago Mr. Gulliver ™ exhibited a Runt which weighed 1 Ib. 14 oz ; and, as ] am informed by Mr. Tegetmeier, two Runts from the south of France were lately exhibited at the Crystal Palace, each of which weighed 2 lbs. 23 0z. ] Elliot, 32 is the standard number ‘Tey YSTsuy—'1z *Biyy "97390°ST13AN7 } athers le te r double row; their permanent fanlike z the tail. + f a © less valued than the position and expansion 0! are arranged in an irregula Cuap. V. DESCRIPTION OF BREEDS. 155 expansion and their upward direction are more remarkable characters than their increased number. The tail is capable of the same move- ments as in other pigeons, and can be depressed so as to sweep the ground. It arises from amore expanded basis than in other pigeons; and in three skeletons there were one or two extra coccygeal vertebree. I have examined many specimens of various colours from different countries, and there was no trace of the oil-gland; this is a curious ease of abortion.’ 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 move- ments of the neck, in the manner of walking, and in the breadth of the breast, the differences so graduate away, that it is impossible to make more than one sub-race. Moore, however, an excellent old authority,“ says, that in 1735 there were two sorts of broad-tailed shakers (7. 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 isan existing German Fantail with a thicker and shorter beak. Sub-race IT, Java Fantail.—Myr. 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. 13 This gland occurs in most birds ; but Nitzsch (in his ‘ Pterylographie,’ 1840, p. 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 desta- tute of an oil-gland, have an unusual number of tail-teathers, namely 16, and in this respect resemble Fantails. 14 See the two excellent editions published by Mr. J. M. Eaton in 1852 and 1858, entitled ‘A Treatise on Fancy Pigeons.’ 156 DOMESTIC PIGEONS: Cuapr. V. Race VI.—Tursir anp Ow1. (Méventauben; pigeons a cravate. ) Feathers divergent along the front of the neck and breast; beak very short, vertically rather thick ; esophagus 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 cesophagus, which causes a movement in the frill. When the cesophagus ofa dead bird isinflated, itisseento be larger than in other breeds, and not so distinctly separated from the crop. The Pouter inflates both its true crop and esophagus; the Turbit inflates in a much less degree the cesophagus alone. The beak of the Turbit is very short, being ‘28 of aninch 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. Rack VII.—Tumsiers. (Tiimmler, or Burzeltauben; cul- butants. ) During flight, tumble backwards ; body generally small ; beak generally short, sometimes excessively short and conical. This race may be divided into four sub-races, namely, Persian, Lotan, Common, and short-faced Tumblers. These sub-races in- clude many varieties which breed true. I have examined eight skeletons of various kinds of Tumblers: excepting in one imperfect and doubtful specimen, the ribs are only seven in number, whereas the rock-pigeon has eight ribs. Sub-race I, Persian Tumblers.—I 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 dovecot pigeon, white and mottled, slightly feathered on the feet, with the beak just perceptibly shorter than in the rock-pigeon, H.M. Consul, Mr. Keith Abbott, informs me that the difference in the length of beak is so slight, that only practised Persian fanciers can distinguish these Tumblers from the common pigeon of the country. He informs me that they fly in flocks high up in the air and tumble well. Some of Cuap. Y. DESCRIPTION OF BREEDS. Log them occasionally appear to become giddy and tumble to the ground, in which respect they resemble some of our Tumblers. African OWL, Fig, 22. Sub-race IT, 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. 158 DOMESTIC PIGEONS: Cuar. ¥ Elliot are white, slightly feathered on the feet, with the feathers on the head reversed ; and they are rather smaller than the rock or dovecot pigeon. The beak is proportionally only slightly shorter and rather thinner than in the rock-pigeon. These birds when gently shaken and placed on the ground immediately begin tumbling head over heels, and they continue thus to tumbie 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 remark- able 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.’ © 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 roli over if only touched on the neck with a rod or wand. Sub-race III, Common English Tumblers. — These birds have exactly the same habits as the Persian Tumbler, but tumble better. The English bird is rather smaller than the Persian, and the beak is plainly shorter. Compared with the rock-pigeon, and propor- tionally 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 featker-footed. They tumble to an extraordinary degree; as Mr. Brent remarks,’® “ 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,”'’ which is a Scotch variety, not 15 English transiation, by F. Glad- win, 4th edition, vol. i. The habit of the Lotan is also described in the seen at any of the Calcutta bird- dealers.” 16 ¢ Journal of Horticulture,’ Oct. Persian treatise before alluded to, published about 100 yearsago: at this date the Lotans were generally white and crested as at present. Mr. Blyth describes these birds in * Annals and Mag. of Nat. Hist.,’ vol. xiv., 1847, p. 104; he says that they “may be 22, 1861, p. 76. 17 See the account of the House- tumblers kept at Glasgow, in the ‘ Cot- tage Gardener,’ 1858, p. 285. Also Mr. Brent’s paper, ‘ Journal of Horti culture,’ 1861, p. 76. Cuap. Y. DESCRIPTION OF BREEDS. 159 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 “oround. Some fly round with the flock, throwing a clean summer- “ sault every few yards, till they are obliged to settle from giddiness “and exhaustion. ‘These are called Air Tumblers, and they com- “monly throw from twenty to thirty summersaults in a minute, “each clear and clean. I have one red cock that J have on two or “three occasions timed by my watch, and counted forty summer- “saults 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 fiying, 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 move- “ment which they seem to try to prevent. I have seen a bird some- “times in his struggles fly a yard or two straight upwards, the “impulse forcing him backwards while he struggles to go forwards. “Tf 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. Sub-race IV. Short-faced Tumblers——These are marvellous birds, and are the gloryand 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 Columbide. Their heads are nearly globular and upright in front, so that some fanciers say’® “the head should resemble a cherry with a barley- corn stuck in it.” These are the smallest kind of pigeons. Mr, Esquilant possessed a blue Baldhead, two years old, which when alive weighed, before feeding-time, only 60z. 5drs.; two others, each weighed 7o0z. We have seen that a wild rock-pigeon weighed 14 oz. 2 drs., and a Runt 384 0z.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 18 J. M. Eaton’s ‘Treatise on Pigeons,’ 1852, p. 9. ‘seiquiny, ystsug peor -110qS— £3 “Bi —$_$____—= Tp OAL OM STLLNE i __ += = Cuap. V. DOMESTIC PIGEONS 160 to have shorter beaks; but proportionally with the size of the body, So, again, the feet of this bird the beak is ‘28 of an inch too short. nap. V. DESCRIPTION OF BREEDS. 161 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 scutelle. 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 occa- sionally 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.” Finally, in regard to the whole group of Tumblers, it is impos- sible 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 genius with the rock-pigeon. The ditferences between the successive steps in this series are not greater than those which may be observed between common dovecot-pigeons (C. livia) brought from different countries. Race VIII.—Inpran FRrint-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 isasmallish 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 « distinct breed. Tro- bably this is the breed seen by Hasselquist in 1757 at Cairo, and said to have been imported from India. Race IX.—Jacopix. (Zopf- or Perrfickentaube ; nonnain.* Feathers of the neck forming a hood; wince ond éail long ; beak moderately short. This pigeon can at once be recognised by its hood, almost enclos- ing the head and meeting in front of the neck. The hood seems to be merely an exaggeration of the crest of reversed feathers on tha back of the head, which is commoa to many sub-varieties, and 19 J. M. Eaton’s Treatise, edit. 1858, p. 76. 1§2 DOMESTIC PIGEONS: — Cuap. Y. which in the Latztaube™ 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 14 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+ inches too long, and the two wings, from tip to tip, 5+ inches too long. In disposition this bird is singularly quiet, seldom flying or moving about, as Bechstein and Riedel have likewise remarked in Germany. 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 LY. 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.—Trumrerer. (Trommeltanbe; pigeon tambour, elouglou.) 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 con- tinued for several minutes; hence their name of Trumpeters. They are also characterised by a tuft of elongated feathers, which curls forward over the base of the beak, and which is possessed by ne other breed. Their feet are so heavily feathered, that they almost appear Jike 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. 70 Neumeister, ‘Taubenzucht,’ Tab. s. 26. Bechstein, ‘ Naturgeschichte 4. fig. i. Deutschlands,’ Band iv. s. 36, 1795. 21 Riedel, ‘ Die Taubenzucht,’ 1824, Crap. V. DESCRIPTION OF BREEDS. 163 Race XI.—Scarcely differing in structure from the wild Columbia livia. Gub-race I, Laughers. Size less than the Rock-pigeon ; voice very peculiar.—As this bird agrees in nearly all its proportions with the ruck-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 Tangher is very different from that of the Trumpeter, yet one of my Trumpefters 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 IT. Common Frill-back (die Strupptaube). Beak rather longer than in the rock-pigeon ; feathers reversed.—This is a consider- ably larger bird than the rock-pigeon, and with the beak, propor- tionally 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 111. Nuns (Pigeons coquilles). These elegant birds are smaller than the rock-pigeon. The beak is actually 1°7, and propor- tionally 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 scutellz 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 inany 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 IV. Spots (die Blasstauben; pigeons heurtés).—These 164 DOMESTIC PIGEONS: Cuap. Ve birds are a very little larger than the rock-pigeon, with the beak a trace smailer 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; and in 1735 Moore remarks that they breed truly, as is the case at the present day. Sub-race V. Swallows.—These birds, as measured from tip to tip of wing, or from the end of the beak to the end of the tail, exceed in size the rock-pigeon; but their bodies are much less bulky; their feet and legs are likewise smaller. The beak is of about the same length, but rather slighter. Altogether their general appear- ance 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* 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 state- ments may be trusted I know not; but the inheritance of any habit may be believed, after what we have seen with respect to the Ground-tumbler of India. MM. Boitard and Corbié describe a pigeon ** 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 Aldroyandi in 1600 to the present day, in the accounts published of the Draijers, Smiters, Finnikins, Turners, Claquers, &c., which are all remark- able 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® briefly describes and figures this bird, with a tail “having 22 Willughby’s‘ Ornithology,’ edited 24 Pigeon pattu plongeur. ‘Les by Ray. Pigeons,’ &c., p. 165. *3 J. M. Eaton’s edition (1858) of 23 * Naturgeschichte Deutschlands, Moore, p. 98. Band iv. s. 47. Cuap. V. DESCRIPTION OF BREEDS. 163 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 Belguim has lately been exhibited at the Philoperisteron Society in London,”® which “ con- joins 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 153 inches from tip to beak to end of tail, and 323 inches from tip to tip of wing; now the wild rock-pigeon weighs 143 ounces, and measures from tip to beak to end of tail 15 inches, and from tip to tip of wing only 262 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 like- wise 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 Columbide, 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, suck 2° Mr. W. B. Tegetmeier, ‘Journal of Horticulture,’ Jan. 20th, 1863, p. 58. - Oa ee he: 166 DOMESTIC PIGEONS: Cuap. V as Turbits and short-faced Tumblers. The remaining do- mestic forms might have been included,in the same genus with the wild rock-pigeon. Indizidual Variability ; variations of a remarkable nature. The differences which we have as yet considered are charac- teristic 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 indi- vidual 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 here- after 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 humerous, 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 Columbide, which never (as far as I can hear) have less than twelve or more than sixteen tail-feathers; and these numbers cha- Unap. V INDIVIDUAL VARIABILITY. 167 racterise, with rare exception, whole sub-families.” 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. Onthe 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 Pald-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 Columbide, as far asI 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 Baldhead- Tumblers. Mr. Brent, however, had an Air-T'umbler (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. Ihave 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 aspecially noticed this in Jacobins. In Mr. Bult’s magnificent col- 27 ¢Coup-d’eil sur l’Ordre des Pi- geons, par C. L. Bonaparte (‘ Comptes Rendus’), 1854-55. Mr. Blyth, in ‘Annals of Nat. Hist.,’ vol. xix., 1847, p. 41, mentions, as a very singular fact, “that of the two species of Ectopistes, which are nearly allied to each other, one should have fourteen tail-feathers, while the other, the passenger pigeon of North America, should possess but the usual numer —twelve.” 168 DOMESTIC PIGEONS: Cuap. V. lection 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 Columbide. 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 itis 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 Faintail 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 scutellz# on the toes often varies in the same breed, and sometimes even differs on the two feet of the same indi- vidual; the Shetland rock-pigeon has fifteen on the middle, and six on the hinder toe; whereas | 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. Onthe 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™ 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 28 Described and figured in the ‘ Poultry Chronicle,’ vol. iii., 1855, p. 82. Cuap. V. SINGULAR VARIATIONS. 169 at the Philoperisteron Soc. I bred a mongrel pigeon which had fibrous feathers, and the wing and tail-feathers so short and imper- fect that the bird could not fly even a foot in height. There are many singular and inherited peculiarities in the piumage 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.”?9 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.°° 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 earriage,*! 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 amcunt 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 29 “The Pigeon Book,’ by Mr. B. P. 51 “A Treatise on the Almond-Tum. Brent, 1859, p. 41. bler, by J. M. Eaton, 1852, p. 8, et 30 “Die ctaarhalsige Taube. Das passim, Ganze, &c.,’ s. 21, tab. i. fig, 4. 170 DOMESTIC PIGEONS: Cuapr. V. 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 sueeessful breeder of prize Fantails, informs me that his male birds often have a greater number of tail-feathers than the females. Mr. Eaton asserts *? 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 repro- duction, 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 ** 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** that in certain light-coloured pigeons the males have their feathers striated with black, and these striz increase in size at each moult, so that the male ultimately becomes spotted with black. With Carriers, the wattle, both 32 A Treatise, &c., p. 10. generally females, and of the ease 33 Boitard and Corbie, ‘Les Pigeons,’ with which a race thus characterised &e., 1824, p. 173. could be produced. Bonizzi (see 34 ¢Te Pigeon Voyageur Belge? ‘ Variazioni dei Columbi domestici :’ 1865, p. 87. I have given im my Padova, 1873) states that certain ‘Descent of Man’ (6th edit. p.466) coloured spots are often different in some curious cases, on the authority the two sexes, and the certain tints of Mr. Tegetmeier, of silver-coloured are commoner in females than in male (i.e. very pale blue) birds being pigeons. Cuap. V. OSTEOLOGIAL DIFFERENCES. 17 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 Columbide.*® Osteological Characters. In the skeletons of the various breeds there is much varia- bility; 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, enas, palumbus, and turtur, which are ranked by all systematists in two or three distinct though allied genera, that the differences are extremely slight, certainiy 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. 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 35 Prof. A. Newton (‘ Proc. Zoolog, _ family ef the Treronide the sexes often Soc.,’ 1865, p. 716) remarks that he differ considerably in colour, See knows no species which present any also on sexual differences in the Colum- remarkablesexual distinction; but Mr. _ bide, Gould, ‘Handbook to the Birds Wallace informs me, that in the sub- of Australia,’ voi. ii. pp. 109-149. 2 DOMESTIC PIGEONS: Cuar. V. 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 elon- Fig. 24.—Skuills of Pigeons viewed laterally, of natural size. A. Wild Kock-pigeon, Columba livia. b. Short-faced Tumbler. C: English Carrier. D. Bagadvtten Currier. gation of the bones of the face, the space between the orbits is pro- portionally a little narrower than in the rock-pigeon. In the Baga- dotten the upper mandible is remarkably arched, and the premaxil- lary bones are proportionally broader In the Short-faced Tumbler 1s t -™” Cuapr. V. OSTEOLOGICAL DIFFERENCES. 173 the skull 1s 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 pro- minent edges of the eye-orbits is depressed. In the Barb the pre- maxillary 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. In the lower jaw, the articular surface is proportionably smaller in many breeds than in the rock-pigeon ; and the vertical diameter, Fig. 25.—Lower jaws, seen from above, of natural size. A. Rock-pigeon. B, Runt, C. Barb, more especially of the outer part of the articular surface, is con- siderably 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 re- markable manner; and the superior margin of the ramus, beyond the middle, is reflexed in an equally remarkable manner, as may be seen in the accompanying figures, in comparison with the jaw of the rock-pigeon. ‘This 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 fig. 26 of the head of a Runt seen from above; here a wide open space may be observed on each side, between the edges of the lower jaw and of the premaxillary bones. i ne cy ie 174 DOMESTIC PIGEONS: Cap. V. 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 Fig. 27.—Lateral view of jaws, of natural size. A. Rock-pigeon. B. Short-faced Tumbler. C. Bagadotten Carrier. that no open space is left. The degree of downward curvature of the distal half of the lower jaw also differs to an extra- ordinary degree in some breeds, as may be seen in the drawings (fig. A) of the rock- Fig. 26.—Skull of Runt. seen pigeon, (B) of the Short-faced Tumbler from above, of natural size, = ee pone pene, showing the reflexed margm ald (Cc) of the Bagadotten Carrier of Neu- of the distal portion of the meister. In some Runts the symphysis of lower jaw. 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. Vertebree.—All the breeds have twelve cervical vertebre.*° 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 verteh-ee are always eiglt. In the rock-pigeon all eight bear ribs; the eight 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 36 [ am not sure that I havede- ferent rules, and, as I use the same signated the different kinds of vertebre terms in the comparison of all the correctly: but I observe that different skeletons, this, I hope, will not anatomists foliow in this respect dit- signify. Cap. V. OSTEOLOGICAL DIFFERENCES, 1s seventh pair had distinct processes. In many breeds there are only seven ribs,as in seven out of eight skeletons of various Tumblers, and iu 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 ihe 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 teud to form an ossified arch. The development of the arch, formed by the hypapophyses of the third and fourth dorsal vertebree, also varies considerably, as does the size of the hypapo- physis of the fifth vertebra. The rock-pigeon has twelve sacral vertebre; 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 vertebre. Jn 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 vertebre. The caudal vertebre 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 vertebree. I have seen Bue in a Nun and J acobin. 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 vertebre and ribs which I have observed :— | | | | - | Pouter, from | ‘Tumbler, Bussorah | | | EEE soe Mr. Bult. | Iutch Roller. Carrier. ier. | | Cervical Veitebree | 12 12 | 12 | | H H The ee bore | a small rib. | Dorsal Vertebrze | 8 | 8 8 8 » Ribs 8 8 7 Al The 6th Pair with The 6th and The 6th and | The 6th and processes, the 7th 7th-pair with 7th pair with- | 7th pair with- | pair without a | processes. out processes. | out processes. process. | | Sacral Vertebre 12 14 1i il | Caudal Vertebrze 7 [28 on Ger ai 7 | Total Vertebree 3y | 42 or 43 38 38 176 DOMESTIC PIGEONS: The pelvis differs very little in any breed. The anterior margin of the ilium, however, is sometimes a little more equally rounded on both sides than in the rock-pigeon. The ischium is also frequently rather more elongated. The obturator-notch is sometimes, aS in many Tumblers, less developed than in the rock-pigeon. The ridges on the ilium are very prominent in most Runts. In the bones of the extremities. I could detect no difference, except in their pro- Fig. 28.—Scapule, of natural. portional lengths ; size. A. Rock-pigeon. b. for instance, the Suort-faced Tumbler. 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 the woodcut, fig. 28, the scapulz of the rock-pigecn (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, forma more perfect cavity some Tumblers than in the rock-pigeon: in Pouters these processes are larger and dif ferently shaped, and the exterior angle of the extremity of the coracoid, which is articulated to the sternum, is squarer. 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. Infig. 29,B and c represent the furcula of two Fantails; and Fic. 29.—Yrureala, of natural size. A. SLore-faced jo um- B and C Fantail. D. it will be seen that the divergence in B is rather less even than in the Cuap. V. CORRELATION OF GROWTH. 177 furculum of the short-faced, small-sized Tumbler (4), whereas the divergence in c equals that in a rock-pigeon, or in the Pouter (p), though the latter is a much larger bird. The extremities of the furcu- lum, 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 ease 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 indi\iduals 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 con- tinued 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 tavu 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 Oi. 178 DOMESTIC PIGEONS: Cuap. V. 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 woodcut No. 27) in the curvature of the lower jaw in the rock-pigeon, the Tumbler, and Baga- dotten 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 woodent No. 25) is not strictly correlated with the width or divergence (as may be clearly seen in woodcut No. 26) of the premaxillary benes, 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 b dy is a selected cha- racter, and the ribs, as we have seen, have generally become very broad, with the seventh pair furnished with processes; the Cuap. V. CORRELATION OF GROWTH. V7Y 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 inde- pendently 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 37 reduced the standard for Bald-head Tumblers from ten to nine white Hight-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 inexpli- cable 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 ; 37 J, M. Eaton’s Treatise, edit. 1858, p. 78. 180 DOMESTIC PIGEONS: Cap. V. 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 cor- relation 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 scutelle or scales covering the toes and tarsi have not only decreased or increased in size, but Hkewise in number. To give a single instance, I have counted eight scutellz 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 scutelle 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 Ejjects of Disuse.—In the following discussion on the relative proportions of the feet, sternum, furculum, scapule, 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. 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 un- fortunately (except in a few cases) not to the root of the tail; I Cuap. V ON THE EFFECTS OF DISUSE. 181 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 ; TABLE. I. Pigeons with thetr beaks generally shorter than that of the Rock-piyeon, proportionally to the size of their bodies. Difference between actual and calculated Actual length of feet, in Name of Breed. length | proportion to length ot of feet and size of body Feet in the Rock-pigeon, Too short | Too long Wild rock-pigeon (mean measurement) .. 2°02 by by Short-faced Tumbler, bald head 1°57 0-11 almond 1:60 0°16 Tumbler, red ° macpie ss 1°75 0-19 red common (by standard to end of tail). 3 peaahiee Sta aes 1°85 0:07 ts common bald-head 1°85 0°18 - roller 1°80 0°06 Puchi. 1°75 OFA ¥ 1:80 | 0-01 : 1°84 0°15 Jac obin . 1°90 0:02 er umpeter, white 2°02 0°06 mottled .. 1 95 0°18 Fantail (by standard to end of tail) 1°85 0 15 3 a 95 0°15 oe E crested var. 45 95 0°0 00 Indian Frill-back - 1-380 0°19 Be English Frill-back 2°10 0°03 LT eal 1°82 0°02 Laugher 1°65 0°16 Barb 2°00 0°03 Fe “ 2-00 re 0°03 Spot 1-90 0 02 ae 5 rae 1:90 0°07 Swallow, red 1°85 0°18 ss 3 blue oe oe ee 2-00 DO 0°03 POUT iene pie sf ea mee 2°42 Bc 0-11 ze German 2°30 a“ 0°09 Bussorah Carrier.. aed eP| | 0°09 Number of specimens .. .. .. 28 | 22 On a x ie = f 4 ’ 1&2 DOMESTIC PIGEONS: Cuap. V. 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 ofthe 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. TaBe IT. Pigeons with thetr beal:s longer than that of the Rock-pigeon, proportionally to the size of their bodies. | Difference between ; actual and calculated Actnal | length of feet, in Name of Breed. length proportion to length of © ; feet and size of body Feet | in the Rock-pigeon. =: . Too short Too long Wild rock-pigeon (mean measurement) .. 2°02 by | aby Gamers oo eee ee Nae oe Vee eee eee POss8 - 2°60 0-25 e he Slee! 2°40 0-21 pe Dragon 2°25 0°06 Bagadotten Carrier 2°80 0°56 Scanderoon, white Spare 2°80 0°3 * Pigeon cygne.. -.. .. «. | 2°85 0°29 Runt ots eae 0°27 Number of specimens .. .._ .. | 3 3) en 8 Tn these two tables we see in the first column the actual length of the feet in thirty-six birds belonging to various breeds, and in the two other columns we see by how much the feet are too shore or too long, according to the size of bird, in comparison with the rock-pigeon. In the first table twenty-two specimens have their Cuap, V. ON THE EFFECTS OF DISUSE. 1838 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 eases can be explained ; for instance, with Pouters the legs and feet are selected for length, and thus any natural tendency to a dimi- nution 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. Tn the second table we have eight birds, with their beaks much longer than in the rock-pigeon, both actually and proportionally with the size of body, and their feet are in an equally marked manner longer, namely,in proportion, on an average by ‘29 of an inch. I should here state that in Table I. there are a few partial exceptions to the beak being proportionally shorter than in the rock-pigeon : thus the beak of the English Frill-back is just perceptibly longer, and that of the Bussorah Carrier of the same length or slightly longer, than in the rock-pigeon. The beaks of Spots, Swallows, and Laughers are only a very little shorter, or of the same proportional length, but slenderer. Nevertheless, these two tables, taken conjointly, indicate pretty plainly some kind of correlation between the length of the beak and the size of the feet. Breeders of cattle and horses believe that there is an analogous connection between the Jength of the limbs and head; they assert that a race-horse with the head ofa 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, ** and that this reduction has acted by correlation on the beaks of the great majority of the birds in Table I. When, on the other hand, the beak bas been much elongated by the continued selection of successive shght increments of length, the feet by corre- lation 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 nas 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. 3° In an analogous, but converse, aHied groups, have larger feet. Seé manner, certain natural groups of Prince Bonaparte’s ‘ Coup-d’eil sur the Columbide, from being more ter- _‘|’Order des Pigeons.’ restrial in their habits than other 184 DOMESTIC PIGEONS: Cuar. V. 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 iess than the wild rock-pigeon. Hence it seemed to me probable that all the parts cf the skeleton subservient to flight would be found to be reduced in size. Withrespect tothe sternum, I have carefully measured its extreme length in tweive birds of different breeds, and in two wd rock-pigeons from the Shetland Islands. For the proportional comparison I have tried three standards of measurement, with ali 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. Length of Sternum. | Actual | Too | Actual Name of Breed. pene Short by Name of Breed. eee short by Wild Rock-pigeon | 2°55 | .. || Barb | 2-35 | 0-34 Pied Scanderoon .. | 2°80 | 0°60 | Nun ss ee | 2°27 | 0-15 Bagadotten Carrier | 2°80 | 9°17 | German Pouter .. | 2°36 | 0°54 Dragon 2-4] Oc41 4 Jacobim :.. sa") ce | 2°33 0°22 Carrier .. .. .. | 2°75 | 0°35 | English Frill-back | 2-40 | 0-43 Short faced Tumbler | 2-5 = 0°28 Swallow Seen | eee | 0:17 This table shows that in these twelve breeds the sternum is of an average one-third of an inch (exactly 352) 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- cighth 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 twoof 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 shght, 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-pigecn. I have measured the length of the scapula in nine different large Cuap. V. ON THE EFFECTS OF DISUSE. 185 and small-sized breeds, and in all the scapula is prcportioually 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 sborter. Thus in a Runt, which measured from tip to tip of wings 383 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 264 inches. Ina 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 cnly 23 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, scapule, 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 13 inch; whilst in the five birds they were less in length by only 8 of aninch. AsI was much surprised that the wings of closely confined birds should thus so frequentiy 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 L86 DOMESTIC PIGEONS: Cuap. V. 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,*? 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 scapule 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 scapulz 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. ‘lhe 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 3° It perhaps deserves notice that would, therefore, appear as if, during besides these five birds two of the the reduction of their beaks, their 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 wings had retained a little of that excess of length which is characterise tic of their nearest relations and pros genitors, Cap. V. SUMMARY OF DIFFERENCES. 187 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 in- dependently 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 develop ment of the wattle. The size and form of the cesophagus 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 vertebrx, and the length of the sternum, all vary. ‘The number and size of the coccygeal vertebra vary, apparently in correlation with the increased size of the tail. ‘The size and shape of the perfora- tions 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. ‘he 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 occasion- ally vary in number, apparently in correlation with the le.gth of the wing. The length of the leg and the size of the feet, and, in connection with the latter, the number of the scutellz, all vary. A web of skin sometimes connects the bases of the two inner toes, and almost invariably the twa outer toes when the feet are feathered. The size of the body differs greatly : a Runt has been known 188 DOMESTIC PIGEONS. Cuap, V. to weigh more than five times as much as a Short-faced Tumbler. The eggs differ in size and shape. According te Parmentier,*? some races use much straw in building their nests, and others use little; but I cannot hear of any recent corroboration of this statement. ‘The length of time required for hatching the eggs is uniform in all the breeds. The period at which the characteristic plumage of some breeds is acquired, and at which 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 faney pigeons up to the proper standard of excellence than the so- called toy pigeons, which differ from each otker 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 secon- dary sexual characters,*? of which the aboriginal rock-pigeon displays not a trace. 40 Temminck, ‘Hist. Nat. Gen. des bet ween the males and females, as are Pigeuns et des Gallinacés, tom. i, not directly connected with the act of 1813, p. 170. reproduction, as the tail of the peas 41 This term was used by John cock, the horns of deer, &c. Hunter for such differences in structure Cuap. VI. DOMESTIC PIGEONS: THEIR PARENTAGE. 189 GHAPERR: VA: PIGEONS—continued. ON THE ABORIGINAL PARENT-STOCK OF THE SEVERAL DOMESTIC RACES— HABITS OF LIFE—WILD RACES OF THE ROCK-PIGEON—_DOVECOT-PIGEONS— PROOFS OF THE DESCENT OF THE SEVERAL RACES FROM COLUMBA LIVIA —FERTILITY OF THE RACES WHEN CROSSED—REVERSION TO THE PLUMAGE OF THE WILD ROCK-PIGEON—CIRCUMSTANCES FAVOURABLE TO THE FOR- MATION OF THE RACES—ANTIQUITY AND HISTORY OF THE PRINCIPAL RACES—MANNER OF THEIR FORMATION—SELECTION—UNCONSCIOUS SE- LECTION—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 ot 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! 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, 1 Temminck, ‘ Hist. Nat. Gén. des Pigeons,’ &c., tom. i. p. 191. Crap. VI 190 DOMESTIC PIGEONS: was evident.2, 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? has been assured that the wild C. livia, var. intermedia, sometimes roosts in trees. I may here give a curious instance of compulsion leading to changed habits: the banks of the Nile above lat. 28° 30' are perpendicular for a long distance, so that when the river is full the pigeons cannot alight on the shore to drink, and Mr. Skirving repeatedly saw whole flocks settle on the water, and drink whilst they floated down the stream. These flocks seen from a distance resembled flocks of gulls on the surface of the sea. If any domestic race had descended from a species which was not social, or which built its nest and roosted in trees,* 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 occa- sionally even young fowls, when the hen gives the danger- ery, run away and try to hide themselves, like young par- tridges or pheasants, In order that their mother may take 2 [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. 3 ¢Annals and Mag. of Nat. Hist.,’ 2nd ser., vol: xx., 1857, p. 509 ; and in « late volume of the Journal of the Asiatic Society. * 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. Crap. VI. THEIR PARENTAGE. 191 flicht, of which she has lost the power. The musk-duck (Cairina moschata) in its native country often perches and roosts on trees,> and our domesticated musk-ducks, though such sluggish birds, “are fond of perching on the tops of barns, walls, &c., and, if allowed to spend the night in the hen-house, the female will generally go to roost by the side of the hens, but the drake is too heavy to mount thither with ease.” ® We know that the dog, however well and regularly fed, often buries, like the fox, any 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 domesticatcd 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 re- marked, is not likely to have been the parent of our domestic breeds, which thrive in the hottest countries. Secondly, the C. rupestris, of Central Asia, which is intermediate’ between the C. leuconota and livia; but has nearly the same coloured tail 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 m 5 Sir R. Schomburgk, in ‘ Journal 6 Rey. E. S. Dixon, ‘Ornamental R. Geograph. Soc.,’ vol. xiii., 1844, Poultry,’ 1848, pp. 63, 66. p. 32. 7 Proc. Zoolog. Soc., 1859, p. 400. 1$2 DOMESTIC PIGEONS: Cuap. VI. fact belong to the genus Carpophaga. Fourthly, Columba guine:, which ranges from Guinea* 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 Sirictcenas 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 characterized by the feathers of the neck having peculiar notched tips,—a character not observed in any domestic race. Fifthly, the Columba enas 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 breecs. This bird agrees in every essential character with the breeds which haye been only slightly modified. It differs from all other species in being of a slaty-blue colour, with two black bars on the wings, aud with the croup (or loms) 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 ? C. amalie, but this species has not been admitted as distinct by other ornithologists. Graba’® 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” as C. ajinis, but 1s now no longer considered by him as a distinct species. C. afinis 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 8 Temminck, ‘ Hist. Nat. Gén. des Pigeons,’ tom. i.; also ‘ Les Pigeons, par Mme. Knip and Temminck. Bona- parte, however, in his ‘ Coup-d'eil,’ believes that two closely allied species are confounded together under this name. The C. lewcocephala of the West Indies is stated by Temminck to be a rock-pigeon ; but 1am informed by Mr. Gosse that this is an error. ° ‘Handbuch der Naturgesch. Vogel Deutschlands.’ 10 *Tagebuch, Reise nach Faro,’ 1830, s. 62. 11 «Annals and Mag. of Nat. Hist.’ vol. xix. 1847, p. 102. This excellent paper on pigeons is well worth con- sulting. COnap. VI. THEIR PARENTAGE. 193 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 te 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 ” says that at Islay fully half the wild rock-pigeons were chequered. Colonel King, of Hythe, stocked his dovecot with young wild birds which he himself procured from nests at the Orkney Islands; and several specimens, kindly sent to me by him, were all plainly chequered. As we thus see that chequered birds occur mingled with the true rock-pigeon at three distinct sites, namely, Faroe, the Orkney Islands, and Islay, no importance can be attached to this natural variation in the plumage. Prince C. L. Bonaparte, a great divider of species, enumerates, with a mark of interrogation, as distinct from CC. livia, the C. turricola of Italy, the C. rupestris of Daouria, and the C. schimperi of Abys- sinia; but these birds differ from C. ‘¢via in characters of the most trifling value. Inthe British Museum there is a chequered pigeon, probably the C. schimperi of Bonaparte, from Abyssinia. ‘To these may be added the C. gymnocyclus of G. R. Gray from W. Africa, which is slightly more distinct, and has rather more naked skin round the eyes than the rock-pigeon ; but from information given me by Dr. Daniell, it is doubtful whether this is a wild bird, for dovecot-pigeons (which I have examined) are kept on the coast of Guinea. The wild rock-pigeon of India (C. intermedia of Strickland) has been more generally accepted as a distinct species. It 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.livix, Moreover we shall immediately have proof that the blue and white croup is a highly variable character; and Bechstein “ asserts that with dovecot-pigeons in Germany this is the most variable of all the characters of the plumage. Hence it may be concluded that C. intermedia cannot be ranked as specifically distinct from C. livia, In Madeira there is a rock-pigeon which a few ornithologists have suspected to be distinct from C. livia. I have examined numerous specimens collected by Mr. E. V. Harcourt and Mr. Mason. They are rather smaller than the rock-pigeon from the Shetland Islands, and their beaks are plainly thinner, but the thickness of the beak varied in the several specimens. In plumage there is remarkable 12 Natural History of Ireland, geons,’ ‘Comptes Rendus,’ 1854-55. Birds, vol. ii. (1850), p. 11. For 14¢ Naturgeschichte. Deutschlands, Graba, see previous reference. Band iv. 1795, s. 14. 13 ¢Coup-d’eil sur |’Ordre des Pi- 194 DOMESTIC PIGEONS: Cuar. VL 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. efinis 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. liv/a, affinzs, 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 dovecot- pigeons, which are kept in various parts of the world, are descended from one or from several of the above-mentioned wild varieties of C. livia, no one who compares them will doubt. But before making a few remarks on dovecot-pigeons, it should be stated that the wild rock-pigeon has been found easy to tame in several countries. We have seen that Colonel King at Hythe stocked his dovecot more than twenty years ago with young wild birds taken at the Orkney Islands, and since then they have creatly multiplied. The accurate Macgillivray © asserts that he completely tamed a wild rock- -pigeon in the Hebrides; and several accounts are on records of these pigeons having bred in doyecots 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 asserts that wild birds come frequently to the dovecots and mingle freely with their inhabitants. In the ancient ‘ Ayeen Akbery’ it is written that, if a few wild pigeons be taken, “ they are speedily joined by a thousand others of their kind.” Dovecot-pigeons are those which are kept in doyvecots in a semi- domesticated state ; for no special care is taken of them, and they procure their own food, except during the severest weather. In England, and, judging from MM. Boitard and Corbié’s work, in France, the common dovecot-pigeon exactly resembles the chequered 13 ‘History of British Birds,’ vol.i. | rock-pigeon came and settled in his pp. 275-284. Myr. Andrew Duncan dovecot in Balta Sound in the Shet- tamed a rock-pigeon in the Shetland land Islands, and bred with his Islands. Mr. James Barclay,and Mr. pigeons; he has also given me other Smith of Uyea Sound, both say that instances of the wild rock-pigeon the wild rock-pigeon can be easily having been taken young and breed- tamed; and the former gentleman ing in captivity. asserts that the tamed birds breed 16 “Annals and. Mag. of Nat. four times a year. Dr. Lawrence History, vol. xix. 1847, p. 103, and Edmondstone informs me that a wild vol. for 1857, p. 512. Cuar. VI. THEIR PARENTAGE. 195 variety of C. livia ; but Ihave seen dovecots brought from Yorkshire without any trace of chequering, like the wild rock-pigeon of the Shetland Islands, The chequered dovecots from the Orkney Islands, after having been domesticated by Colonel King for more than twenty years, differed slightly from each other in the darkness of their plumage and in the thickness of their beaks ; the thinnest beak being rather thicker than the thickest one in the Madeira birds. In Germany, according to Bechstein, the common dovecot- pigeon is not chequered. In India they often become chequered, and sometimes pied with white; the croup also, as I am informed by Mr. Blyth, becomes nearly white. I have received from Sir. J. Brooke some dovecot-pigeons, which originally came from the S. Natunas Islands in the Malay Archipelago, and which had been crossed with the Singapore dovecots: they were small and the darkest variety was extremely like the dark chequered variety with a blue croup from Madeira; but the beak was not so thin, though decidedly thinner than in the rock-pigeon from the Shetland Islands. A dovecot- pigeon sent to me by Mr. Swinhoe from Foochow, in China, was likewise rather small, but differed in no other respect. I have also received through the kindness of Dr. Daniell, four living dovecot- pigeons from Sierra Leone,” these were fully as large as the Shetland rock-pigeon, with even bulkier bodies. In plumage some of them were identical with the Shetland rock pigeon, but with the metallic tints apparently rather more brilliant ; others had a blue croup, and resembled the chequered variety of C. intermedia of India; and some were so much chequered as to be nearly black. In these four birds the beak differed slightly in length, but in all it was decidedly shorter, more massive, and stronger than in the wild rock-pigeon from the Shetland Islands, or in the English dovecot. When the beaks of these African pigeons were compared with the thinnest beaks of the wild Madeira specimens, the contrast was great ; the former being fully one-third thicker in a vertical direction than the latter; so that any one at first would have felt inclined to rank these birds as specifically distinct; yet so perfectly graduated a series could be formed between the above-mentioned varieties, that it was obviously impossible to separate them. - To sum up: the wild Columba livia, including under this name C. affinis, intermedia, and the other still more closely- affined geographical races, has a vast range from the southern coast of Norway and the Faroe Islands to the shores of the Mediterranean, to Madeira and the Canary Islands, to Abys- sinia, India, and Japan. It varies greatly in plumage, being 17 Domestic pigeons of the common published in 1746; they are said, in kind are mentioned as being pretty | accordance with the name which they numerous in John Barbut’s ‘Descrip- _ bear, to have been imported. tion of the Coast cf Guinea’ (p, 215), 196 DOMESTIC PIGEONS: Cuap, VI. in many places chequered with black, and having either a white or blue croup or loins; it varies also slightly in the size of the beak and body. Dovecot-pigeons, which no one disputes are descended from one or more of the above wild forms, present a similar but greater range of variation in plumage, in the size of body, and in the length and thickness of the beak. There seems to be some relation between the croup being blue or white, and the temperature of the country inhabited by boih wild and dovecot pigeons; for nearly all the dovecot-pigeons in the northern parts of Europe have a white croup, lke that of the wild European rock- pigeon ; and nearly all the dovecot-pigeons of India havea blue croup like that of the wild C. intermedia of India. As in various countries the wild rcck-pigeon has been found easy to tame, it seems extremely probable that the dovecot-pigeons throughout the world are the descendants of at least two and perhaps more wild stocks; but these, as we have just seen, cannot be ranked as specifically distinct. With respect to the variation of C. livia, we may without fear of contradiction go one step further. Those pigeon- fanciers who believe that all the chief races, such as Carriers, Pouters, Fantails, &c., are descended from distinct aboriginal stocks, yet admit that the so-called toy-pigeons, which differ from the rock-pigeon in little except colour, are descended from this bird. By toy-pigeons are meant such birds as Spots, Nuns, Helmets, Swallows, Priests, Monks, Porcelains, Swa- bians, 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 dove- cot-pigeons. Although we may safely admit that dovecot- pigeons, which vary slightly, and that toy-pigeons, which Cuap. VI. THEIR PARENTAGE. 197 vary in a greater degree in accordance with their more highly- domesticated condition, are descended from @. 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 descen- dants 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, asin 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? Iam aware that some naturalists, following Pallas, believe that crossing gives a strong tendency to variation, indepen- dently of the characters inherited from either parent. ‘I hey 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 ina future chapter I shall have to recur to this subject. For our present purpose the point is not material. The question which concerns usis, whether or not many new and important characters have 198 DOMESTIC PIGEONS: Ciap. VI, 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 cross- ing 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 differ- ences 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 he unknown to orni- thologists. The hypothesis that such species formerly existed, but have become extinct, 1s 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 Cuap. VI. THEIR PARENTAGE. 199 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 lving 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. Second/y.—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 quad- rupeds. Nevertheless I have met with accounts showing that the common fowl has become feral in South America and perhaps in West Africa, and on several islands: the turkey was at one time almost feral on the banks of the Parana; and the Guinea-fowl has become perfectly wild at Ascension and in Jamaica. In this latter island the peacock, also, “has become a maroon bird.” The common duck wanders from its home and becomes almost wild in Norfolk. Hybrids between the common and musk-duck which have become wild have been shot in North America, Belgium, and near the Caspian Sea. The goose is said to have run wild in La Plata. The common dovecot- pigeon has become wild at Juan 200 DOMESTIC PIGEONS: Cuap. VL 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.!* 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 under- gone 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 intenticnally or by chance, a most abnormal set of pigeons; for that species resembling such birds as Pouters, Fantails, Carriers, Barbs, Short-faced Tumblers, Turbits, &c., would be in the highest degree abnormal, as compared with all the existing members of the great pigeon family, cannot be doubted. Thus we should have to believe that man not only formerly succeeded in thoroughly domesticating several highly abnormal species, but that these same species have since all become extinct, or 18 With respect to feral pigeons —for Juan Fernandez, see Bertero in ‘ Annal. des Sc. Nat.,’ tom. xxi. p. 351. For Norfolk Islands, see Rev. E. 8. Dixon in the ‘ Dovecote,’ 1851, p. 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 ‘ Annais of Nat. Hist.,’ vol. xx., 1857, p-911. For Scotland, see Macgillivray, ‘British Birds,’ vol. i. p. 275; also Thompson’s ‘Nat. Hist. of Ireland, Birds,’ vol. ii. p. 11. For ducks, see Rey. E. S_ Dixon, ‘ Ornamental Poultry,’ 1847, p. 122. For the feral hybrids of the common and musk- ducks, see Audubon’s ‘ American Or- nithology,’ and Selys-Longchamp’s ‘Hybrides dans la Famille des Ana- tides.’ For the goose, Isidore Geoffroy St.-Hilaire, ‘ Hist. Nat. Gén.,’ tom. iii. p- 498. For guinea-fowls, see Gosse’s. ‘Naturalist’s Sojourn in Jamaica,” p- 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, p. 42. For the turkey I rely on oral information; I ascer- tained that they were not Curassows. With respect to fowls I will give the references in the next chapter. Cuap. VI. THEIR PARENTAGE. 201 are at least now unknown. ‘This double accident is so ex- tremely 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 ce:tainly 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 cha racteristic 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 deve- lopment of the crop in Pouters, in the length of the beak in Tumblers, in the state of the wattle in Carriers, &c. If these characters are the result of successive variations added together by selection, we can understand why they should be so variable: for these are the very parts which have varied since the domestication of the pigeon, and therefore would be likely still to vary; these variations moreover have been recently, and are still being accumulated by man’s selection ; therefore they have not as yet become firmly fixed. Fifthly.—All the domestic races pair readily together, and, what is equally important, their mongrel offspring are per- fectly fertile. To ascertain this fact I made many experi- ments, which are given in the note below; and recently Mr. Tegetmeier has made similar experiments with the same result..? The accurate Neumeister asserts that when dovecots © T have drawn out a long table of | doubtedly have thus united all. The the various crosses made by fanciers between the several domestic breeds bat I do not think it worth while pub- lishing. I have myself made for this special purpose many crosses, and all were perfectly fertile. I have united -n one bird five of the most distinct races, and with patience I might un- 10 case of five distinct breeds being blended together with unimpaired fer- tility is important, because Gartner has shown that it is a very general, theugh not, as he thought, universal rule, that complex crosses between several species are excessively steriie. I have met with only two or thres 202 DOMESTIC PIGEONS: Cua. VL are crossed with pigeons of any other breed, the mongrels are extremely fertile and hardy.*? MM. Boitaid and Corbié?! affirm, after their great experience, that the more distinct the breeds are which are crossed, the more productive are their mongrel! offspring. J 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, &c., the fact of their perfect, or even increased, fertility when intercrossed in the most complicated manner becomes a strong argument in favour of their having all descended from a single species. This argument is rendered much stronger when we hear (I append in a note *? all the cases which I have collected) that cases of reported sterility in the off- spring of certain races when crossed. Pistor (‘Das Ganze der Feldtau- benzucht,’ 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 tertile. Temminck has stated (‘ Hist. Nat. Gén. des Pigeons,’ tom. i. p- 197) that the Turbit or Owl will not cross readily with other breeds: but my Turbits crossed, when left free with Almond Tumblers and with Trumpeters; the same thing has occurred (Rev. E. S. Dixon. ‘The Dovecot,’ p. 107) between Turbits and Dovecots and Nuns. I have crossed Turbits with Barbs, as has M. Boitard (p. 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. iv. s. 44. Turbits (Riedel, s. 26) have been crossed with Pouters and ith Jaccbips, 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. 8. Dixon’s explanation of such statements is correct, viz. that individual birds both with Turbits and other breeds are occasionally sterile. 20 *Das Ganze der Taubenzucht,’ seg hs 21 «Les Pigeons,’ &e., p. 35. 22 Domestic pigeons pair readily with the allied C. e@nas (Bechstein, * Naturgesch. Deutschlands,’ B. iv. 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. eas and a male Ant- werp Carrier) paired with a Dragon, but never laid eggs. Bechstein fur- ther states (s. 26) that the domestic pigeon will cross with C. palumbus, Lurtur risoria and TL. vulgaris, but nothing is said of the fertility of the hybrids, and this would have been mentioned had the fact been ascer- tained. In the Zoological Gardens Cuar. VL THEIR PARENTAGE. 203 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 differ- ences, 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 Columbide ; 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 court- ing the females; and all (with the exception of Tiumpeters (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. anasand gymnophthalmos were sterile. In Loudon’s ‘ Mag. of Nat. Hist.’ voi. vii. 1854, p. 154, it is said that a male hybrid (from TYurtur vulgaris male, and the cream-coloured 7. risorix female) paired during two years with a female TZ. risoria, and the latter laid many eggs, but all were sterile. MM. Boitard and Corbié (‘ Les Pigeons,’ p- 235) state that the hybrids from these two turtle-doves are invariably sterile both inter se and with either pure parent. The experiment was tried by M. Corbié “ avec une espéce d’obstination ;” and likewise by M. Mauduyt, and by M. Vieillot. Tem- minck also found the hybrids from these two species quite barren. There- fore, 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’ news- paper (in a letter dated Nov. 10th, 18538) 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 7. risoria: it would be an unparalleled fact if the same two species sometimes produced extremely fertile, and some- times extremely barren, offspring. In the MS. report from the Zoological Gardens it is said that hybrids from Turtur vulgaris and suratensis, and from ZT. vulgaris and Ectopistes migras torius, were sterile. Two of the latter male hybrids paired with their pure parents, viz. Zurtur vulgaris and the ketopistes, and likewise with 7. risoria and with Columba enas, and many eggs were produced, but all were barren. At Paris, hybrids have been raised (Isid. Geoffrey Saint-Hilaire, ‘Hist. Nat. Generale,’ tom. iii. p. 180) from Tutur auritus with T. can- bayensis and with 7. suratensis; but nothing is said of their fertilitv. At the Zoological Gardens of London the Goura coronata and victorie produced a hybrid which paired with the pure G. coronata, and laid several eggs, but these proved barren. In 1860 Columba gymnophthalmos and maculosa pyro: duced hybrids in these same gardens. 204 DOMESTIC PIGEONS: Cuap. VI. 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. Hach race presents nearly the same range of variation in colour; and in most of the races we have the same singular correlation between the develop- ment 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 Columbidz. In those races which present some remarkable deviation of structure, such as In the tail of Fantails, crop of Pouters, beak of Carriers and Tumblers, &c., the other parts remain nearly unaltered. Now every eae 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 election; 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 cer- tainly lead to other changes of structure thronghout the whole organization. On the other hand, if the several races of the pigeon have been produced by man threugh 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 f 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 Cuap. VI. THEIR REVERSION IN COLOUR. 205 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.” 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,‘ of 73 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 ex- 2eption signifies the less asthe Swallow approaches closely in structure to C. livia. bars are replaced by bars of various colours. The figures given by Neu- meister are sufficient to show that, if the wings alone are blue, the black wing-bars appear. 24 T 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 ex- amined had white, and two had blue In many sub-varieties the black 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 au- thentic accounts of at least two in- stances of the blue variety with black bars having appeared in England; blue Jacobins were bred by Mr. Brent from two black birds. I have seen common Tumblers, both Indian and English, and Short-faced Tumblers, of a blue colour, with black wing-bars, with the black bar at the end of the tail, and with the outer tail-feathers edged with white; the croup in all was blue, or extremely pale blue, never absolutely white. Blue Barbs and Trumpeters seem to be excessively rare; but Neumeister, who may be implicitly trusted, figures blue varie- ties of both, with black wing-bars. Mr. Brent informs me that he has seen a 206 DOMESTIC PIGEONS: Cuap. VI 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 equaily common in the wild C. livia, in dovecot-pigeons, and in all the most highly modified races. Thus, in all, the croup varies from white to blue, being most frequently white in Europe, and very generally blue in India.” We have seen that the wild C. livia in Europe, and dovecots in all parts of the world, often have the upper wing-coverts chequered with black; and all the most distinct races, when blue, are occasionally chequered in precisely the same manner. Thus I have seen Pouters, Fantails, Carriers, Turbits, Tumblers (Indian and English), Swallows, Bald-pates, and other toy-pigeons blue and chequered ; and Mr. Esquilant has seen a chequered Runt. I bred from two pure blue Tumblers a cheyuered bird. The facts hitherto given refer to the occasional appearance in pure races of blue birds with black wing-bars, and likewise blue Barb; and Mr. H. Weir, asl am Madras. A slaty-blue and chequered informed by Mr. Tegetmeier, once bred a silver (which means very pale blue) Barb from two yellow birds. 25 Mr. Blyth informs me that all the domestic races in India have tke croup Slue; but this is not invariable, for I possess a very pale blue Simmati pigeon with the croup perfectly white, sent to me by Sir W. Elliot from 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 (im- ported into Amoy, and thence sent me) has a perfectly white croup. Cur. VI. THEIR REVERSION IN COLOUR. 207 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, some- times chequered, with black wing-bars, &c.; or if not of a blue colour, yet with the several characteristic marks more or jess plainly developed. I was led to investigate this subject from MM. Boitard and Corbié?® having asserted that from crosses between certain breeds it is rare to get anything but bisets or dovecot pigeons, which, as we know, are blue birds with the usual characteristic marks. We shall here- after 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 com- plicated descent, namely, from a black Barb, a Spot, and 26 “Les Pigeons,’ &c., p. 37. 208 DOMESTIC PIGEONS: Cuar. VL 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?’ 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 1735.78 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. Ina second brood from the same parents a brown bird was produced, with several white feathers confined to the croup. 27 “Treatise on Pigeons,’ 1858, p. 28 J. Moore’s ‘ Columbarium,’ 1735; 145. in J. M. Eaton’s edition, 1852, p. 71. Cuap. VL THEIR REVERSION IN COLOUR. 209 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 *° 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 29 Tcould give numerous examples ; grey. Another mongrel whose four two will suffice. A mongrel, whose grandparents were a red Ruut, white 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 vail and tail-coverts were dark bluish- Trumpeter, white Fantail, and the same blue Pouter, was pure white tal 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 210 DOMESTIC PIGEONS: Cuar. VL 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 dovecot-pigeon and the chequered wild variety of the rock-pigeon, the statement before referred to by MM. Boitard and Corbié would almost suffice; but I will give three instances of the appearance of such birds from crosses in which one alone of the parents or great-grandparents was blue, but not chequered. Icrosseda male blue Turbit with a snow-white Trumpeter, and the following year with a dark, leaden-brown, Short-faced Tumbler; the offspring from the first cross were as perfectly chequered as any dovecot-pigeon ; and from the second, so much so as to be nearly as black as the most darkly chequered rock-pigeon from Madeira. Another bird, whose great-grandparents were a white Trumpeter, a white Fantail, a white Red-spot, a red Runt, and a blue Pouter, was slaty-blue and chequered exactly like a dovecot-pigeon. I may here 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 0. livia, no breeder will doubt that the occasional appearance of blue birds thus characterised Cuap. VI. THEIR REVERSION IN COLOUR. Dat is accounted for on the well-known principle of “ throwing back” or reversion. Why crossing should give so strong a tendency to reversion, we do not with certainty know; but abundant evidence of this fact will be given in the following chapters. It is probable that I might have bred even for a century pure black Barbs, Spots, Nuns, white Fantails, Trumpeters, &c., without obtaining a single blue or barred bird; yet by crossing these breeds I reared in the first and second generation, during the course of only three or four years, a considerable number of young birds, more or less plainly coloured blue, and with most of the characteristic marks. When black and white, or black and red birds, are crossed, it would appear that a slight tendency exists in both parents to produce blue offspring, and that this, when com- bined, overpowers the separate tendency in either parent to produce black, or white, or red offspring. If we reject the belief that all the races of the pigeon are the modified descendants of C. livia, and suppose that they are descended from several aboriginal stocks, then 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 Columbide presents these combined cha- racters; and it would not be possible to find any other instance of several species identical in plumage, yet as different in important points of structure as are Pouters, Fantails, Carriers, Tumblers, &c. Or lastly, we may assume that all the races, whether descended from ©. livia or from several aboriginal species, although they have keen bred with so much care and are so highly valued by fanciers, have all been crossed within a dozen or score of generations with CU. livia, and have thus acquired their tendency to produce | (4 DOMESTIC PIGEONS: Cuar. VI blae birds with the several characteristic marks. J 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 azd 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 improba- bility 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 confirne- Cusp. VI. THEIR REVERSION IN COLOUR. DMN ment 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 cha- racter; 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 unprobability 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, &c., 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 214 DOMESTIC PIGEONS: Cuap, VI. 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,°° 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. I+ 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, shonld we hesitate to believe in that greater amount of variation which is necessary for the production of the eleven chief races? It should be borne in mind that in two of the most strongly-marked races, namely, Carriers and Short-faced ‘Tumblers, the extreme forms can be connected with the parent-species by graduated differences not greater than those which may be observed between the dovecot- pigeons inhabiting different countries, or between the various kinds of toy-pigeons,—gradations which must certainly be attributed to variation. That circumstances have been eminently favourable for the modification of the pigeon through variation and selec- tion 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 s.c.;%! but Mr. Birch, of the British Museum, informs me that the pigeon appears in a bill of fare in the previous dynasty. Domestic pigeons are mentioned in Genesis, Leviticus, and Isaiah.*? In the time of the Romans, 30 It deserves notice, as bearing on predicament. This is the case,as Mr. 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 2re ix the same Blyth has remarked to me, with Treron, Palumbus, and Turtur. 31 ¢ Denkmiler,’ Abth. ii. Bl. 70. 32 The ‘ Dovecote,’ by the Rev. E. S. Dixon, 1851, pp. 11-13. Adolphe Pictet (in his ‘Les Origines Indo Cuar. VI. Pilea FORMATION OF RACES, as we hear from Pliny,?? immense prices were given for pigeons; ‘nay, they are come to this pass, that they can reckon up their pedigree and race.’ In India, about the year 1600, pigeons were much valued by Akber Khan: 20,000 birds were carried about with the court, and the merchants brought valuable collections. “The monarch of Jran 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.” 4 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 in- vocation, “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 citles are eager fanciers. Mr. Layard informs me Européennes, 1859, p. 399) states domestication of the pigeon in the that there are in the ancient Sanscrit East. jJanguage between 25 and 30 names for the pigeon, and other 15 or 16 Persian names ; none of these are com- mon to the European languages. This fact indicates the antiquity of the 33 Enolish translation, 1601, Book Xo Chips SEK VIN. 34“ Ayeen Akbery,’ translated ky F, Gladwin, 4to edit., vol. i. p. 270. 216 DOMESTIC PIGEONS: Cuar. VL 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 passion- ately devoted to the breeding of pigeons. Hear how en 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.” °° 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 hfe would be likely to cause variability. Pigeons have been domesticated for nearly 5U00 years, and have been kept in many places, so that the numbers reared under domestication must have been enor- 1ous: 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 facilty. Pigeons, differentiy from any other domesticated animal, can easily be mated for © J. M. Eaton, ‘Treatise on the Almond Tumbler,’ 1851; Preface, p. vi. Gear: Vs. HiSTORY OF THE PRINCIPAL RACES. 2A. life, and, though kept with other pigeons, rarely prove un- faithful 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.*® 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 ofany other domesticated animal. Someof the cases are inter- esting 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 charac- terised in 1735; and Laughers were apparently known in India before the year 16C0. Spots in 1676, and Nuns in the time of Aldrovandi, before 1600, were coloured exactly as they new 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,*’ 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 63 or 67 long “must be allowed to be avery good one.” Mr. Bult, the most 36 As in the following discussion I completed in the year 1858. often speak of the present time, I 37 “ Ornithologie,’ 1600, vol. ii. p should state that this chapter was 360. 218 DOMESTIC PIGEONS: Cuap. VIL. 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 73 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 Buii’s birds measured 73 inches in length. The extremely slight improvement in Pouters, except in the length of the leg, during the last 125 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* there was a change of fashion, stouter and more feathered legs being preferred to thin and nearly naked legs. Fantcils—The first notice of the existence of this breed is in India, before the year 1600, as given in the ‘Ayeen Akbery;** at this date, judging from Aldrovandi, the breed was unknown in Europe. In 1677 Willughby speaks of a Fantail with 26 tail-feathers; in 1735 Moore saw one with 36 feathers; and in 1824 MM. Boitard and Corbié assert that in France birds can easily be found with 42 tail- feathers. In England, the number of the tail-feathers is not at present somuch 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 intro- duction into Europe ; and some, said to have been brought from . Calcutta, which J 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 scnt 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 land cf pigeon, and the bill is 38 not in- variably 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 230 DOMESTIC PIGEONS. Cuap. VL 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, aud 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 ex- tinct. 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. Jn 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 ex- emplified 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 Cuap. VI. MANNER OF FORMATION OF RACES. 231 into slightly different sub-breeds, such as the common English Tumbler, the Dutch Roller, the Glasgow House- tumbler, and the Long-faced Beard Tumbler, &c.; and in the course of centuries, unless fashions greatly change, these sub- oreeds will diverge through the siow 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, tugether with the extinciion 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 cha- racters. 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 preferrmg 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 extinc- tion 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 inthe 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. Look- mg back to the time of Aldrovandi, we find that there existed in Europe, before the year 1600, four breeds which 2a2, DOMESTIC PIGEONS. Cuap. VL 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 recognized 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 domes- ticated 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 ex- tinction of a race within historical times is an event hkely to be noticed; but its gradual and scarcely sensible modifi- cation 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 conver- sion 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 srowth 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 organisa- tion, it is not surprising that dovecot-pigeons have remained Sep Ti. Cuar. VI. MANNER OF FORMATION OF RACES. Dae unaltered from time immemorial ; and that some toy-pigeons, which differ in little else besides colour from the dovecot- pigeon, have retained the same character for several centuries. for when one of these toy-pigeons had once become beautifully and symmetrically coloured,—when, for instance, a Spot had been produced with the crown of its head, its tail, and tail- coverts of a.uniform colour, the rest of the body being snow- white,—no alteration or improvement would be desired. On the other hand, it is not surprising that during this same interval of time our highly-bred pigeons have undergone an astonishing amount of change; for in regard to them there is no defined limit to the wish of the fancier, and there is no known limit to the variability of their characters. What is there to stop the fancier desiring to give to his Carrier a longer and longer beak, or to his Tumbler a shorter and shorter beak? nor has the extreme limit of variability in the beak, if there be any such limit, as yet been reached. Not- withstanding 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 Columbide, 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 tve, or in habits of life, as in roosting and building in trees. But the above objection shows how com- pletely the principle of selection has been misunderstood. It is not likely that characters selected by the caprice of man should resemble differences preserved under natural conditions either from being of direct service to each species, or from standing in correlation with other modified and serviceable structures. Until man selects birds differing in the relative length of the wing-feathers or toes, &c., no sensible change in these parts should be expected. Nor could man do anything 234 DOMESTIC PIGEONS. Cuap. VL unless these parts happened to vary under domestication :. I do not pcsitively 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 scutelle. 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 ereatly 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 de- scended from the Columba livia, including under this name certain wild races. But the differences between the latter throw no ight whatever on the characters which distinguish the domestic races. In each breed or sub-breed the individual birds are hore variable than birds in a state of nature; and occasionally they vary in a sudden and strongly-marked manner. ‘This plasticity of organization apparently results from changed conditions of life. Disuse has reduced certain parts of the body. Correlation of growth so ties the organisa- tion 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. Sclection = Cuap. VI. MANNER OF FORMATION OF RACES, 235 is followed methodically when the fancier tries to improve and modify a breed according to a prefixed standard of excel- lence; 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. 236 FOWLS. CHAPTER VII. FOWLS. BRIEF DESCRIPTIONS CF THE CHIEF BREEDS—-ARGUMENTS IN FAYOUR OF THEIR DESCENT FROM SEVERAL SPECIES—-ARGUMENTS IN FAVOUR OF ALL THE BREEDS HAVING DESCENDED FROM GALLUS BANKIVA—REYVERSION 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, VERTEBR#Z, 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 descrip- tion of them.! From what I have read and seen of specimens brought from several quarters of the world, I believe that most of the chief kinds have been imported into England, but many sub-breeds are probably still unknown here. The following discussion on the origin of the various breeds and on their characteristic differences does not pretend to com- pleteness, 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 subordinaticn 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; 1 I have drawn up this briefsynop- _ likewise assisted me in every possible sis from various sources, but chiefly way in obtaining for me information from information given me by Mr. and specimens. I must not let this Tegetmeier. This gentleman has opportunity pass without expressing kindly looked through this chapter; my cerdial thanks to Mr. B. P. Brent, and from his well-known knowledge, a well-known writer on poultry, for the statements here given may be continuous assistance and the gift of fully trusted. Mr. Tegetmeier has many specimens. Cuap. VIL. DESCRIPTION OF BREEDS. Zan bat I have great doubts whether this is a natural arrange- ment, 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 unsatis- factory. 1. GamME BrreEeD.—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, &c., with their legs of various colours. 2. Matay Bree. ’ 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, DorKine BreEp.—Size great; body square, compact; feet with an additional toe; comb well developed, but varies much in form; wattles well developed; colour of plumage various. Skull remarkably broad between the orbits. Of English origin. The white Dorking may be considered as a distinct sub-breed, being a less massive bird. 5. SPANISH Brexp (fig 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 238 FOWLS. Cuar. VIL 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. Fig. 30.—Spanish Fowl. 6, HampurGH Breen (fig. 31).—Size moderate; comb flat, pro- duced backwards, covered with numerous small points; wattle of moderate dimensions; ear lobe white; legs blueish, thin. Do not incubate. Skull, with the tips of the ascending branches of the premaxillary and with the nasal bones standing a little separate from each other; anterior margin of the frontal bones less depressed than usual. Cuap. VIL DESCRIPTION OF BREEDS. 239 There are two sub-breeds; the spangled Hamburgh, of English origin, with the tips of the feathers marked with a dark spot; and the penciled 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-yarieties. Black Hamburghs have been produced by a cross with the Spanish breed. 7. CrestED og Potisa Breep (fig. 32).—Head with a large, rounded crest of feathers, supported on a hemispherical protuberance Sh \ AA Fig. 31—Hamburgh Fowl. 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. ‘lhe orifice of the nostrils raised and crescentic. Beak short. Comb absent, or small and of cre- scentic 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. 240 FOWLS. Cuar. VIL 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 sameremarkable peculiarities of structure as in the true Polish fowl. Sub-breed (a2) Sultans—A Turkish breed, resembling white Polish fowls with a large crest and beard with short and well- Fig. 32.—Polish Fowl. feathered legs. The tail is furnished with additional sickle feathers Do not incubate.* Sub-breed (0) Pturmigans.—An inferior breed closely allied to 2 The best account of Sultans is by kindness the examination of some Miss Watts in ‘The Poultry Yard,’ specimens of this breed. 1856, p. 79. I owe to Mr. Brent’s Cuap. VIL DESCRIPTION OF BREEDS. 241 the last, white, rather small, legs much feathered, with the crest pointed ; comb small, cupped; watties 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. J have seen an allied white, tailless breed from Turkey. Sub-breed (7) Créve-ceur.—A French breed of large size, barely capable of flight, with short black legs, head crested, comb produced into two points or horus, sometimes a little branched like the horns of astag; both beard and wattles present. Eggs large. Disposition quiet.? Sub-breed (e) //orned fowl.—With a small crest; comb preduced into two great points, supported on two bony protuberances. Sub-breed (f) Houwdan.—A French breed; of moderate size, short- legged with five toes, well developed; plumage invariably mottled with black, white,and straw-yetlow; head furnished with a crest, on a triple comb placed transversely ; both wattles and beard present.* Sub-breed (7) 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 BreEp.—Originally from Japan,> 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. RumpiEss Fowus.—These are so variable in character® that they hardly deserve to be called a breed. Anyone who will examine the caudal vertebree will see how monstrous the breed is. 10. CREEPERS ok JUMPERS.—These are characterized 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. li. Frizzuep or Carrre Fowis.—Not uncommon in India, with the feathers curling backwards, and with the primary feathers of the wing and tail imperfect; periosteum of bones black. 3 A good description, with figures, is given of this snb-breed in the ‘Journal of Horticulture,’ June 10th, 1862, p. 206. * A description, with figures, is given of this breed in ‘Journal of Horticulture,’ June 3rd, 1862, p. 185. Some writers describe the comb as two-horned. 5 Mr. Crawfurd, ‘ Descript. Dict. of the Indian Islands,’ p. 113. Ban- tams are mentioned in an ancient native Japanese Encyclopedia, as I am informed by Mr. Birch of the British Museum. § ‘Ornamental and Demestie Pou try,’ 1848. Fee aa 9A? + FOWLS. Cuar. VIL 12. Smx Fowts.—Feathers silky, with the primary wing and tail-feathers impericct; 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. Soory Fowris.—An Indian breed, having the peculiar appear- ance 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 neariy 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. 8. Dixon’ argues strongly on this side of the question; and one fancier even denounces the opposite con- clusion 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 ereat difference between the chief kinds, and they ask with force, can differences in climate, food, or treatment have pro- duced 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 correla- tion of growth—of the long-continued use and disuse of paris, 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 7 ‘Ornamental and Domestic Poultry,’ 1848. Cuap. VIL THEIR PARENTAGE. 243 though they are well aware that their birds differ individually and that by selecting the best birds for a few generations they, ean improve their stocks. An amateur writes* 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 improba- bility 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 siven showing not only that careful breeding, but that actual selection was practised during ancient periods, and by barely civilized 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 recom- mends as the best, those sorts that have five toes and white ears.’° In the fifteenth century several breeds were known and described in Europe; and in China, at nearly the same period, seven kinds were named. A more striking case is that at present, in one of the Philippine Islands, the semi-barbarous inhabitants have distinct native names for no less than nine sub-breeds of the Game fowl.!® Azara,!! 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. Wiil it then be pretended that those persons who, 8 Ferguson’s ‘ Illustrated Series of of the Domesticated Animals to Civili- Rare and Prize Poultry,’ 1854, p. vi. zation,’ separately printed, p. 6; first Preface. read before the Brit. Assoc. at Oxford, ® Rev. E. S. Dixon, in his ‘Orna- 1860. mental Poultry,’ p. 203, gives an ac- 1 “Quadrupédes du Paraguay,’ tem eount of Columella’s work. ii. p. 324, 10 Mr. Crawfura ‘On the Relation D44 FOWLS. Cuap, VIL in ancient times and in semi-civilized countries took pains to keep the breeds distinct, and who therefore valued them, would not occasionally have destroyed inferior birds and occa- sionally 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 medify an old breed according to some ideal standard of excellence. He who cared for poultry would merely wish to obtain, and afterwards te 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 lus 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. IH, 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 cha- racters, though at first only slight in degree; from the effects of the use and the disuse of parts; possibly from the direct effects of changed climate and food; from correlation of Cuap. VIL. THEIR PARENTAGE. 245 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 unconsciovs selection carried on during many generations, there is no insuperable difficulty, to the best of my judgment, in believ- ing 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,” 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 lamine, trans- versely banded with three colours; and I have met no authentic account of any such character having been observed in any domestic breed.“ 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 “ 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 12 ¢Proc. Zoolog. Soc.’ 1832, p. red game-hen, and they exhibited the 151, true character of those of G. sonne- 13 These feathers have been de- ratii, except that the horny lamine scribed by Dr. W. Marshall,‘Der Zoolog. | were much smaller. Garten,’ April 1874, p. 124. I ex- 14 See also an excellent letter on amined the feathers of some hybrids the Poultry of India, by Mr. Biyth, raised in the Zoological Gardens in ‘Gardiner’s Chronicle,’ 1851, p. between the male G. sonneratii and a 619. 246 Cuap. VIL. FOWLS. more or less infertile. Nearly similar experiments have recently been tried on a great scale in the Zoological Gardens with almost the same result.’ 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 ‘nfer se. From these facts, and from the above-mentioned strong!y-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. stanleyiz ; this species approaches so closely (except in the colouring of the comb) to the domestic fowl, that Messrs. Layard and Kellaert '® 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. stanleyiz. This species, then, may in all pro- bability 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 Tam informed by Mr. Crawfurd,” hybrids are commonly raised between the male G. variuvs 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 thonght to be specifically distinct, and were named G. eneus. Mr. Blyth and others believe that the G. temminckii'*® (of which the history isnot 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. J may just allude to the G. giyanteus, so often referred to in works on poultry as a wild species ; but Marsden the first describer, speaks of it as a tame breed; and the specimen in the British Museum evidently has the aspect of a domestic variety. 55 Mr. S. J. Salter, in ‘ Natura! History Review,’ April 1863, p. 276. 16 See also Mr. Layard’s paper in ‘Annals and Mag. of Nat. History,’ 2nd series, vol. xiv. p. 62. 17 See also Mr. Crawfurd’s ‘ Descrip- tive Dict. of the Indian Islands,’ 1856, p. 113. 18 Described by Mr. G. R. Gray, ‘Proc. Zoolog. Soc,’ 1849, p. 62. 18 The passage from Marsden 1s given by Mr. Dixon in his ‘ Poultry Beok,’ p. 176. No ornithologist now ranks this bird as a distipet species. Cae. VIL. THEIR PARENTAGE. 247 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 archipelego 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” 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, &c. 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.*!_ Mr. Brent, however, has remarked that these black feathers remain in the wild bird after the development oi 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 20 “Coup-d’eil général sur |'Inde 21 Mr. Blyth, in ‘ Annals and Mag. Archipélagique,’ tom. iii. (184), p. of Nat. Hist.,’ 2nd ser., vol. i. (1848), 177; sce also Mr. Blyth in ‘Indian p. 455 Sporting Review,’ vel. ii. p. 5, 1856, 248 FOWLS. Cuar. VII 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 1o maturity ; but as he remarks, “no wild gallinaceous bird thrives well at first on hard grain.” Mr. Blythalso 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.~ Sir Walter Elliot informs me that the hen of a native domestic breed of Pegu is undistinguish- able 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. Mr. Crawfurd remarks that from etymology it might be argued that the fowl was first domesticated by the Malays and Javanese.** 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,?° who are familiar with G. bankiva, believe that it is the parent of most or all our 22 Crawfurd, ‘ Dese. Dict. of Indian Islands,’ 1856, p. 112. Journ. of Lit. and Science.’ vol. xxii. p. 2, speaking of G. bankiva, says, “ un- 23 In Burmah, as I hear from Mr. Blyth, the wild and tame poultry con- stantly cross together, and irregular transitional forms may be seea. 24 Ibid. p. 113. %® Mr. Jerdon, in the ‘ Madras questionably the origin of most of the varieties of our common fowls.” For Mr. Blyth, see his excellent article in ‘Gardener’s Chron.,’ 1851, p. 619; ana in ‘ Annals and Mag. of Nat. Hist.,’ vol. xx., 1847, p. 388. Cuap. VIL, 249 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, how- ever, 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 Fast. 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,”® generally have a restricted range: we sce this well illustrated in India, where the genus Gallus in- habits 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 THEIR PARENTAGE. unknown species of the genus. that Gallas should 9 < 6 ¢Gardiner’s Chronicle’ 1851, p. 619. 27 [| have consulted an eminent authority, Mr. Sclater, on this subject, and he thinks that I have not expressed myself too strongly. Iam 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 probabiy 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. Alien (‘Narrative of Niger Expedition,’ 1848, vol. ii. p. 42) de- scribes wild fowls on Ilha dos Rollas, an island near St. Thomas’s on the west coast of Africa; the natives in- 12 it is, also, as improbable inhabit South America2’ as that a formed him that they had escaped from a vessel wrecked there many years ago; they were extremely wild and had “a ery quite different to that ot the domestic fowl,” and their ap- pearance was somewhat changed. Hence it is not a little doubtful, not- withstanding 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, p. 62), and in the Ladrones (Anson’s Voyage). Those found in the Pellew Islands Crawfurd) are believed to be feral 250 FOWLS. . Cuap. VII. humming-bird should be found in the Old World. From the vharacter 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. Elyth and Crawtfurd, who have attended to this subject, doubt whether Gallus ever existed in a wild state even as far west as Persia. Although the earhest 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 pro- bably have become known. Mr. Birch, of the British Museum, has translated for me passages from a Chinese Encyclopedia 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 im- portance 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 and lastly, it is asserted that they but whether this is correct I know have become feral in New Zeaiapd, not. Cuar. VI. REVERSION AND ANALOGOUS REVERSION. 251 crossed with each other, or when crossed, with the exception of G. bankiva, with the domestic fowl, produce infertile hybrids. Finally, we have not such good evidence with fowls as with pigeons, of all the breeds having descended from a single primitive stock. In both cases the argument of fertility must go for something; in both we have the im- probability of man having succeeded in ancient times in thoroughly domesticating several supposed species,—most of these supposed species being extremely abnormal as compared with their natural allies,—all being now either unknown or extinct, though the parent-form of 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 hblack- 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,”?’ when 8 Mr. Hewitt, in ‘The Poultry Book,’ by W. B. Tegetmeier, 1866, p. 248, p25) ae FOWLS._ Ouav. VIE, two or three years old, not uncommonly become ruddy; these latter Bantams occisionally “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 Ham- burgh 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 hving 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 egos 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 hroadly bordered with very pale yellowish-brown ; so that in Cuap. VII. REVERSION AND ANALOGOUS REVERSION. 293 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° 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 pale orange- coloured hackles, chiefly on the loins, and the other an abundance of fine orange-red hackles on the neck, loins, and upper wing-coverts. Here again we have a more decided, though partial, reversion to the colours of G. bankiva. This second cock was in fact coloured like an inferior “ pile Game cock ;”—now this sub-breed can be produced, as I am in- formed 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 29 ‘Journal cf Horticulture,’ Jan. 14th, 1862, p. 325. 254 FOWLS. Cuar. VII. old, one of them assumed yellowish-white hackles, and thus resembled in a considerable degree the cross from the Ham- burgh 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 ex- traordinary 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 hkewise 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 suv 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 shightly brown or sooty. Several hens, which were for a long time snow- Cuar. VII. REVERSION AND ANALOGOUS REVERSION. 255 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 longi- tudinal stripes so general with gallinaceous birds; but this did not occur in @ 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 ethan 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*? 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, 39 “Die Hithner- und Pfaucuzucht,’ W.B. Tegetmeier, 1866, p. 222. I am Ulm, 1827,s. 17. For Mr. Hewitt’s indebted to Mr. Orton for a letter on statement with respect to the white the same subject. Silk fowl, see the ‘ Pou’try Book,’ by Z56 FOWLS. Cuap. VII ith the mottling partially and obscurely arranged in trans- verse 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. Teget- meier 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. bankira ; nor does it often follow, as I hear from Mr. Teget- meier, from crossing distinct breeds; but it 1s 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 Cuap. VII- REVERSION AND ANALOGOUS VARIATION. 257 ou their wings.*! The editor of the ‘Poultry Chronicle’ # remarks that all the breeds which properly have red ear- lappets occasionally produce birds with white ear-lappets. 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, wil 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 31 Dixon, ‘Ornamental and Do- ‘Prize Poultry,’ p. 260. mestic Poultry,’ pp. 253, 324, 335. 32 ¢ Poultry Chronicle,’ vol. ii. For game fowls, sce Ferguson on 71. 258 FOWLS. Cuar. VIL. 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.—Riitimeyer found no remains of the fowl in the ancient Swiss lake-dwellings ; but, according to Jeitteles,*? such have certainly since been found associated with extinct animals and prehistoric remains. It is, there- fore 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 sB.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 33 “Die vorgeschichtlichen Alter- aversion. The natives of the Pellew thiimer,’ IJ. Theil, 1872, p. 5. Dr. Islands would not eat the fowl, nor will Pickering, in his ‘Races of Man,’ the Indians in of S. 1850, p. 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 tev. S. Erhardt that on the east coast of Africa, from 4° to 6° south of the equator, most of the pagan tribes at the present day hold the fowl ir some parts America. For the ancient history of the fowl, see also Volz, ‘ Beitrage zur Culturgeschichte,’ 1852, s. 77; and Isid. Geoffroy St.-Hilaire, ‘Hist. Nat Gén.,’ tom. iii. p. 61. Mr. Crawfurd has given an admirable history of the fowl in his paper ‘On the Relation of Domesticated Animals to Civilisa- tion,’ 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 lnsti- tuses of Manu. Cuap. VII. THEIR HISTORY. 259 by Julius Cesar. 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 Encyclopedia, 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 re- ferred to in an old Japanese Encyclopedia, as I am informed by Mr. Birch. In the Chinese Encyclopzedia 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 de- scribes 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 wonld 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 *‘ considers 4 “Ornamental and Domestic Poul- 312. For Golden Hamburghs, sce try, 1847, p. 185; for passages Alom’s ‘Natural History of Birds, translated from Columella, sce p. 3 vols., with plates 1731-38. 260 FOWLS. CuHap. Vil. 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 belheving that all the breeds are descended frem 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. Eqqs.—Myr. Dixon remarks® that “to every hen belongs an indiy idual 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 acquain- tance.” 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 re- cognised. The eggs of differently sized breeds naturally differ much in size; but apparently, not always in strict relation to the size ot 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 ;% 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 35 ‘Ornamental and Demestic Poul- informed, cannot generally be trusted try,’ p. 152. He gives, however, figures and mneb 88 Ferguson on ‘Rare Prize Pow'- information on eggs. See pp. 34 and try, p. 297 This writer, I am 235 on the eggs of the Game fowl 261 Cuav. VII. DIFFERENCES BETWEEN THE BREEDS. vary considerably in character; for instance, Mr. Ballance states 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.*® Tbe colour diltfers considerably,—the Cochins laying buff-coloured eggs; the Malays a paler variable butf; 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. Iam 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. 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 37, See ‘Poultry Book,’ by Mr. Tegetmeier, 1866, pp. 81 and 78. 38 ‘The Cottage Gardener,’ Oct. 1855, p. 13. On the thinness of the eggs of Game-fowls, see Mowbray on Poultry, 7th edit., p. 13. 38 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, sce Tegetmeier’s * Poultry Book,’ 1866, p, apie) EP 262 FOWLS. Cuar. VIL 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 longitudiual 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 atime 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 lam 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 tne back. Gold- spangled Polish chickens (Tegetmeier) are of a warm russet brown ; and silver-spangled Polish chickens are grey, sometimes Sees with dashes of ochre on the head, wings, and breast. Cuckoo an 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;*° but the crest, which is supported on the protuberance, is at first feebly developed, nor does it attain 40 As I hear from Mr. Tegetmeier ; crest, see ‘Poultry Chronicie, vel, see also ‘Proc. Zoolog. Soc.’ 1656, p. ii. p. 132. 366. On the late development of the Cuap. VIL SEXUAL DIFFERENCES, 263 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 erow 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.*! Cochins are characterised by a small tail, and in the young cocks the tail is developed at an unusually late period. Game fowls are notorious for their pugnacity ; and the young cocks crow, clap their little wings, and fight obstinately with each other, even whilst under their mother’s care.** ‘I have often had,” says one author,** “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 gallina- ceous 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 unnaturaliy transferred to the young male chickens. In the same manner, it is probable that the extraordinary de- velopment of the comb in the Spanish cock has been un- intentionally 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; 41 On these points, see ‘Poultry tic Poultry,’ p. 273. Chronicle,’ vol. iii. p. 166; and Teget- 43 Ferguson on Rare and Prize meier’s ‘ Poultry Book,’ 1866, pp. 105 Poultry, p. 261. and 121. ** Mowbray on Poultry, 7th edit. #2 Dixon, ‘Ornamental and Domes- 1834, p. 15. 264 FOWLS. Cuar. VIL 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, differs much in colour. In our domestic breeds the difference 1s 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 differencein 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.*? A similar relation is also evident when we com- pare the several varieties of Cochins. In the two sexes of gold and silver-spangled and of buff Polish fowls, there is puch 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 lkewise a-con- siderable degree of similarity between the two sexes. In pencilled Hamburghs, on the other hand, there is much dis- similarity ; 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 resemblage in plumage to the females, are often called hennies. There is much diversity of opinion whether these males arein any degree sterile; that they some- 45 See the full description of the meier’s ‘ Poultry Book,’ 1866, p. 131 varieties of the Game-breed, in Teget- For Cuckoo Dorkings, p. 97. Car. VII. SEXUAL DIFFERENCES. 265 times are partially sterile seems clear,*® 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-lke 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.*’ 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** 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.” *° 46 Mr. Hewitt in Tegetmeier’s a-dozen cocks thus sacrificed. ‘Poultry Book,’ 1866, pp. 246 and 48 «Proceedings of Zoolog. Soc.’ 156. For hen-tailed game-cocks, see March, 1861, p. 102. The engraving p- 131. of the hen-tailed cock just alluded te 47 «The Field,’ April 20th, 1861. was exhibited before the Society. the writer says he has seen halt- 49 «The Field,’ April 20th, 1861 266 - FOWLS. Cuap, VIL 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 seer, as far as the colour of plumage is con- cerned, and so it is with the other secondary sexual characters. Firstly, the comb differs much in the various breeds,*° and its form is eminently characteristic of each kind, with the exception of the Dorkings, in which the form has not been as yet 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 ina 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.” *! In some breeds the comb is duuble, and when‘the two ends are cemented together it forms a “ cup-comb;” in the “rose comb” it is depressed, covered with small pro- jections, and produced backwards ; in the horned and eréve- ceur 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 deve- loped, 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 ** that they are in consequence hable 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 3” Jam much indebted to Mr. Brent 51 The ‘ Poultry Book,’ by Teget- for an account, with sketches, of all meier, 1866, p. 234. the variations of the comb known to 52 «Die Hiihoer- und Pfanenzucht, him, and likewise with respect to the 1827, s. 11. tail as presently to be given. Cuar. VIE. SEXUAL DIFFERENCES. 267 are short and sti in Malays, and absent in Hennies. As in some orders male birds display extraordinarily-shaned feathers, such as naked shafts with discs at the end, &c., the following case may be worth giving. In the wild Gallus bankiva and in our domestic fowls, the barbs which arise from each side of the extremities of the hackles are naked or not clothed with barbules, so that they resemble bristles; but Mr. Brent sent me some scapular hackles from a young Birchen Duckwing Game cock, in which the naked barbs became densely re- clothed with barbules towards their tips; so that these tips, which were dark coloured with a metallic lustre, were sepa- rated from the lower parts by a symmetrically-shaped trans- parent zone formed of the naked portions of the barbs. Hence the coloured tips appeared hke 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 Games; but droop much in Malays and insome 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 tas 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 ; ** 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 Ency- clopeedia. Their occurrence may be considered as a case of 53 “Poultry Chronicle,’ vol. i. p. position of the spurs in Dorkings, see 695. Mr. Brent has informed me of ‘Cottage Gardener,’ Sept. 18th, 1860, the same tact. With respect to the p. 380. 268. FOWLS. Cuap. VIL 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 Gallinacex, 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 en- tirely takes the place of the crest.”°* 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 lke- wise been largely transferred to the female ;°* and she some- times even possesses the eminently masculine character of spurs. Many cases are on record of fertile hens being furnished 54 Dixon, ‘Ornamental and Domes- _ bative, that it is now generally the tic Poultry,’ p. 320. practice to exhibit each hen in @ 55 Mr. Tegetmeier informs me that _—_ separate pen. Game hens have been foun? so com- Guar, VIL. EXTERNAL DIFFERENCES. 269 with spurs; and in Germany, according to Bechstein,*® the spurs in the Silk hen are sometimes very long. He mentions aiso another breed similarly characterised, in which the hens are excellent layers, but are apt to disturb and break their egos owing to their spurs. Mr. Layard*’ 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 aid 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 ereat 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,°®* 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 care- fully 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. 56 ¢ Naturgeschichte Deutschlands,’ Band iii. (1723), s. 359, 407. 57 On the Ornithology of Ceylon in ‘Annals and Mag. of Nat. History.’ 2nd series, vol. xiv. (1854), p. 63. °8 “Handbuch der yergleich. Ana- tomie,’ 1805, p. 85, note. Mr. Teget- meier, who gives in ‘Proc. Zoolog. Soc.,’ Nov. 25th, 1856, a very interest- ing account of the skulls of Polish fowls, not knowing of Bechstein’s ac- count, has disputed the accuracy of Blumenbach’s statement. For Bech- stein, see ‘ Naturgeschichte Deutsch- lands,’ Band iii. (1793), s. 399, note. 1 may add that at the first exhibition of Poultry at the Zoological Gardens, in May, 1845, I saw some fowls, callnd Friezland fowls,of which the hens were crested, and the cocks furnished with a comb. J ee . = R 270 FOWLS. Cuapr. VII. 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 wiih the Sexes, between the Breeds and between inaividual Birds. The size of the body differs greatly. Mr. Tegetmeier has knowu 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 Minoreas.*® A good observer® 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, 5° *Cottage Gardener,’ Jan. 3rd, before the Dublin Nat. Hist. Soc., 1860, p. 218. quoted in ‘Cottage Gardener,’ 1554, 8® Mr. Williams, in a paper read_ p. 161. Cuap. VII. EXTERNAL DIFFERENCES. Del as M. Godron © remarks the three principal types of skin in man- kind. 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 cres- eentic outline. The primary wing-feathers are short in Cochins; in % male, which must have been more than twice as heavy as G. oankiva, these featherg 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 number9; 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 callinaceous species the relative length and number of the main wing and tail-feathers are extremely constant. The tail differs much in erestness 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.°? An eminent physiologist * has recently spoken of this breed as a distinct species ; had he examined the deformed state of the os coccyx he would never have come to this conclusion ; he was probably misled by the statement, which may be found in some works, that tailless fowls are wild in Ceylon; but this statement, as I have been assured by Mr. Layard and Dr. Kellaert who have so closely studied the birds of Ceylon, is utterly false. The tarsi vary considerably in length, being relatively to the $1 cleaned and viewed from above, presented the appearance of an open basin. ‘The change in the whole internal form of the skull is sur- prisingly great. The brain is modified in a corresponding manner, as is shown in the two longitudinal sections, which deserve attentive consideration. ‘The upper and antericr 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, asI 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 (fig. 35), a perfect gra- dation 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® 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.” 6° “«Naturgeschichte Deutschlands,? have received communications to a Band iii. (1793), s. 400. similar effect from Messrs, Brent and 7@ The ‘Field, May 11th, 1861. I Tegetmeier. Cuap. VIL. OSTEOLOGICAL DIFFERENCES. PT 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 ex- tremities : 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 ASS SS SSS Wigs CRG —) BECO TMA 22 y Saris © )) 2 Fig. 35.—Longitudinal sections of Skull, of natural size, viewed laterally. A. Polish Cock. Bb. C in. Coel: ai : Z ge Cock, selected for comparison with the above from being of nearly the same 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. Hach nasal bone properly sends out in front two long 278 FOWLS. Cuap. VII. 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 BEET a crescentic outline. I must still say a few words on some of the foreign Crested breeds. The skull of a crested, rnmpless, white Turkish fowl was very slightly protuberant, and but little perforated; the ascending branches of the premaxillary was well developed. In another Turkish breed, called Ghoondooks, the skull was considerably protu- berant and perforated; the ascending branches of the premaxillary were so much aborted that they projected only +,th of aninch; 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 pro- tuberance 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, Cuap. VIL OSTEOLUGICAL DIFFERENCES. 279 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 hat 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 per- forated 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. Tegetineier informs me that if chickens lately hatched be selected with a large bony protuberance. when adult aA Ss = = ZA ~ S N dd gd idseii LZ Fig. 36.—Skull of Horned Fowl, of natural size, viewed from above, a little obliquely. (In the pussession of Tegetmeier.) they will have a large crest. There can be no doubf that in former times the breeder of Polish fowls attended solely to the crest, and not tothe skull; nevertheless, by increasing the crest, in which he has been wonderfully successful, he has unintentionall y made the skull protuberant to an astonishing degree; and through correlation of growth, he has at the same time affected the form and relative con- nexion 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 AD of the whole skull together with the shape of the rain. Vertehre.—In G. bankiva there are fourteen cervical, seven dorsal with ribs, apparently fifteen lumbar and sacral, and six caudal 280 FOWLS. Cuap. VIL vertebrae ;7 but the lumbar and sacra! are so much anchylosed that Iam not sure of their number, and this makes the comparison of the total number of vertebrze In the several breeds difficult. I have spoken of six caudal vertebra, because the basal one is almost completely anchylosed with the pelvis; but if we consider the number as seven, the caudal vertebree agree in all the skeletons. The cervical vertebre 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, forall the cervical vertebree 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 vertebre,’* we need feel no surprise at the number of the cervical vertebre in the fowl being, as it appears, variable. There are seven dorsal vertebrz 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 vertebrze were free. In two skeletons, the fifth dorsal was free; generally the sixth is free (as in G. banxkiva), 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 vertebrz 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 71 Tt appears that I have not caudal vertebre in this genus. But correctly designated theseveral groups I have used the same terms in al! the of vertebrae, for a great authority, following descriptions. Mr. W. K. Parker (‘ Transact. Zoolog. 2 Macgillivray, ‘ British Birds,’ vol Soc.,? -ve.. v. p. 198), specifies 16 i. p. 25. eervical, 4 dorsal, 15 lumbar, and 6 Cuap. VIL OSTEOLOGICAL DIFFERENCES. gs 281 pair in other skeletons; in this Game cock, as far as could be judged from the appearance of the lumbar vertebree, 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 vertebre; but they certainly do not correspond in shape or number in the several skeletons. The caudal vertebra 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 vertebre were a little elongated. In three rumpless fowls the caudal vertebre were few in number, and anchylosed together into a misformed mass. In the individual vertebre the differences in structure are very slight. Inthe 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 are 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 hemal descending processes being united to the body of the vertebra _. ; i by asort of buttress. This structure may H Gee ae ae be observed in Cochins, Polish,some Ham- A. Wild Gallus bankiva. B. burghs, and probably other breeds; but Cochin Cock. 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 vertebree; 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. 282 FOWLS. Cuap, VIL The end of the pubic bone presents the greatest difference; being hardly enlarged in the Bankiva; considerably and gradually enlarged in Cochins, and in a lesser degree in some other breeds; and abruptly enlarged in Bantams. In one Bantam this bone extended very little beyond the extremity of the ischium. The whole pelvis in this latter bird differed widely in its proportions, being far broader proportionally to its length than in Bankiva, Sternum.—This bone is generally so much deformed that it is scarcely possible to compare its shape strictly in the several breeds. The form of the triangular ex- tremity of the lateral processes differs considerably, being either almost equilateral or much elon- gated. The front margin of the crest is more or less perpendicular end 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 Ban- kiva, and rounded in the Spanish breed. The jurculum 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 Ban- ee Ot Se mae kiva, much broader in the middle Fig, 2 Estremity of the, Fura of in the Polish fowl, and abruptly Gallus bankiva. B. Spangled Polish narrowed towards the apex In Eur C. Spanish Fowl. D. Dorking the two Sultan fowls. I carefully compared each separate bone of the lez 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. Cxap. VIL OSTEOLOGICAL DIFFERENCES. 283 Finally, I have not examined a sufficient number of skele- tons to say whether any of the foregoing differences, except in the skull, are characteristic of the several breeds. Appa- rently 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 wellas 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 te unnatural conditions of life, and their whole organization 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 284 - FOWLS. Cuar, VIL 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 pro- tuberances 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 1s 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 Paris. 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 jength 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 Cuar. VIL THE EFFECTS OF DISUSE. 289 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 pevine accidentally had wings of slightly greater length than usual; that the measurements are not worth giving. But it ee notice that the Silk and Frizzled fowls, which are quite incapable of flight, had their wings /ess 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 ofnutriment. ‘The wing-bones, however, in both these breeds, are found to be slightly reduced in length when judged by the standard of the length of the sternum or head, relatively to these same parts in G. bankiva., The actual weight of the main bones of the leg and wing in twelve breeds is given in the two first columns in the following table. The calculated weight of the wing-bones relatively to the 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.’? TABue I, Actual Weight of Wing- Weight| Actual | bones relatively tc Names of Breeds. of Weight of | the Leg-bones in Femur | Humerus | comparison with and |and Ulna.| these same bones Tibia. in G. bankiva | Grains. |* Grains. Gallus bankiva .. .. wild male | 86 54 100 Cochin Pe Peet be MNBLE 311 162 83 | Dorking... .. male 597 248 70 Sp: ish (Minorea) .. male 386 183 79 | Gold-Spangled Polish male 145 7d Game, black-breasted male 143 Oe NM Aktye eae eee) ce) gue temale ; 116 80 Sulla. 20. 22 9 male 94 79 Indian Frizzled .. .. male 88 67 Burmese Jumper’... female 36 108 Hamburgh (pencilled) male ; 104 106 - Hamburgh (pencilled) female U0, 108 Silk (black-boned) .. female a7 103 1 2 3 4 5) 6 7 8 9 73 It may be well to explain how leg-hones are to the wing-bones as the calculation has been made for the 86: 54, or as (neglecting decimals) third column. In G. bankiva the 100 : 62;—in Cochinsas 311 : 162, or > es ae 286 FOWLS. Cua. VII. 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 earried to the greatest extent, namely, to thirty-three per cent. of their proper proportional weight. Inthe 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 the foregoing table 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.“ Jam 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 lb. 2 0z., 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 so on for the remainder of the third column in the table. 74 Mr. Blyth (in ‘ Annals and Mag. as 100 : 52 s—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, &c. And now taking 10), 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 of Nat. Hist.,’ 2nd series, vol. i, 1848, p. 456) gives 33 lb. as the weight of a full-grown male G. bankiva; but from what I have seen of the skins and skeletons of various breeds, I cannot believe that my two specimens | ot G. bankiva could have weighed zo much. Onap. VII. THE EFFECTS OF DISUSE. 287 no certain conclusions could be reached. For instance, the legs of the above Dorking cock were nearly three-quarters of an inch too short relatively to the length of the sternum, and more than three- quarters of an inch too long relatively to the length of the skull, in comparison with these same parts in G. bankiva. In the following Table II. in the two first columns we see in inches and decimals the length of the sternum, and the extreme depth of its crest to which the pectoral muscles are attached. In the third 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.”® Tas.eE II. } | | Depth of Crest Depth of | relatively to the peer Crest length of the q of Sternum, in Steruum. | sternum. comparison with G. bankiva. Names of breeds. | Gallus bankiva .. .. male Wochitt. t2.° hice. Me. ale Dorkine 2: 2-85. - cara s.) Take RRs ccm et vee oe Tae iisian eres Meet rs =e amale Game yeiee ccc t tz, Female Ways eee pt eka, Lelale mais ce es ee Male Hrizzled hen 3° tea.) male Burmese Jumper .. .. female CHamburchs) 2 -.... 2... male Hamburch~ <2 =.. ~... female Silk fowl Vapi nes Lemule 1 2 3 4 ) 6 7 8 9 He OT 0 HE HE Ot Or OT OD OD Ot t a i ee ee 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 com- parison 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 75 The third column is calculated on the same principle as explained in the previous foot-note, p. 285. 288 FOWLS. Cuap. VII. 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 deli-_ cate ; 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 lable 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. but Bory de Saint-Vincent °° saw at Madrid gold-fish furnished with a dorsal fin anda triple tail. One variety is characterised by a hump on its back near the head; and the Rev. L. Jenyns°? has described a most singular variety, imported from China, almost globular in form like a Diodon, with “ the fleshy part of the tail as if entirely cut away? the caudal fin being set on a little behind the dorsal and immediately above the anal.” In this fish the anal and caudal fins were double; the anal fin being attached to the body in a vertical line: the eyes also were enormously large and protuberant. Hive-BeErEs. Bers 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 54 ¢Pyoc. Zoolog. Soc.,’ May 25th, 57 ¢Qbservations in Nat. Hist.,’ 1812. 1846, p.211. Dr. Gray has described, °5 Yarrell’s ‘ British Fishes,’ vol. i. in ‘Annals and Mag. of Nat. Hist.,’ . 319. 1860, p. 151, a nearly similar variety, 36 + Dict. Class. d’Hist. Nat.,’ tom. but destitute of a dorsal fin. Vv. p. 276, 314 HIVE-BEES. Cuap. VITT. 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®® says that they are generally larger in the south than in other parts of France ; it has also been asserted that the little brown hees 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®? 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,®° that he has several times seen queen bees of the common kind annu- lated 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 differ- ence in the queens or workers of the same hive. The great apiarian, Dzierzon, In answer to my queries on this subject, says, that in Germany bees of some stocks are decidedly dark, whilst others are remarkable for their yellow colour. Bees also seem to differ in habits in different districts, for Dzierzon adds, “If many stocks with their offspring are more inclined to swarm, whilst others are richer in honey, so that some bee-keepers even distinguish between swarming and honey-gathering bees, this is a habit which has become second nature, caused by the customary mode of keeping the bees 58 “De VEspéce,? 1859, p. 459. implicitly trusted; see ‘Journal of With respect to the bees of Burgundy, Horticulture,’ July 14th, 1863, p. 39. see M. Gérard, art. ‘ Espéce,’ in ‘ Dict. Univers. d’Hist. Nat.’ 3° See a discussion on this subject, in answer to a question of mine, in ‘Journal of Horticulture,’ 1862, pp. 225-242; also Mr. Bevan Fox, ia litto, 1862, p. 284 8° This excellent observer may be 1 “Journal of Horticulture,’ Sept. 9th, 1862, p. 4633 see also Herr Kleine on same subject (Nov. 11th, p. 643), who sums up, that, though there is some variability in colour, no constant or perceptible differences can be detected in the bees of Germany. Cuap. VIIL HIVE-BEES. 315 and the pasturage of the district. For example, what a difference in this respect one may perceive to exist between the bees of the Liineburg heath and those of this country !” aa “ 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 o! dead bees from Jamaica, where they have long been natural- ised, 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 differ- ence. 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 hee.*? 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 domestica- tion. The Egyptian and some other bees are hkewise ranked by Dr. Gerstacker,*? 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 closety connected by inter- mediate forms. I have alluded to a single instance of the separation and 62 Mr. Woodbury has published 63 “Annals and Mag. of Nat. Hist. several such accounts in ‘Journal of 3rd series, vol. xi. p. 339, Horticulture,’ 1861 and 1862. 316 SILK-MOTHS. — Cyan Was preservation of a particular stock of bees. Mr. Lowe“ pro- cured 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 appled exclusively to the workers, for, as we have seen, queens and drones cannot be selected and paired. SrtK-Morus. 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 correspond- ing 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 pro- duced. 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 silk-worms. 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 ® 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, how- 6 ‘The Cottage Gardener,’ May, 8° ¢ Transact. Entomolog. Soce.,’ 3rd 1860, p. 110; and ditto in‘ Journal series, vol. iii. pp. 143-173, and pp. of Hort.,’ 1862, p. 242. 295-3531. Cuar. VIIL. THEIR DIFFERENCES. 317 ever, 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 Con- stantinople in the sixth century, whence it was carried into Ttaly, and in 1494 into France.®® Everything has been favourable for the variation of this insect. It is believed to have been domesticated in China as long ago as 2700 B.c. It has been kept under unnatural and diversified conditions of life, and has been transported into many countries. There is reason to believe that the nature of the food given to the caterpillar influences to a certain extent the character of the breed.®? Disuse has apparently aided in checking the develop- ment 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,®* and the production of egos 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. The following details on the differences between the several breeds are taken, when not stated to the contrary, from M. Robinet’s excellent work,’® which bears every sign of care and large experi- ence. 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 86 Godron, ‘ De l’Espéce,’ 1859, tom. i. p. 460. The antiquity of the silk- worm in China is given on the authority Of Stanislas Julien. 87 See the remarks of Prof. West- wood, Gen. Hearsey, and others, at the meeting of the Entomolog. Soc. of London, July, 1361. 68 See, for instance, M. A..de Quatre- fages’ ‘Etudes sur les Maladies actu- elles du Ver a Soie,’ 1859, p. 101. 69 My authorities for the statements will be given in the chapter on Selec- tion. 70 ‘Manuel de l’Educateur de Vers 4 Soie,’ 1848, als SILK-MOTHS. Cuar, VIIL 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 eges 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 Treveltini races.” 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 tigrée, so called from being marked with transverse black stripes. As the general colour of the caterpillar is not correlated with that of the silk,” 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 df various breeds, are due to reversion; for the caterpillars of several allied wild species of Bombyx are marked and coloured in this manner. He separated some caterpillars with the tiger-like marks, and in the succeeding spring (pp. 149, 298) nearly all the caterpillars reared from them were dark-brindled, and the tints became still darker in the third generation. The moths reared from these caterpillars also became darker, and resembled in colouring the wild &. huftont. 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 Jarge 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 71 Robinet, ibid., pp. 12, 318. I may add that the eggs of N. American silkworms taken to the Sandwich Islands produced moths at very irre- gular 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 ‘Atheneum,’ 1844, p. 329, of J. Jarves’ ‘Scenes in the Sandwich Islands.’ 72 ¢The Art of revring Silk-worms,’ translated from Count Dandolo, 1825, 73 ¢Transact. Ent. Soc.,’ ut supra, pp. 153, 308. Cuap. VIII. THEIR DIFFERENCES. 319 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 occasion- ally 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 ™ 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 expérimentées, ont fait quatre mues a la seconde ou a la troisiéme année, ce qui semble prouver quwil a suffi de les placer dans des conditions favyorables pour leur rendre une faculté qu’elles avaient perdue sous des influences moins favorables.” Cocoons.—The caterpillar in changing into the cocoon loses about 50 per cent. of its weight; but the amount of loss differs in 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 Lace 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 beiug 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 genera- tions, the number of yellow cocoons in one breed has been reduced in France from one hundred to thirty-five in the thousand. According to Robinet, the white race, called Sina, by careful selection during the last seventy-five years, “est arrivée a un tel état de pureté, qu’on ne voit pas un seul cocon jaune dans des maillions de cocons blancs.” 7 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—lI can find no account of any constant difference in tiie moths of the most distinct races. Mrs. Whitby assured me that there was none in the several kinds bred by her; and I have "4 Kobinet, ibid., p. 317 75 Robinet, ibid., pp. 306-317. 320 SILK-MOTHS. Cuap. VIIL received a similar statement from the eminent naturalist, M. de Quatrefages. Captain Hutton also says*® 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 Bombycide “ fly swiftly in the day-time and evening, but the females are usually very sluggish and inactive.” ‘’ 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, hen 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‘*® 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 quwil y a la un véritable arrét de développement partiel.” On the other hand, he describes the female moths of the André Jean breed as having “leurs ailes larges et étalées. Un seul présente quelques courbures irréguliéres et des plis anormaux.” As moths and butter- flies of all kinds reared from wild caterpillars under confinement often have crippled wings, the same cause, whatever it may be, has probably acted on silk-moths, but the disuse of their wings during so many generations has, it may be suspected, likewise come into play. The moths of many breeds fail to glue their eggs to the surface on which they are laid,” but this proceeds, according to Capt. Hutton,*° 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 mul- berry-tree, often commit the strange mistake of devouring the base of the leaf on which they are feeding, and consequently fall 76 Robinet, ibid., p. 15. =e Quatrefages, ‘Btudes,’ &c., pp. — OL 5 yee CULTIVATED PLANTS. Cuap. UX. CHAPTER IX. CULTIVATED PLANTS: CEREAL AND CULINARY PLANTS, PRELIMINARY REMARKS ON THE NUMBER AND PARENTAGE OF CULTIVATED PLANTS——FIRST STEPS IN CULTIVATION — GEOGRAPHICAL DISTRIBUTION OF CULTIVATED PLANTS. CEREALIA.—»vousts 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.—caApBAGES: 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, SOMZ 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 bota- nists belheve 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 de- scended from one species, or from several inextricably com- mingled 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, &e.,and frequently it is not known how far their peculiarities ean be transmitted by seminal generation. Nevertheless. Cuap. IX. PRELIMINARY REMARKS Oue 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 enterimg on details a few general remarks on the origin of cultivated plants may be introduced. M. Alph. De Candolle! in an admirable discussion on this subject, in which he displays a wonderful amount of knowledge, gives a list of 157 of the most useful cultivated plants. Of these he believes that 85 are almost certainly known in their wild state; but on this head other competent judges? entertain ereat doubts. Of 40 of them, the origin is admitted by M. De Candolle to be doubtful, either from a certain amount of dissimuarity 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 con- dition. But it should be observed that he does not include in his lst 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® 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 1 Cnar. IX. PEAS. 345 This latter plant, in its cultivated state, differs in scarcely any character from the wild English carrot, except in general luxuri~ ance 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.*® 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. Carriére, 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 (#. sativus) in their roots, as well as the wonderful Chinese variety, R. cawdatus: (see ‘ Journal d’ Agriculture pratique,’ t. i., 1869, p. 159; also a separate essay, ‘ Origine des Plants Domestiques, 1869.) Raphanus raphanistrum aiid 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, p. 482) has now shewn that these differences, re- markable as they are, graduate away, the fruit of R. caudatus being intermediate. By cultivating 2. ruphanistrum during several generations (ibid., 1875, p. 9), Professor Hoffman also obtained plants bearing fruits like those of R. sativus, Fea (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.*! 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. Alefeld has recently studied * the genus with care, and, after having cultivated about fifty varieties, concludes that certainly they all belong to the same species. It is an interest- ing fact already alluded to, that, according to O. Heer,** the peas found in the lake-habitations of Switzerland of the Stone and Bronze ages, belong to an extinct variety, with exceedingly small 81 Alph. De Candolle, ‘Géograph. Bot., 960. Mr. Bentham (‘ Hort. Journal,” vol. ix, (1855), p. 141) believes that garden and field peas 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 raisedseveral distinct varieties. 89 Loudon’s ‘Encyclop. of Garden- ng, p- 39. 16 belong to the same species, and in this respect he differs from Dr. Targioni. 82 «Botanische Zeitung,’ 1860, s. 204. 83 “Die Pflanzen der Pfahlbauten, 1865, s. 22. 346 CULINARY PLANTS. Cuap. IX. 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,**—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 géant sans parchemin, which bears purple flowers, the. leafiets 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 Leguminose. 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 Pots nain hatif; but Hairs Dwarf Monmouth, which has large leaves, likewise has large flowers. The calyx in the Victoria Marrow is large, and in Bishop's Long Ped 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 Pots 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 Pos géant it 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 81 A variety called the Rounciva _ series), vol. i., 1835, p. 374, from attains this height, as is stated by Mr. which paper I have taken some facts Gordon in ‘ Transact. Hort. Soc.’ (2nd Cuar IX. PEAS, | 347 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. q A Ny Ay a th | i a. <> Ss = == — Se = <= —S ——— SSS = ~~ ——— > Q a Us = = & —— LS = SSS — =——— SSS ——— SSS SS SSS ——SS== Ss [SS <= 55 —2 Nh ) dH ; yu) i hh A Mt MiLTA } i SSS Z —=——F = = — = eas —=—= Fig. 41._Pods and Peas. I. Queen of Dwarfs. II. American Dwarf. III. Thurstun’s Reliance.—1V Pois Géant sais parchemin. a. Dan U’Rourke Pea. b, Queen of Dwarts Pea. c, Kuight’s Tall White Marrow. d. Lewis’s Negro Pea. 548 CULINARY PLANTS. ; Cuap. IX. 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, tegether with red passing through fine purple into a dark chocolate tint. These colours are either uniform or distributed in dots, strisz, 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-peus, with purple flowers, thin-skinned pods of an extraordinary shape, including large, dark-purple peas, grew wild by the side of the lowly Quzen of the Dwar/s, 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, ike 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*® 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 Siue Dwarf, which came out about the year 1820.°%° But the greater number of varieties have a singularly short existence: thus Loudon remarks* 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 SS 4 Pan, Pract 1729, p. 196. 87 «Encyclopedia of Gardening,’ p 86 ‘Gardener’s Magazine,’ vol. i, 823. 1826, p. 153. Cuap. IX. PEAS, 349 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 Leguminose, 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.** 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 * that varieties thus circumstanced would have oc- easionally eressed; and I shall give in the eleventh chapter two cases of this having occurred, as shown (in a manner hereafter to be ex- plained) by the pollen of the one variety having acted directly on the seeds of the other. Whether many of the new varieties which in- cessantly 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 °° of Knight’s four kinds of marrows, says, they have acquired a famous history, but their glory has departed. ; With respect to Beans (ba vulgaris), I will say but little. Dr. Alefeld has given 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 *8 See Dr. Anderson to the same ‘Gardener’s Chronicle,’ 1857, Oct. effect in the ‘ Bath Soc. Agricultural 25. Papers,’ vol. iv. p. 87. °0 “Gardener’s Chronicle,’ 1865, p. ®8 I have published full details of 387. experiments on this subject in the *! ¢ Bonplandia,’ x., 1862, s. 348 200 CULINARY PLANTS. Cuar. CX Switzerland *? by a peculiar and now extinct variety producing very small beans. . Pituto (Solanum tuberosum)—There is little doubt about the parentave 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.** The varieties cultivated in Britain are numerous; thus Lawson *® gives a de- scription 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.*° The fruit or berries also differ, but only in a slight degree.*’ The varieties are liable in very different degree to the attack of the Colorado potato-beetle.*$ 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 *’ 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 92 Heer,‘ Die Pflanzen der Pfahl- auten,’ 1866, s. 22. 93 Mr. Bentham informs me that in Poitou and the adjoining parts of 1845, p. 285. Sabine, in ‘Transact Hort. Soc.,’ vol. v. p. 249. 8° “Synopsis o the Vegetable Products of Scotland,’ quoted in France, varieties of Phaseolus vulgaris are extremely numerous, and so dif- ferent that they were described by Savi as distinct species. Mr. Bentham believes that all are descended from an unknown eastern species. Al- though the varieties differ so greatly in stature and in their seeds, “ there is a remarkable sameness in the ne- glected characters of foliage and tiowers, and especially in the brac- teoles, an insignificant character in the eyes even of botanists.” %4 Darwin, ‘ Journal of Researches,’ Wilson’s ‘ British Farming,’ p. 317. 86 Sir G. Mackenzie, in ‘ Gardener’s Chronicle,’ 1845, p. 790. 87 Putsche und Vertuch, ‘ Versuch einer Monographie der Kartoffeln,’ 1819,s.9,15. See also Dr. Anderson’s ‘ Recreations in Agriculture,’ vol. iy. Dd 20- 88 Walsh, ‘The American Entomvu- logist,’ 1859, p. 160. Also S. Tenney, ‘The American Naturalist,’ May, 1871, pura = °° * Gardener's Chronicle,’ 1862, p. 1052. Cuap. IX. POTATOES. Bt 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, &c., by which means the same individual is exposed during a length of time to diversified conditions, seedling potatoes generally display innumerable slight differences. Several varieties, even when propagated by tubers, are far from constant, as will be seen in the chapter on Bud-variation. Dr. Anderson’” procured seed from an Irish purple potato, which grew far from any other kind, so that it could not at least in this generation have been crossed, yet the many seedlings varied in almost every possible respect, so that “scarcely two plants were exactly alike.” Some of the plants which closely resembled each other above ground, pro- duced extremely dissimilar tubers; and some tubers which 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, 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.” 100 < Bath Society Agricult. Papers,’ 101 « Gardener’s Chronicle,’ 183, v. vol. v. p. 127. And ‘Recreations in 643, Agriculture,’ vol. v. p. 86. SA FRUITS. Cuap. X. CHAPTER X. PLANTS continued — FRUITS—ORNAMENTAL TREES—FLOWERS. FRUITS—crapres—VARY IN ODD AND TRIFLING PARTICULARS. MUL-= BERRY—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—SINGUIAR VARIETIES OF. APPLE.—— PEAR. STRAWBERRY—INTERBLENDING OF THE ORIGINAL FORMS. GOUSEBERRY—STEADY INCREASE IN SIZE OF THE FRUIT—VARIETIES OF. WALNUT. NOT. CUCURBITACEOUS PLANTS—-WONDERFCUL VARIA- TION OF. ORNAMENTAL TREES—THEIR VARIATION IN DEGREE AND KIND— ASH-TREE—SCOTCH-FIR—HAWTHORN. FLOW ERS—aMt.LtTIPLeE ORIGIN OF MANY KINDS—VARIATION IN CONSTITU- TIONAL PECULIARITIES—KIND OF VARIATION. R0SES—-SEVERAL SPECIES CULTIVATED. PANSY. DAHLIA. HYACINTH —HISTORY AND VARIATION OF. The Vine (Vitis vinifera). —TuHeE best authorities consider all our erapes as the descendants of one species which now grows wild in western Asia, which grew wild during the Bronze age in Italy,! and which has recently been found fossil in a tufaceous deposit in the south of France.2, 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* 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,* 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 varietics are 1 Heer, ‘Pflanzender Pfahlbauten, Saporta on the ‘Tertiary Plants of 1866, s. 28. France.’ 2 Alph. De Candolle, ‘Géograph. 3 Godron, ‘ De l’Espéce,’ tom. ii. p. Bot.. p. 872; Dr. A. ‘Targioni- 100. Tozzetti, in ‘Jour. Hort. Soc.,’ vol ix. * See an account of M. Vibert’s ex- p. 135. For the fossil vine found by periments, by Alex. Jordan, in ‘ Méin. Dr. G. Planchon, see ‘Nat. Hist. de )Acad. de Lyon,’ tom. ii. 1852, p. Review, 1865, April, p. 224. See 108. also the valuable works of M. de Cuap. XN. VINES. Os nr ay produced almost every year; for instance,? a golden-coloured variety has been recently raised in England from a black grape without the aid of a cross. Van Mons® reared a multitude of 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 characte1 both in the fruits and foliage. The cultivated varieties are extremely numerous; Count Odar 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 IT will not enter on this subject, nor on the many and great dif- ferences between the varieties. I will merely specify a few curious and trifling peculiarities, all taken from Odart’s highly esteemed work,’ for the sake of showing the diversified variability of this plant. Simon has classed grapes into two main divisions, those with downy leaves, and those with smooth leaves, but he admits that in one variety, namely the Rebazo, the leaves are either smooth, or downy; and Odart (p. 70) states that some varieties have the nerves alone, and other varieties their young leaves, downy, whilst the old ones are smooth. The Pedro-Ximenes grape (Odart, p. 397) presents a peculiarity by which it can be at once recognised amongst a host of other varieties, 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 (p. 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 gzapes have founded their main divisions on the berries being either round or oblong; and Odart admits the value of this character; yet there is one variety, the Maccabeo (p. 71), which often produces small round, and large oblong, berries in the same bunch. Certain grapes called Nebbiolo (p. 429) present a constant character, sufficient for their recognition, namely, “ the slight adherence of that part of the pulp which surrounds the seeds to the rest of the berry, when cut through transversely.” A Rhenish variety is mentioned (p. 228) which likes a dry soil; the fruit ripens well, but at the moment of maturity, if much rain falls, the berries are apt to rot; on the other hand, the fruit of a Swiss variety (p. 243) is valued for well sustaining prolonged humidity. This Jatter 5 ¢ Gardener’s Chronicle,’ 1864, p. _p. 290. £88. 7 Odart, ‘Ampélographie Untver 6 ¢ Aybres Fruitiers,’ 1836, tom. ii. _ selie,’ 1849. 354 Cuar, X. variety sprouts late in the spring, yet matures its fruit early ; other varieties (p. 362) have the fault of being too much excited by the April sun, and in consequence suffer from frost. A Styrian variety ‘p. 254) has brittle foot-stalks, so that the clusters of fruit are oiten blown eff; 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® 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 distinet 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. 2 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® 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 reso 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,” “so many varieties have been produced by cultivation that it is difficult to ascertain whether they FRUITS : 8 M. Bouchardat, in ‘ Comptes Ren- dus,’ Dec. Ist, 1851, quoted in ‘ Gar- dener’s Chron.,’ 1852, p. 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, p. 63, and ‘ Fifth Re- port,’ 1873, p. 66. ® «Etudes sur les Maladies actuelles du Ver 4 Soie,’ 1859, p. 321. 10 “Productive Resources of In lia, p- 130. Cuap. X. ORANGE GROUP. 305 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,™ 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” 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 2 wild state, is a distinct species; though its distinctness is doubted by another writcr “of great authority on such matters,” namely, Dr. Buchanan Hamilton. Alph. De Candolle,® 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 py seed. Sweet and bitter oranges ~ differ in no important respect except in the flavour of their fruit, but Gallesio * is most emphatic that both kinds can be propagated by seed with absolute certainty. Consequently, in accordance with his simple rule, he classes them as distinct species; as he does sweet and bitter almonds, the peach and nectarine, &c. He admits, however, that the soft-shelled pine-tree produces not only soft- shelled but some hard-shelled seedlings, so that a little greater force in the power of inheritance would, according to this rule, raise a soft-shelled pine-tree into the dignity of an aboriginally created species. The positive assertion made by Macfayden” that a iraite * ridu i? Citrus,= 1814: 12 Mr. Bentham, ‘ Review of Dr. A, ‘ Teoria della Riproduzione Vegetale,’ 1816. I quote chiefly from. this second werk. In 1839 Gallesio pub- lished in folio ‘Gli Agrumi dei Giard. Bot. di Firenze,’ in which he gives a curious diagram of the supposed selationship of all the forms, Targioni-Tozzetti, ‘ Journal of Hort. DOC VOleix. p. 133. 13 ¢Géograph. Bot.,’ p. 863. 14 ¢ Teoria della Riproduzione,’ pp. 52-57. 15 Hooker’s ‘Bot. Misc.,’ vol. i. p 302; vol. ii. p 111. 306 FRUITS: Cuap. X, 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; tor 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 re- produced 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 gencral 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,® they largely intercross without artificial aid; thus he positively states that seeds taken from lemon-trees (C. /lemonum) growing mingled with the citron (C. medic’), 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 swect orange, which grew cluse 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, &c., frequently cross in a state of nature. If indeed it were proved that plants of the orange tribe raised from these crosses were even partially sterile, it would be a strong argument in favour of their rank as species. Gallesio asserts that this is the case; but he does not distinguish between sterility from hybridism and from the effects of culture; and he almost destroys the force of this statement by another,” 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’* impregnated an orange with pollen from a lemon, and tlie 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 shail fully discuss in the following chapter. 16 ‘Teoria della Riproduzione,’ p. 53. 17 Gallesio, ‘Teoria della Riproduzione,’ p. 69, 18 Ibid. p. 67. et hoe Cuap. X. PEACH AND NECTARINE. ool The second remarkable fact is, that two supposed hybrids” (for their hybrid nature was not ascertained), between an orange and either a lemon or citron, produced on the same tree leaves, fiowers, 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,” is an analogous case, I know not. But to this subject I shall have to recur. I will conclude by giving from A. Risso*! a short account of a very singular variety of the common orange. It is the “ citrus aurantium fructu vuriubili,” which on the young shoots produces rounded-oyal 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- ereen colour embroidered with yellow, borne on footstalks without wings. The fruit whilst young is pear-shaped, yellow, longitu- dinally 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,* 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. Andrew Knight,” 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 1s a modified almond; and in this he has been followed by various authors.2* A first-rate peach, almost globular in shape, formed of soft and sweet 19 Gallesio, ‘Teoria della Ripro- duzione,’ pp. 75, 76. 20 ¢Gardener’s Chronicle,’ 1841, p. G13. 71 ¢ Annales du Muséum,’ tom. xx. p. 188. 22 “Journal of Hort. Soc.,’ vol. ii. series), vol. i. 1835, p. 56. See also p. 283. ‘ Cat. of Fruit in Garden of Hort. Soc.,’ 86 Alph, de Candolle, ‘Géograph. 3rd edit. 1842. 366 FRUITS : Cuap. X. In thecase of the almond, bitterness has been thought by some high authorities to indicate specitic difference. In N. America the Roman apricot-endures ‘“ cold and unfavour- able situations, where no other sort, except the Masculine, will succeed; and its blossoms bear. quite a severe frost without injury.” ® According to Mr. Rivers,®® 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,” ten varieties of the apricot, very different from each other, are cultivated, and all are raised from seed, excepting one, which is budded. Plums (Prunus insititia)—Formerly the sloe, P. spinosa, was thought to be the parent of all our plums; but now this honour is 1. Bullace Plum, 6. Denyer’s Victoria. 7. Diamond. Fig. 43.—Plum Stones, of natural size, viewed laterally. t 2. Shropshire Damson. 3. Blue Gage. 4. Urieans, 5. Elvas. very commonly accorded to P. insititia or the bullace, which is found wild in the Caucasus and N.-Western India, and is natural- ised in England.” It is not at all improbable, in accordance with some observations made by Mr. Rivers,” 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, 68 Downing, ‘The Fruits ot Ame- vinces,’ vol. i. 1841, p. 295. rica,’ 1845, p. 157: with respect to 71 See an excellent discussion on the Alberge apricot in France, see p. 53: 8 ¢ Gardener’s Chronicle. |n 3 564. 70 «Travels in the Himalayan Pro- 7=-=59p90 1863, p- this subject in Hewett C. Watson’s ‘ Cybele Britannica,’ vol. iv. p. 80. ‘2 *Gardener’s Chronicle,’ 1855, p O77 al. Onap, X. PLUMS. 367 Godron remarks ® 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 Mowers 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 “™ 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 ereengage. 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, promi- nence 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 *3 “De l’Espéce,’ tom.ii. p.94. On 278, 284, 310, 314. Mr. Rivers the parentage of our plums, see also’ raised (‘Gard. Chron.,’ 1863, p. 27) Alph. De Candolle, ‘Géograph. Bot.,’ from the Prune-péche, which bears p- 878. Also Targioni-Tozzetti,‘Jour- latge, round, red plums on stout, nal Hort. Soc.,’ vol. ix. p. 164, Also robust shoots, a seedling which bears Babington, ‘ Manual of Brit. Botany,’ — oval, smailer fruit on shoots that are 1851, p. 57. so slender as to be almost pendulous, 74 ¢Fruits of America,’ pp. 276. 368 FRUITS : Cuap. X. 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.” 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.** 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” 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, &c.).—Botanists believe that our cultivated cherries are descended from one, two, four, or even more wild stocks.** That there must be at least two parent species we may infer from the sterility of twenty hybrids raised by Mr. Knight from the morello fertilized by pollen of the Elton cherry; for these hybrids produced in all only five cherries, and one alone of these mn 75 “ Gardener’s Chronicle, 1855, p. ’s ‘Fruit Trees.” p. 278. 77 ¢Gardener’s Chronicle,’ 1863, p. 27. Sageret, in his ‘ Pomologie Phys.,’ p. 345, enumerates five kinds which can be propagated in France by seed: see also Downing’s ‘Fruit Trees of America,’ p. 3035, 312, &e. ‘8 Compare Ot ide) S nica,” vol, i. p. 354 its nearly sessile fruit, ranges farther south than the long-stalked P. acerba, which is entirely absent in Madeira, the Canaries, and apparently in Por- tugal. This fact supports the belief that these two forms deserve to be called species. But the characters separating them are of slight import- ance, and of a kind known to vary in other cultivated fruit-trees. 83 Sve ‘Journ. of Hort. Tour, by Deputation of the Caledonian Hort. 85 Loudon’s ‘ Gardener’s Mag.,’ vol. vi., 1830, p. &3. 87 Sve ‘Catalogue of Fruit in Gar- den of. Hort. Soe.” 1842, and Downine’s ‘ American Fruit Trees.’ 88 Loudon’s - Gardener's Magazine,’ vol. iv., 1828>p. 112. £9 ¢The Culture of the Apple,’ p. 43. Van Mons makes the same remarg on the pear, ‘ Arbres Fruitiers,’ tom. ii, 1836., p. 414. 0 Jindley’s ‘ Horticulture,’ p. 116 Cuar. X. APPLES, 371 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 Calville rouge de Micoud occasionally bears two crops during the same year. The Burr Knot is covered with small exerescences, 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." Mr. Rivers has recently described *? 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® 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 state- ment with respect to a cider apple, and adds that he only once saw these insects just above the stock, but that three days after- wards 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 specific- ally distinct. The famous St. Valery apple must not be passed over; the flower has a double calyx with ten divisions, and fourteen styles sur- mounted by conspicuous oblique stigmas, but is destitute of starnens or corolla. The fruit is constricted round the middle, and is formed of five seed-cells, surmounted by nine other cells.** Not being See also Knight on the Apple-Tree, in ‘Transact. of Hort. Soc.,’ vol. vi. p. 229. Ze omuransact.; Hort. SoC.) vol. 4. 1812, p. 120. 82 ¢ Journal of Horticulture,’ March 13th. 1866, p. 194. 93 «Transact. Hort. Soc.,’ vol. iv. p. 68. For Knight’s case, see vol. vi. p. 547. When the coccus first appeared in this country, it is said (vol. ii. p. 163) that it was more injurious to crab-stocks than to the apples grafted on them. The Majetin apple has been found equally free of the coccus at Mel- bourne in Australia (‘ Gard. Chron.’ 1871, p. 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 jer cent. of lime, while that of the crab exhibited not quite 23 per cent. In Tasmania Mr. Wade (‘ Transact. New Zealand Institute,’ vol. iv., 1871, p- 431) raised seedlings of the Siberian Bitter Sweet for stocks, and he found barely one per cent. of them attacked by the coccus. Riley snows (‘ Fitth Report on Insects of Missouri,’ 1873, p. 87) that in the United States some varieties of apples are highly attrac- tive to the coccus and others very little so. Turning to a very ditferent pest, namely, the caterpillar of a moth (Carpocapsa pomonella), Walsh affirms (‘ The American Entomologist,’ April, 1859, p. 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.” 94 © Mém. de Ja Soc. Linn. de Paris,’ tom. iii., 1825, p. 164; and Seringe, ‘Bulletin Bot.’ 1830, p. 117. V2 FRUITS: Cap. X. provided with stamens, the tree requires artificial fertilisation; and the girls of St. Valery annually go to “futre 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,®? 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 -trictly inherited. No one can raise, for instance, from the seed of ‘he Ribsten 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.°° Yet ii was a mistake to suppose that with most varieties the characters are not to a certain extent nmherited. 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 resembis to a certain extent their parents. We see this indeed in the severa! sub-groups of Russetts, Sweetings, Codlins, Pearmains, Reinettes, &e.,2° which are all believed, and many are known, to be descended from other varieties bearing the same names. cultivated, in its fruit, flowers, and foliage. One of the most celebrated botanists in Europe, M. Decaisne, has carefully studied the many varieties ;** 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. Deecaisne 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.* Strawberries (Fragaria).—This fruit is remarkable on account of the number of species which have been cultivated, and from 95 ¢Gardener’s Chronicle,’ 1849, p. 74 — . #6 R. Thompson, in ‘ Gardener’s Chron.,’ 1850, p. 788. %7 Sageret, ‘Pomologie Physiolo- gique,’ 1830, p. 263. Downing’s ‘Fruit Trees,’ pp. 150, 134, 139, &c. Loudon’s ‘(tardener’s Mag.,’ vol. viii. p. 317. Alexis Jordan, ‘ De l’Origine des diverses Varietés,’ in ‘Mém. de V’Acad. Imp. de Lyon,’ tom. ii., 1852, pp. 95, 114. _ ‘ Gardener’s Chronicle,’ 1859, pp. 774, 788. 98 “Comptes Rendus,’ July 6th, 1853. °° *Gardener’s Chronicle,’ 1856, p. 894; 1857, p. 820; 1862, p. 1199. Cuar. X. STRAWBERRIES. 1d 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. 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 severai species are almost Innumerable. The species consist of, firstly, the wood or Alpine cultivated strawberries, descended froin 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. colina, and little cultivated in England. Thirdly, the Hautbois, from the European £. elutior, Fourthly, the Scarlets, descended from #'. 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.“'! These five or six forms have been ranked by most botanists as specifically distinct; but this may be doubted, for Andrew Knight, who raised no less than 400 crossed straw- berries, asserts that the /. virginiana, chiloensis and grandiflora “may be made to breed together indiscriminately,” and he found, in accordance with the principle of analogous variation, “that similiar varieties could be obtained from the seeds of any one of them.” Since Knight’s time there is abundant and additional evidence !% of the extent to which the American fcrms spontaneously cross. We owe indeed to such crosses most of our choicest existing 100 Most pf the largest cultivated strawberries are the descendants of /. grandiflora or chiloensis, and I have seen no account of these forms in their wild state. Methuen’s Scarlet (Downing, ‘Fruits, p. 527) has “immense fruit of the largest size,” and belongs to the section descended from F. virginiana ; and the fruit of this species, as I hear from Prof. A. Gray, is only a little larger than that of F. vesca, or our common wood- strawberry. 101 ¢Ve Fraisier,’ par le Comte L. de Lambertye, 1864, p. 50. 102 “Transact. Hort. Soc.,’ vol.. iii. 1820, p. 207. 103 See an account by Prof. Decaisne, and by others in ‘Gardener’s Chron- icle,’ 1862, p 335, and 1858, p. 172; and Mr. Barnet’s paper in ‘ Hort. Soc. Transact.,’ vol. vi. 1826, p. 170. a7+t FRUITS: Cuap. X varieties. Knight did not succeed m crossing the European wood- strawberry with the American Scarlet or with the Hautbois. Mr. Williams of Pitmaston, however, succeeded; but the hybrid ofispring from the Hautbois, though fruiting well, never produced seed, with the exception of a single one, which reproduced the parent hybrid form.’* 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.%° We thus see*® 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 some- times 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.” 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 spedes 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.” Although the fruit in the different varieties differs so ereatly 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, isy according to De Jonghe, absolutely the same in all: and this no doubt 104 Transact. Hort. Soc.,’ vol. v. 1824. p. 294. 105 ¢ Journal of Horticulture,’ Dec. 30th, 1862, p. 779. See also Mr. Prince to the same effect, ibid., 1863, p. 418. 106 For additional evidence see ¢ Journal of Horticulture,’ Dec. 9th, 1862, p. 721. 107 «Te Fraisier,’ par le Comte Le de Lambertye, pp. 221, 230. 108 ¢ Transact. Hort. Soc.,’ vol. vi. . 200. 109 «Gardener’s Chron.,’ 173. 1858, p Cuap. X. STRAWBERRIES. 37) may be accounted for by the seed being of no value, and conse- quently 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 Linnzus 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? A variety raised by Mr. Myatt,"' 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 /. collina. The Red Bush Alpine strawberry (one of the F. vescw 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 pro- pagated by seed it often degenerates and produces plants with runners. 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,”® and she produced three calves which had on the same side of the head, instead of a horn, a small bony lump attached 23 These statements are taken from the following works in oraer:—Youatt on *The Horse,’ p. 48; Mr. Darvill, in ‘The Veterinary,’ vol. viii. p. 50. With respect to Robson, see ‘The Veterinary,’ vol. iii. p. 580; Mr. Lawrence on ‘ The Horse,’ 1829, p. $; ‘The Stud Farm,’ by Cecil, 1821 Baron Cameronn, quoted in ‘The Veterinary,’ vol. x. p. 500. *4 ‘Recreations in Agriculture and Nat. Hist.,’ vol. i. p. 68. *5 “Ueber die Eigenschaften,’ &c., 1828, s. 107. *6 Bronn’s ‘ Geschichte der Natur,’ Band ii. s. 132. Car. XIL. : INHERITANCE. 457 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. Poly- dactylism graduates *’ by multifarious steps from a mere cutaneous appendage, not including any bone, to a double hand. But an ad- ditional 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. Gene- rally, through the law of correlation, both hands and both feet are similarly affected. Dr. Burt Wilder has tabulated” 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 normaly exceeded by any existing mammal, bird, or reptile. Nevertheless, supernumerary digits are strongly inherited ; they have been transmitted through five genera- tions; and in some cases, after disappearing for one, two, or even three generations, have reappeared through reversion. These facts are rendered, as Protessor 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 1-82nd 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 dizits are often amputated soon after birth, and can seldom have peen strengthened by use. Dr. 27 Vrolik has discussed this point pp. 684. at full length in a work published 28 « Massachusetts Medical Society,” in Dutch, from which Sir J. Paget vol. ii. No. 3; and ‘Proc. Boston has kindly translated for me passages. Soc. of Nat. Hist.,’ vol. xiv.. 1871, p. See, also, Isidore Geoffroy St. Hilaire’s 154. ‘Hist. des Anomaties,’ 1832, tom. 3 £58 INHERITANCE. Cuap. RIT 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 inherit- ance. 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,” thickened joints, crooked fingers, &c., 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.” 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 (Sa/amandra 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 metamor- phosis, 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, &c., are often accompanied by polydactylism.** 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 On the inheri- 22 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,’ Ap. 1872. 30 For these several statements, sze Dr. Struthers, ‘ Edinburgh New Phil. Journal,’ July, 1863, especially on intermissions in the line of descent. Prof. Huxley, ‘ Lectures on our Know- ledge of Organic Nature,’ 1853, p. 97. With respect to inheritance, see Dr. Prosper Lucas, ‘ L’Hérédité Nat.,’ tom. i. p. 325. Isid. Geoftroy, ‘ Anom.,’ tom. i. p. 701. Sir A. Carlisle, in ‘Phil. Transact.,? 1814, p. 94. A. Walker, on ‘Intermarriage,’ 1838, p. 140, gives a case of five genera- tions ; as does Mr. Sedgwick, in ‘ Brit. and Foreign Medico-Chirurg. Review,’ April, 1855, p. 462. tance of other anomalies in the ex- tremities, see Dr. H. Dobell, in vol. xlvi. of ‘ Medico-Chirurg.Transactions,’ 1863; also Mr. Sedgwick, in op. cit., April, 1863, p. 460. With respect to additional digits in the negro, see Prichard, ‘ Physical History of Man- kind.” Dr. Dieffenbach (‘Jour. Royal Geograph. Soc.,’ 1841, p. 208) says this anomaly is not uncommon with the Pulynesians of the Chatham Islands; and I have heard of several cases with Hindus and Arabs. 31 Meckel and Isid G. St. Hilaire insist on this fact. See, also M. A. Roujou, ‘Sur quelques Analogies du Type Humain,’ p. 61; published, I believe, in the ‘ Journal of the Anthro. polog. Soc. of Paris,’ Jan. 1872. Cuar. XIL. INHERITANCE. 459 four-toed breeds. 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 per- fectly, 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. 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 re- produced a nail. Dr. Struthers mentions a case of the partial re-growth 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 ampu- tation; 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, to- gether 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 de- gree 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 hesi- tation, 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** states that brothers and sisters of the same 32 ¢The Poultry Chronicle,’ 1854, p- 909. 33 The statements in this paragraph are taken from Isidore Geoffroy St. Hilaire, ‘ Hist. des Anomalies,’ tom. i. pp. 688-693. Mr. Goodman gives, ‘Phil. Soc. of Cambridge,’ Nov. 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. 34 ¢ Medical Notes and Reflections,’ 1839, pp. 24, 34. See, also, Dr. P Lucas, ‘ L’Héréd. Nat.,’ tom. ii. p. 33. 460 INHERITANCE. Cuap. XI. 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 pecuhar 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** 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 °° 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 surpris- ing 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 35 ¢Du anger des Mariages Con- Chirurg. Review, July, 1833, pp sanguins,’ 2nd edit., 1862, p. 103. 183, 189. 3°‘ British and Fereign Medico- Cras. XII. INHERITANCE. 461 capacity in the individual: thus Vilmorin *’ 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 te 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. ven 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.** Mr. Rivers fertilized, as he informs me, the flowers of a new Beigian weeping thorn (Crateyus 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,” 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.*#? Lastly, a weeping or rather a prostrate yew (Tu«xus baccaia) 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.4 These facts, it might have been thought, would have been sufficient similar statement in ‘Proc. Nat. of Philadelphia,’ 1872, p. 255. 37 Verlot, ‘La Product. des Varié- tes, 1865, p. 32. 38 Loudon’s ‘ Gard. Mag.,’ vol. xii., 1336, p. 368. 39 Verlot, ‘La Product. des Vari¢- tés,’ 1865, p. 94. 49 Bronn’s ‘Geschichte der Natur,’ BD. li.s. 121. Mr. Meenan makes a 41 Rey. W. A. Leighton, ‘ Flora of Shropshire,’ p. 497; and Charles- worth’s ‘Mag. of Nat. Hist.,’ vol. i. 1837, p. 30. I possess prostrate trees produced from these seeds. 462 INHERITANCE. Cuap. XII. 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 # 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. Ander- son, 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.* 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 (/’. 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 seed- ling out of above twenty thousand ; whereas the weeping variety of a second species of ash, which could not, whilst grown in the same garden, retainits 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 scedlings of a variety of the Cerasuspadus, with yellow fruit, bore yellow fruit: one-twelfth of the seedlings of the variety of Cornus mascula, with yellow fruit, came true : * and lastly, all the trees raised by my father from a yellow-berried holly (lex aquifolium), founds wild, 4? Verlot, op. cit., p. 93. 1833, p. 597. #3 For these several statements, sez #4 These statements are taken from Loudon’s ‘Gard. Magazine, vol. x. Alph. De Candolle, ‘ Bot. Géograph., 1834, pp. 408, 180; and vol. ix., p. 1083, Cuar. XII INHERITANCE. 463 produced yellow berries. Vilmorin * observed in a bed of Saponaria calabricu 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 Yugetes 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 mtermediate 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 generaily transmit their colour much more truly than any other variety. This fact probably stands in close relation with one observed by Verlot,*® namely, that flowers which are normally white rarely vary into any other colour. J 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 plaut, 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;”*7 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. 45 Verlot, op. cit., p. 38. 47 Alph. De Candolle, ‘ Géograph. %@ Op. cit., p. 59. Bot.,’ p. 1082. 464 INHERITANCE. Cuap. XM. 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.*® 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 vulyaris), from their reciprocally dimorphic structure; *° 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 singie 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 (Velphinium consolida and ajacis), of which common seedlings preseit a greater diversity of colour than any other plant known to me; yet on procuring seed of five named German varieties of D. conso/ida, 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 dis- tinguished botanist maintains that the annual species of Delphinium are always self-fertilised; therefore I may mention that thirty-twe 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, &c., 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 48 See ‘Cottage Gardener,’ April 49 Darwin, in ‘Journal of Proce. 10, 1860, p. 18, and Sept. 10, 1861, Linn. Soc. Bot.’ 1862, p. 94. p. 456; ‘ Gard. Chron.,’ 1845, p. 102. SerAP. XII. INHERITANCE. 465 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.”° In a few rare cases peculiarities fail to be inherited, appa- rently from the force of inheritance being too strong. I have been assured by breeders of the canary-bird that to get a good jonguil-coloured bird it does not answer to pair two jonquils, as the colour then comes out too strong, or 1s 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: °! 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 exces- sive, and the crest itself fails to be developed. Mr. Hewitt, speaking of Laced Sebright Bantams, says ** that, ‘why this should be so I know not, but Iam 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 asingle instance: not onescholar out of 148, who were at the same time in the London Institu- tion, was thé 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 50 Hofacker, ‘Ueber die Eigen- that he believes that these statements schaften,’ &c., s. 10. are correct. 51 Bechstein, ‘ Naturgesch. Deutsch- 52 ¢The Poultry Book,’ by W. B. lands,’ b. iv. s. 462. Mr. Brent, a Tegetmeier, 1866, p. 245. great breeder of canaries, informs me 21 166 INHERITANCE. Cuar. XIL two deaf and dumb children were produced. Mr. Sedgwick,*3 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,°* 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 cata- logue 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, 1t was until lately difficult to come to any definite conclusion. Some mutilations have been practised for a vast number of generations without any in- herited result. Godron remarks *° 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 mutila- tions have ever been inherited.°® Adhesions due to in- 53 ¢ British and Foreign Med.- 55 Nevertheless Mr. Wetherell states, Chirurg. Review,’ July, 1861, pr. 200-204. Mr. Sedgwick has given such full details on this subject, with ample references, that I need refer to no other authorities. 54 Mr. Sproule, in ‘ British Medical Journal,’ April 18, 1863. 53 De 1]’Espéce,’ tom. ii., 1859, p. 299. ‘Nature,’ Dec. 1870, p. 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.” Cuar. XII. INHERITANCE. 467 flammation 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 5’ that Jews are often born in Germany in a condition rendering circum- cision 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 coincidences, 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 aifected 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-Séquard®® gives the following summary of his observations on guinea-pigs; and this summary is so important that I will quote the whole :— 5” ‘Philosoph. Mag.’ vol. iv., 1799, 1875, p. 7. The extracts are from Dd. 3. this last paper. It appears that 58 ¢ Proc, Royal Soc.,? vol. x. p. Obersteiner, ‘Stricker’s Med. Jahr- 297. ‘Communication to the Brit. biicher,’ 1875, No, 2, has confirmea Assoc,’ 1870. ‘The Lancet,’ Jan. Brown-Séquard’s observations. 468 INHERITANCE. Cuap, XIL “Ist. 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. “Ath, 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 cf 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 haying been made in most cases only on one of the corpora restiformia. “ 6th. Hoeematomaand 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 anesthetic from a section of the sciatic nerve alone, or of that nerve and also of the crural. Some- times, 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-Séquard 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-Séquard speaks of such cases as one of the rarer forms of inheritance. It isa still more interesting fact-— Csap. XII. INHERITANCE. 469 “That the sciatic nerve in the congenitally toeless animal has inherited the power of passing through ali 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 acertain order.” In most of the cases of inheritance recorded by Brown-Sé- quard only one of the two parents had been cperated 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. Proper Lucas has collected a long list of inherited injuries. A few instances will suffice. A cow lost a horn from an accident with consequent suppur- _ ation, 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. Blumen- bach 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.5® 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 Some curious cases are 5° This last case is quoted by Mr. Sedgwick in ‘British and Foreign Medico-Chirurg. Review,’ April, 1861, p- 484. For Blumenbach, see above- cited paper. See, also, Dr. P. Lucas, ‘Traité de l’Héréd. Nat.,’ tom. ii. p. 492, Also, ‘Transact. Linn. Soc.,’ vol. ix. p. 323. given by Mr. Baker in the ‘ Veterinary,’ vol. xiii. p. 723. Another curious case is given in the ‘Annales deg Scienc. Nat.,’ Ist series, tom. xi. p 324, 470 INHERITANCE. Cuar. XIL 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, &c., amputated or injured, produc- ing offspring with the same parts ill-formed; but as it is not very rare for similar malformations to appear spontaneously, al] 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 ;” © 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-Séquard’s observa- tions, that the effects of injuries, especially when followed by disease, or perhaps exclusively when thus followed, are occasionally inherited.*! 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, reduced on the same part of these feathers, it seems extremely probable, 6° «The Dog,” by Stonehenge, 1867, p. 118. 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 as Mr. Salvin remarks (‘ Proc. Zoolog. Soc.’ 1873, p. 429), that this is due to the inherited effects of long-continued mutilation. Cuap, XII. NON-INHERITANCE. 471 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 Carriére,* 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 trans- mitted, 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, &c., 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 prin- ciple of reversion. But scarcely a single seedling will be found perfectly to resemble the parent-form; and this 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 mani- festly affected, they produce seedlings which vary greatly, 62 ¢ Production et Fixation des Varictés,’ 1865, p. 72. 472 INHERITANCE. Cuap. XII. departing from the parental type in many characters.® 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, &c., 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 con- ditions, and are not liable to such extreme variability; there- fore 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 com- monly 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 in- herited. 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 trans- mitted. 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 cau be assigned. The effects of injuries or mutilations are occasionally inherited; and we 63 Downing, * Fruits of America,’ p. 5: Sageret, ‘Pom. Phys.,’ pp. 43, 72. Crap, XII. INHERITANCE, A473 shall see in a future chapter that the long-coniinued use and disuse of parts produces an inherited effect. ven those cha- racters 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, con- ditions 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 cr more remote ancestors instead of its immediate parents. This latter - subject will be discussed in the following chapter. END OF VOL. TI. Darwin, Chas Variation « 6019 dom RARIES \s and plan N LIB f anima STITUTIO CY i ks yr a Sp lg i AN IN variation 0 nhbae 4 {e) 0) 7) f= x re = = tte 7) >