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The Variation of Animals and Plants Under Domestication, Vol. I.
by Charles Darwin
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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;[400] but the crest, which is supported on the protuberance, is at first feebly developed, nor does it attain its full size until the second year. The Spanish cock is pre-eminent for his magnificent comb, and this is developed at an unusually early age; so that the young males can be distinguished from the females when only a few weeks old, and therefore earlier than in other breeds; they likewise crow very early, namely, when about six weeks old. In the Dutch sub-breed of the Spanish fowl the white ear-lappets are developed earlier than in the common Spanish breed.[401] Cochins are characterised by a small tail, and in the young cocks the tail is developed at an unusually late period.[402] Game fowls are notorious for their pugnacity; and the young cocks crow, clap their little wings, and obstinately fight with each other, even whilst under their mother's care.[403] "I have often had," says one {251} author,[404] "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." With the males of all gallinaceous birds the use of their weapons and pugnacity is to fight for the possession of the females; so that the tendency in our Game chickens to fight at an extremely early age is not only useless, but is injurious, as they suffer so much from their wounds. The training for battle during an early period may be natural to the wild Gallus bankiva; but as man during many generations has gone on selecting the most obstinately pugnacious cocks, it is more probable that their pugnacity has been unnaturally increased, and unnaturally transferred to the young male chickens. In the same manner, it is probable that the extraordinary development of the comb in the Spanish cock has been unintentionally transferred to the young cocks; for fanciers would not care whether their young birds had large combs, but would select for breeding the adults which had the finest combs, whether or not developed at an early period. The last point which need here be noticed is that, though the chickens of Spanish and Malay fowls are well covered with down, the true feathers are acquired at an unusually late age; so that for a time the young birds are partially naked, and are liable to suffer from cold.

Secondary Sexual Characters.—The two sexes in the parent-form, the Gallus bankiva, differ much in colour. In our domestic breeds the difference is never greater, but is often less, and varies much in degree even in the sub-breeds of the same main breed. Thus in certain Game fowls the difference is as great as in the parent-form, whilst in the black and white sub-breeds there is no difference in plumage. Mr. Brent informs me that he has seen two strains of black-breasted red Games, in which the cocks could not be distinguished, whilst the hens in one were partridge-brown and in the other fawn-brown. A similar case has been observed in the strains of the brown-breasted red Game. The hen of the "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 {252} 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.[405] A similar relation is also evident when we compare the several varieties of Cochins. In the two sexes of gold and silver-spangled and of buff Polish fowls, there is much general similarity in the colouring and marks of the whole plumage, excepting of course in the hackles, crest, and beard. In spangled Hamburghs, there is likewise a considerable degree of similarity between the two sexes. In pencilled Hamburghs, on the other hand, there is much dissimilarity; the pencilling which is characteristic of the hens being almost absent in the males of both the golden and silver varieties. But, as we have already seen, it cannot be given as a general rule that male fowls never have pencilled feathers, for Cuckoo Dorkings are "remarkable from having nearly similar markings in both sexes."

It is a singular fact that the males in certain sub-breeds have lost some of their secondary masculine characters, and, from their close resemblance in plumage to the females, are often called hennies. There is much diversity of opinion whether these males are in any degree sterile; that they sometimes are partially sterile seems clear,[406] but this may have been caused by too close interbreeding. That they are not quite sterile, and that the whole case is widely different from that of old females assuming masculine characters, is evident from several of these hen-like sub-breeds having been long propagated. The males and females of gold and silver-laced Sebright Bantams can be barely distinguished from each other, except by their combs, wattles, and spurs, for they are coloured alike, and the males have not hackles, nor the flowing sickle-like tail-feathers. A hen-tailed sub-breed of Hamburghs was recently much esteemed. There is also a breed of Game-fowls, in which the males and females resemble each other so closely that the cocks have often mistaken their hen-feathered opponents in the cock-pit for real hens, and by the mistake have lost their lives.[407] The cocks, {253} 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[408] 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."[409]

I have remarked in my 'Origin of Species' that secondary sexual characters are apt to differ much in the species of the same genus, and to be unusually variable in the individuals of the same species. So it is with the breeds of the fowl, as we have already seen, as far as the colour of plumage is concerned, and so it is with the other secondary sexual characters. Firstly, the comb differs much in the various breeds,[410] and its form is eminently characteristic of each kind, with the exception of the Dorkings, in which the form has not been as yet determined on by fanciers, and fixed by selection. A single, deeply-serrated comb is the typical and most common form. It differs much in size, being immensely developed in Spanish fowls; and in a local breed called Red-caps, it is sometimes "upwards of three inches in breadth at the front, and more than four inches in length, measured to the end of the peak behind."[411] In some breeds the comb is double, and when the two ends are cemented {254} together it forms a "cup-comb;" in the "rose-comb" it is depressed, covered with small projections, and produced backwards; in the horned and creve-coeur fowl it is produced into two horns; it is triple in the pea-combed Brahmas, short and truncated in the Malays, and absent in the Guelderlands. In the tasselled Game a few long feathers arise from the back of the comb; in many breeds a crest of feathers replaces the comb. The crest, when little developed, arises from a fleshy mass, but, when much developed, from a hemispherical protuberance of the skull. In the best Polish fowls it is so largely developed, that I have seen birds which could hardly pick up their food; and a German writer asserts[412] that they are in consequence liable to be struck by hawks. Monstrous structures of this kind would thus be suppressed in a state of nature. The wattles, also, vary much in size, being small in Malays and some other breeds; they are replaced in certain Polish sub-breeds by a great tuft of feathers called a beard.

The hackles do not differ much in the various breeds, but are short and stiff in Malays, and absent in Hennies. As in some orders of birds the males display extraordinarily-shaped feathers, such as naked shafts with discs at the end, &c., the following case may be worth giving. In the wild Gallus bankiva and in our domestic fowls, the barbs which arise from each side of the extremities of the hackles are naked or not clothed with barbules, so that they resemble bristles; but Mr. Brent sent me some scapular hackles from a young Birchen Duckwing Game cock, in which the naked barbs became densely reclothed with barbules towards their tips; so that these tips, which were dark coloured with a metallic lustre, were separated from the lower parts by a symmetrically-shaped transparent zone formed of the naked portions of the barbs. Hence the coloured tips appeared like little separate metallic discs.

The sickle-feathers in the tail, of which there are three pair, and which are eminently characteristic of the male sex, differ much in the various breeds. They are scimitar-shaped in some Hamburghs, instead of being long and flowing as in the typical breeds. They are extremely short in Cochins, and are not at {255} all developed in Hennies. They are carried, together with the whole tail, erect in Dorkings and Games; but droop much in Malays and in some Cochins. Sultans are characterized by an additional number of lateral sickle-feathers. The spurs vary much, being placed higher or lower on the shank; being extremely long and sharp in Games, and blunt and short in Cochins. These latter birds seem aware that their spurs are not efficient weapons; for though they occasionally use them, they more frequently fight, as I am informed by Mr. Tegetmeier, by seizing and shaking each other with their beaks. In some Indian Game-cocks, received by Mr. Brent from Germany, there are, as he informs me, three, four, or even five spurs on each leg. Some Dorkings also have two spurs on each leg;[413] and in birds of this breed the spur is often placed almost on the outside of the leg. Double spurs are mentioned in the ancient Chinese Encyclopaedia. Their occurrence may be considered as a case of analogous variation, for some wild gallinaceous birds, for instance, the Polyplectron, have double spurs.

Judging from the differences which generally distinguish the sexes in the Gallinaceae, certain characters in our domestic fowls appear to have been transferred from the one sex to the other. In all the species (except in Turnix), when there is any conspicuous difference in plumage between the male and female, the male is always the most beautiful; but in golden-spangled Hamburghs the hen is equally beautiful with the cock, and incomparably more beautiful than the hen in any natural species of Gallus; so that here a masculine character has been transferred to the female. On the other hand, in cuckoo Dorkings and in other cuckoo breeds the pencilling, which in Gallus is a female attribute, has been transferred to the male: nor, on the principle of analogous variation, is this transference surprising, as the males in many gallinaceous genera are barred or pencilled. With most of these birds head ornaments of all kinds are more fully developed in the male than in the female; but in Polish fowls the crest or top-knot, which in the male replaces the comb, is equally developed in both sexes. In certain {256} sub-breeds, which, from the hen having a small crest, are called lark-crested, "a single upright comb sometimes almost entirely takes the place of the crest in the male."[414] From this latter case, and from some facts presently to be given with respect to the protuberance of the skull in Polish fowls, the crest in this breed ought perhaps to be viewed as a feminine character which has been transferred to the male. In the Spanish breed the male, as we know, has an immense comb, and this has been partially transferred to the female, for her comb is unusually large, though not upright. In Game-fowls the bold and savage disposition of the male has likewise been largely transferred to the female;[415] and she sometimes even possesses the eminently masculine character of spurs. Many cases are on record of hens being furnished with spurs; and in Germany, according to Bechstein,[416] the spurs in the Silk-hen are sometimes very long. He mentions also another breed similarly characterized, in which the hens are excellent layers, but are apt to disturb and break their eggs owing to their spurs.

Mr. Layard[417] 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 finds that the same rule holds good with this breed near Calcutta. The males and females, on the other hand, of the black-boned European breed, with silky feathers, do not differ from each other; so that in the one breed black skin and bones, and the same kind of plumage, are common to both sexes, whilst in the other breed these characters are confined to the female sex.

At the present day all the breeds of Polish fowls have the great bony protuberance on their skulls, which includes part of the brain and supports the crest, equally developed in both sexes. {257} But formerly in Germany the skull of the hen alone was protuberant: Blumenbach,[418] who particularly attended to abnormal peculiarities in domestic animals, states, in 1813, that this was the case; and Bechstein had previously, in 1793, observed the same fact. This latter author has carefully described the effects of a crest on the skull not only in fowls, but in ducks, geese, and canaries. He states that with fowls, when the crest is not much developed, it is supported on a fatty mass; but when much developed, it is always supported on a bony protuberance of variable size. He well describes the peculiarities of this protuberance, and he attended to the effects of the modified shape of the brain on the intellect of these birds, and disputes Pallas' statement that they are stupid. He then expressly states that he never observed this protuberance in male fowls. Hence there can be no doubt that this remarkable character in the skulls of Polish fowls was formerly in Germany confined to the female sex, but has now been transferred to the males, and has thus become common to both sexes.

External Differences, not connected with the sexes, between the breeds and between individual birds.

The size of the body differs greatly. Mr. Tegetmeier has known a Brahma to weigh 17 pounds; a fine Malay cock 10 pounds; whilst a first-rate Sebright Bantam weighs hardly more than 1 pound. During the last 20 years the size of some of our breeds has been largely increased by methodical selection, whilst that of other breeds has been much diminished. We have already seen how greatly colour varies even within the same breed; we know that the wild G. bankiva varies slightly in colour; we know that colour is variable in all our domestic animals; nevertheless some eminent fanciers have so little faith in variability, that they have actually argued that the chief Game sub-breeds, which differ from each other in nothing but colour, are descended from distinct wild species! Crossing often causes strange modifications of colour. Mr. Tegetmeier informs me that when buff and white Cochins are crossed, some of the {258} chickens are almost invariably black. According to Mr. Brent, black and white Cochins occasionally produce chickens of a slaty-blue tint; and this same tint appears, as Mr. Tegetmeier tells me, from crossing white Cochins with black Spanish fowls, or white Dorkings with black Minorcas.[419] A good observer[420] 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 collars and some spangled feathers with almost obliterated markings;" so that a new variety arose in this singular manner. The skin in the different breeds differs much in colour, being white in common kinds, yellow in Malays and Cochins, and black in Silk fowls; thus mocking, as M. Godron[421] remarks, the three principal types of skin in mankind. The same author adds, that, as different kinds of fowls living in distant and isolated parts of the world have black skin and bones, this colour must have appeared at various times and places.

The shape and carriage of the body and the shape of the head differ much. The beak varies slightly in length and curvature, but incomparably less than with pigeons. In most crested fowls the nostrils offer a remarkable peculiarity in being raised with a crescentic outline. The primary wing-feathers are short in Cochins; in a male, which must have been more than twice as heavy as G. bankiva, these feathers were in both birds of the same length. I have counted, with Mr. Tegetmeier's aid, the primary wing-feathers in thirteen cocks and hens of various breeds; in four of them, namely in two Hamburghs, a Cochin, and Game Bantam, there were 10, instead of the normal number 9; but in counting these feathers I have followed the practice of fanciers, and have not included the first minute primary feather, barely three-quarters of an inch in length. These feathers differ considerably in relative length, the fourth, or the fifth, or the sixth, being the longest; with the third either equal to, or considerably shorter than the fifth. In wild gallinaceous species the relative length and number of the main wing and tail-feathers are extremely constant.

The tail differs much in erectness and size, being small in Malays and very small in Cochins. In thirteen fowls of various breeds which I have examined, five had the normal number of 14 feathers, including in this number the two middle sickle-feathers; six others (viz. a Caffre cock, Gold-spangled Polish cock, Cochin hen, Sultan hen, Game hen, and Malay hen) had 16; {259} and two (an old Cochin cock and Malay hen) had 17 feathers. The rumpless fowl has no tail, and in a bird which I kept alive the oil-gland had aborted; 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.[422] An eminent physiologist[423] has recently spoken of this breed as a distinct species; had he examined the deformed state of the os coccyx he would never have come to this conclusion; he was probably misled by the statement, which may be found in some works, that tailless fowls are wild in Ceylon; but this statement, as I have been assured by Mr. Layard and Dr. Kellaert, who have so closely studied the birds of Ceylon, is utterly false.

The tarsi vary considerably in length, being relatively to the femur considerably longer in the Spanish and Frizzled, and shorter in the Silk and Bantam breeds, than in the wild G. bankiva; but in the latter, as we have seen, the tarsi vary in length. The tarsi are often feathered. The feet in many breeds are furnished with additional toes. Golden-spangled Polish fowls are said[424] to have the skin between their toes much developed; Mr. Tegetmeier observed this in one bird, but it was not so in one which I examined. In Cochins the middle toe is said[425] to be nearly double the length of the lateral toes, and therefore much longer than in G. bankiva or in other fowls; but this was not the case in two which I examined. The nail of the middle toe in this same breed is surprisingly broad and flat, but in a variable degree in two birds which I examined; of this structure in the nail there is only a trace in G. bankiva.

The voice differs slightly, as I am informed by Mr. Dixon, in almost every breed. The Malays[426] have a loud, deep, somewhat prolonged crow, but with considerable individual differences. Colonel Sykes remarks that the domestic Kulm cock in India has not the shrill clear pipe of the English bird, and "his scale of notes appears more limited." Dr. Hooker was struck with the "prolonged howling screech" of the cocks in Sikhim.[427] The crow of the Cochin is notoriously and ludicrously different from that of the common cock. The disposition of the different breeds is widely different, varying from the savage and defiant temper of the Game-cock to the extremely peaceable temper of the Cochin. The latter, it has been asserted, "graze to a much greater extent than any other varieties." The Spanish fowls suffer more from frost than other breeds.

Before we pass on to the skeleton, the degree of distinctness of the several breeds from G. bankiva ought to be noticed. Some {260} writers speak of the Spanish as one of the most distinct breeds, and so it is in general aspect; but its characteristic differences are not important. The Malay appears to me more distinct, from its tall stature, small drooping tail with more than fourteen tail-feathers, and from its small comb and wattles; nevertheless one Malay sub-breed is coloured almost exactly like G. bankiva. Some authors consider the Polish fowl as very distinct; but this is a semi-monstrous breed, as shown by the protuberant and irregularly perforated skull. The Cochin, with its deeply furrowed frontal bones, peculiarly shaped occipital foramen, short wing-feathers, short tail containing more than fourteen feathers, broad nail to the middle toe, fluffy plumage, rough and dark-coloured eggs, and especially from its peculiar voice, is probably the most distinct of all the breeds. If any one of our breeds has descended from some unknown species, distinct from G. bankiva, it is probably the Cochin; but the balance of evidence does not favour this view. All the characteristic differences of the Cochin breed are more or less variable, and may be detected in a greater or lesser degree in other breeds. One sub-breed is coloured closely like G. bankiva. The feathered legs, often furnished with an additional toe, the wings incapable of flight, the extremely quiet disposition, indicate a long course of domestication; and these fowls come from China, where we know that plants and animals have been tended from a remote period with extraordinary care, and where consequently we might expect to find profoundly modified domestic races.

Osteological Differences.—I have examined twenty-seven skeletons and fifty-three skulls of various breeds, including three of G. bankiva: nearly half of these skulls I owe to the kindness of Mr. Tegetmeier, and three of the skeletons to Mr. Eyton.

The Skull differs greatly in size in different breeds, being nearly twice as long in the largest Cochins, but not nearly twice as broad, as in Bantams. The bones at the base, from the occipital foramen to the anterior end (including the quadrates and pterygoids), are absolutely identical in shape in all the skulls. So is the lower jaw. In the forehead slight differences are often perceptible between the males and females, evidently caused by the presence of the comb. In every case I take the skull of G. bankiva as the standard of comparison. In four Games, in one Malay hen, in an {261} African cock, in a Frizzled cock from Madras, in two black-boned Silk hens, no differences occur worth notice. In three Spanish cocks, the form of the forehead between the orbits differs considerably; in one it is considerably depressed, whilst in the two others it is rather prominent, with a deep medial furrow; the skull of the hen is smooth. In three skulls of Sebright Bantams the crown is more globular, and slopes more abruptly to the occiput, than in G. bankiva. In a Bantam or Jumper from Burmah these same characters are more strongly pronounced, and the supra-occiput is more pointed. In a black Bantam the skull is not so globular, and the occipital foramen is very large, and has nearly the same sub-triangular outline presently to be described in Cochins; and in this skull the two ascending branches of the premaxillary are overlapped in a singular manner by the processes of the nasal bone, but, as I have seen only one specimen, some of these differences may be individual. Of Cochins and Brahmas (the latter a crossed race approaching closely to Cochins) I have examined seven skulls; at the point where the ascending branches of the premaxillary rest on the frontal bone the surface is much depressed, and from this depression a deep medial furrow extends backwards to a variable distance; the edges of this fissure are rather prominent, as is the top of the skull behind and over the orbits. These characters are less developed in the hens. The pterygoids, and the processes of the lower jaw, relatively to the size of the head, are broader than in G. bankiva; and this is likewise the case with Dorkings when of large size. The terminal fork of the hyoid bone in Cochins is twice as wide as in G. bankiva, whereas the length of the other hyoid bones is only as three to two. But the most remarkable character is the shape of the occipital foramen: in G. bankiva (A) the breadth in a horizontal line exceeds the height in a vertical line, and the outline is nearly circular; whereas in Cochins (B) the outline is sub-triangular, and the vertical line exceeds the horizontal line in length. This same form likewise occurs in the black Bantam above referred to, and an approach to it may be seen in some Dorkings, and in a slight degree in certain other breeds.



Of Dorkings I have examined three skulls, one belonging to the white sub-breed; the one character deserving notice is the breadth of the frontal bones, which are moderately furrowed in the middle; thus in a skull which was less than once and a half the length of that of G. bankiva, the breadth between the orbits was exactly double. Of Hamburghs I have examined four skulls (male and female) of the pencilled sub-breed, and one (male) of the spangled sub-breed; the nasal bones stand remarkably wide apart, but in a variable degree; consequently narrow membrane-covered spaces fare left between the tips of the two ascending branches of the premaxillary {262} bones, which are rather short, and between these branches and the nasal bones. The surface of the frontal bone, on which the branches of the premaxillary rest, is very little depressed. These peculiarities no doubt stand in close relation with the broad flattened rose-comb characteristic of the Hamburgh breed.



I have examined fourteen skulls of Polish and other crested breeds. Their differences are extraordinary. First for nine skulls of different sub-breeds of English Polish fowls. The hemispherical protuberance of the frontal bones[428] may be seen in the accompanying drawings, in which (B) the skull of a white-crested Polish fowl is shown obliquely from above, with the skull (A) of G. bankiva in the same position. In fig. 35 longitudinal sections are given of the skulls of a Polish fowl, and, for comparison, of a Cochin of the same size. The protuberance in all Polish fowls occupies the same position, but differs much in size. In one of my nine specimens it was extremely slight. The degree to which the protuberance is ossified varies greatly, larger or smaller portions of bone being replaced by membrane. In one specimen there was only a single open pore; generally, there are many variously-shaped open spaces, the bone forming an irregular reticulation. A medial, longitudinal, arched ribbon of bone is generally retained, but in one specimen there was no bone whatever over the whole protuberance, and the skull when cleaned and viewed from above presented the appearance of an open basin. The change in the whole internal form of the skull is surprisingly great. The brain is modified in a corresponding manner, as is shown in the two longitudinal sections, {263} which deserve attentive consideration. The upper and anterior cavity of the three into which the skull may be divided, is the one which is so greatly modified; it is evidently much larger than in the Cochin skull of the same size, and extends much further beyond the interorbital septum, but laterally is less deep. Whether this cavity is entirely filled by the brain, may be doubted. 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 entirely 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 gradation in the configuration of each part of the internal surface could be traced. In the Polish skull, with a small protuberance, the ridge between the anterior and middle cavities was present, but low; and in the Sultan this ridge was replaced by a narrow furrow standing on a broad raised eminence.



{264}

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[429] 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.[430]

To return to the skull. The posterior part, viewed externally, differs little from that of G. bankiva. In most fowls the posterior-lateral process of the frontal bone and the process of the squamosal bone run together and are ossified near their extremities: this union of the two bones, however, is not constant in any breed; and in eleven out of fourteen skulls of crested breeds, these processes were quite distinct. These processes, when not united, instead of being inclined anteriorly as in all common breeds, descend at right angles to the lower jaw; and in this case the longer axis of the bony cavity of the ear is likewise more perpendicular than in other breeds. When the squamosal process is free, instead of expanding at the tip, it is reduced to an extremely fine and pointed style, of variable length. The pterygoid and quadrate bones present no difference. The palatine bones are a little more curved upwards at their posterior ends. The frontal bones, anteriorly to the protuberance, are, as in Dorkings, very broad, but in a variable degree. The nasal bones either stand far apart, as in Hamburghs, or almost touch each other, and in one instance were ossified together. Each nasal bone properly sends out in front two long processes of equal lengths, forming a fork; but in all the Polish skulls, except one, the inner process was considerably, but in a variable degree, shortened and somewhat upturned. In all the skulls, except one, the two ascending branches of the premaxillary, instead of running up between the processes of the nasal bones and resting on the ethmoid bone, are much shortened and terminate in a blunt, somewhat upturned point. In those skulls in which the nasal bones approach quite close to each other or are ossified together, it would be impossible for the ascending branches of the premaxillary to reach the ethmoid and frontal bones; hence we see that even the relative connection of the bones has been changed. Apparently in consequence of the branches of the premaxillary and of the inner processes of the nasal bones being somewhat upturned, the external orifices of the nostrils are upraised and assume a crescentic outline.

I must still say a few words on some of the foreign Crested breeds. The skull of a crested, rumpless, white Turkish fowl is very slightly protuberant, and but little perforated; the ascending branches of the premaxillary {265} are well developed. In another Turkish breed, called Ghoondooks, the skull is considerably protuberant and perforated; the ascending branches of the premaxillary are so much aborted that they project only 1/15th of an inch; and the inner processes of the nasal bone are so completely aborted, that the surface where they should have projected is quite smooth. Here then we see these two bones modified to an extreme degree. Of Sultans (another Turkish breed) I examined two skulls; in that of the female the protuberance was much larger than in the male. In both skulls the ascending branches of the premaxillary were very short, and in both the basal portion of the inner processes of the nasal bones were ossified together. These Sultan skulls differed from those of English Polish fowls in the frontal bones, anteriorly to the protuberance, not being broad.

The last skull which I need describe is a unique one, lent to me by Mr. Tegetmeier: it resembles a Polish skull in most of its characters, but has not the great frontal protuberance; it has, however, two rounded knobs of a different nature, which stand more in front, above the lachrymal bones. These curious knobs, into which the brain does not enter, are separated from each other by a deep medial furrow; and this is perforated by a few minute pores. The nasal bones stand rather wide apart, with their inner processes, and the ascending branches of the premaxillary, upturned and shortened. The two knobs no doubt supported the two great horn-like projections of the comb.



From the foregoing facts we see in how astonishing a manner some of the bones of the skull vary in Crested fowls. The protuberance may certainly be called in one sense a monstrosity, as being wholly unlike anything observed in nature: but as in ordinary cases it is not injurious to the bird, and as it is strictly inherited, it can hardly in another sense be called a monstrosity. A series may be formed commencing with the black-boned Silk fowl, which has a very small crest with the skull beneath penetrated only by a few minute orifices, but with no other change in its structure; and from this first stage we may proceed to fowls with a moderately large crest, which rests, according to Bechstein, on a fleshy mass, but without any {266} protuberance in the skull. I may add that I have seen a similar fleshy or fibrous mass beneath the tuft of feathers on the head of the Tufted duck; and in this case there was no actual protuberance in the skull, but it had become a little more globular. Lastly, when we come to fowls with a largely developed crest, the skull becomes largely protuberant and is perforated by a multitude of irregular open spaces. The close relation between the crest and the size of the bony protuberance is shown in another way; for Mr. Tegetmeier informs me that if chickens lately hatched be selected with a large bony protuberance, when adult they will have a large crest. There can be no doubt that in former times the breeder of Polish fowls attended solely to the crest, and not to the skull; nevertheless, by increasing the crest, in which he has wonderfully succeeded, he has unintentionally made the skull protuberant to an astonishing degree; and through correlation of growth, he has at the same time affected the form and relative connexion of the premaxillary and nasal bones, the shape of the orifice of the nose, the breadth of the frontal bones, the shape of the post-lateral processes of the frontal and squamosal bones, the direction of the axis of the bony cavity of the ear, and lastly the internal configuration of the whole skull together with the shape of the brain.

Vertebrae.—In G. bankiva there are fourteen cervical, seven dorsal with ribs, apparently fifteen lumbar and sacral, and six caudal vertebrae;[431] but the lumbar and sacral are so much anchylosed that I am not sure of their number, and this makes the comparison of the total number of vertebrae in the several breeds difficult. I have spoken of six caudal vertebrae, because the basal one is almost completely anchylosed with the pelvis; but if we consider the number as seven, the caudal vertebrae agree in all the skeletons. The cervical vertebrae are, as just stated, in appearance fourteen; but out of twenty-three skeletons in a fit state for examination, in five of them, namely, in two Games, in two pencilled Hamburghs, and in a Polish, the fourteenth vertebra bore ribs, which, though small, were perfectly developed with a double articulation. The presence of these little ribs cannot be considered as a fact of much importance, for all the cervical vertebrae bear representatives of ribs; but their development in the fourteenth vertebra reduces the size of the passages in the transverse processes, and makes this vertebra exactly like the first dorsal vertebra. The addition of these little ribs does not affect the fourteenth cervical alone, for properly the ribs of the first true dorsal vertebra are destitute of processes; but in some of the skeletons in which the fourteenth cervical bore little ribs, the first pair of true ribs had well-developed processes. When we know that the sparrow has only nine, and the swan twenty-three cervical vertebrae,[432] we need feel no surprise at the number of the cervical vertebrae in the fowl being, as it appears, variable.

There are seven dorsal vertebrae bearing ribs; the first dorsal is never {267} anchylosed with the succeeding four, which are generally anchylosed together. In one Sultan fowl, however, the two first dorsal vertebrae were free. In two skeletons, the fifth dorsal was free; generally the sixth is free (as in G. bankiva), but sometimes only at its posterior end, where in contact with the seventh. The seventh dorsal vertebra, in every case excepting in one Spanish cock, was anchylosed with the lumbar vertebrae. So that the degree to which these middle dorsal vertebrae are anchylosed together is variable.

Seven is the normal number of true ribs, but in two skeletons of the Sultan fowl (in which the fourteenth cervical vertebra was not furnished with little ribs) there were eight pairs; the eighth pair seemed to be developed on a vertebra corresponding with the first lumbar in G. bankiva; the sternal portion of both the seventh and eighth ribs did not reach the sternum. In four skeletons in which ribs were developed on the fourteenth cervical vertebra, there were, when these cervical ribs are included, eight pairs; but in one Game-cock, in which the fourteenth cervical was furnished with ribs, there were only six pairs of true dorsal ribs; the sixth pair in this case did not have processes, and thus resembled the seventh pair in other skeletons; in this game-cock, as far as could be judged from the appearance of the lumbar vertebrae, a whole dorsal vertebra with its ribs was missing. We thus see that the ribs (whether or not the little pair attached to the fourteenth cervical vertebra be counted) vary from six to eight pair. The sixth pair is frequently not furnished with processes. The sternal portion of the seventh pair is extremely broad in Cochins, and is completely ossified. As previously stated, it is scarcely possible to count the lumbo-sacral vertebrae; but they certainly do not correspond in shape or number in the several skeletons. The caudal vertebrae are closely similar in all the skeletons, the only difference being, whether or not the basal one is anchylosed to the pelvis; they hardly vary even in length, not being shorter in Cochins, with their short tail-feathers, than in other breeds; in a Spanish cock, however, the caudal vertebrae were a little elongated. In three rumpless fowls the caudal vertebrae were few in number, and anchylosed together into a misformed mass.



In the individual vertebrae the differences in structure are very slight. In the atlas the cavity for the occipital condyle is either ossified into a ring, or is, as in Bankiva, open on its upper margin. The upper arc of the spinal canal is a little more arched in Cochins, in conformity with the shape of occipital foramen, than in G. bankiva. In several skeletons a difference, but not of much importance, may be observed, which commences a the fourth cervical vertebra, and is greatest at about the sixth, seventh, or eighth vertebra; this consists in the haemal descending processes being united to the body of the vertebra by a sort of buttress. This structure may be observed in Cochins, Polish, some Hamburgh, and probably other breeds; but is absent, or barely developed, in Game, Dorking, Spanish, Bantam, and {268} 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 spine; the outline, however, does differ in being more truncated in Bantams, and more rounded in certain breeds, as in Cochins. The outline of the ischiadic foramen differs considerably, being nearly circular in Bantams, instead of egg-shaped as in the Bankiva, and more regularly oval in some skeletons, as in the Spanish. The obturator notch is also much less elongated in some skeletons than in others. The end of the pubic bone presents the greatest difference; being hardly enlarged in the Bankiva; considerably and gradually enlarged in Cochins, and in a lesser degree in some other breeds; and abruptly enlarged in Bantams. In one Bantam this bone extended very little beyond the extremity of the ischium. The whole pelvis in this latter bird differed widely in its proportions, being far broader proportionally to its length than in Bankiva.



Sternum.—This bone is generally so much deformed that it is scarcely possible to compare its form strictly in the several breeds. The shape of the triangular extremity of the lateral processes differs considerably, being either almost equilateral or much elongated. The front margin of the crest is more or less perpendicular and varies greatly, as does the curvature of the posterior end, and the flatness of the lower surface. The outline of the manubrial process also varies, being wedge-shaped in the Bankiva, and rounded in the Spanish breed. The furcula 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 coracoids present no difference worth notice. The scapula varies in shape, being of nearly uniform breadth in Bankiva, much broader in the middle in the Polish fowl, and abruptly narrowed towards the apex in the two Sultan fowls.

I carefully compared each separate bone of the leg and wing, relatively to the same bones in the wild Bankiva, in the following breeds, which I thought were the most likely to differ; namely, in Cochin, Dorking, {269} Spanish, Polish, Burmese Bantam, Frizzled Indian, and black-boned Silk fowls; and it was truly surprising to see how absolutely every process, articulation, and pore agreed, though the bones differed greatly in size. The agreement is far more absolute than in other parts of the skeleton. In stating this, I do not refer to the relative thickness and length of the several bones; for the tarsi varied considerably in both these respects. But the other limb-bones varied little even in relative length.

Finally, I have not examined a sufficient number of skeletons to say whether any of the foregoing differences, except in the skull, are characteristic of the several breeds. Apparently some differences are more common in certain breeds than in others,—as an additional rib to the fourteenth cervical vertebra in Hamburghs and Games, and the breadth of the end of the pubic bone in Cochins. Both skeletons of the Sultan fowl had eight dorsal vertebrae, and the end of the scapula in both was somewhat attenuated. In the skull, the deep medial furrow in the frontal bones and the vertically elongated occipital foramen seem to be characteristic of Cochins; as is the great breadth of the frontal bones in Dorkings; the separation and open spaces between the tips of the ascending branches of the premaxillaries and nasal bones, as well as the front part of the skull being but little depressed, characterise Hamburghs; the globular shape of the posterior part of the skull seems to be characteristic of laced Bantams; and lastly, the protuberance of the skull with the ascending branches of the premaxillaries partially aborted, together with the other differences before specified, are eminently characteristic of Polish and other Crested fowls.

But the most striking result of our examination of the skeleton is the great variability of all the bones except those of the extremities. To a certain extent we can understand why the skeleton fluctuates so much in structure; fowls have been exposed to unnatural conditions of life, and their whole organisation has thus been rendered variable; but the breeder is quite indifferent to, and never intentionally selects, any modifications 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 points,—fluctuate in our domestic fowls in the same manner as the several parts of the skeleton. An additional toe is a "point" in Dorkings, and has become a fixed character, but is variable in {270} Cochins and Silk-fowls. The colour of the plumage and the form of the comb are in most breeds, or even sub-breeds, eminently fixed characters; but in Dorkings these points have not been attended to, and are variable. When any modification in the skeleton is related to some external character which man values, it has been, unintentionally on his part, acted on by selection, and has become more or less fixed. We see this in the wonderful protuberance of the skull, which supports the crest of feathers in Polish fowls, and which by correlation has affected other parts of the skull. We see the same result in the two protuberances which support the horns in the horned fowl, and in the flattened shape of the front of the skull in Hamburghs consequent on their flattened and broad "rose-combs." We know not in the least whether additional ribs, or the changed outline of the occipital foramen, or the changed form of the scapula, or of the extremity of the furcula, are in any way correlated with other structures, or have arisen from the changed conditions and habits of life to which our fowls have been subjected; but there is no reason to doubt that these various modifications in the skeleton could be rendered, either by direct selection, or by the selection of correlated structures, as constant and as characteristic of each breed, as are the size and shape of the body, the colour of the plumage, and the form of the comb.

Effects of the Disuse of Parts.

Judging from the habits of our European gallinaceous birds, Gallus bankiva in its native haunts would use its legs and wings more than do our domestic fowls, which rarely fly except to their roosts. The Silk and the Frizzled fowls, from having imperfect wing-feathers, cannot fly at all; and there is reason to believe that both these breeds are ancient, so that their progenitors during many generations cannot have flown. The Cochins, also, from their short wings and heavy bodies, can hardly fly up to a low perch. Therefore in these breeds, especially in the two first, a considerable diminution in the wing-bones might have been expected, but this is not the case. In every specimen, after disarticulating and cleaning the bones, I carefully compared the relative length of the two main bones of the wing to each other, and of the two main bones of the leg to each other, with those of G. bankiva; and it was surprising to see (except in the case of the tarsi) how exactly the same relative length had been retained. This fact is curious, from showing how truly the proportions of an organ may be inherited, although not fully exercised during many generations. I then compared in several breeds the {271} length of the femur and tibia with the humerus and ulna, and likewise these same bones with those of G. bankiva; the result was that the wing-bones in all the breeds (except the Burmese Jumper, which has unnaturally short legs) are slightly shortened relatively to the leg-bones; but the decrease is so slight that it may be due to the standard specimen of G. bankiva having accidentally had wings of slightly greater length than usual; so that the measurements are not worth giving. But it deserves notice that the Silk and Frizzled fowls, which are quite incapable of flight, had their wings less reduced relatively to their legs than in almost any other breed! We have seen with domesticated pigeons that the bones of the wings are somewhat reduced in length, whilst the primary feathers are rather increased in length, and it is just possible, though not probable, that in the Silk and Frizzled fowls any tendency to decrease in the length of the wing-bones from disuse may have been checked through the law of compensation, by the decreased growth of the wing-feathers, and consequent increased supply of nutriment. The wing-bones, however, in both these breeds, are found to be slightly reduced in length when judged by the standard of the length of the sternum or head, relatively to these same parts in G. bankiva.

The actual weight of the main bones of the leg and wing in twelve breeds is given in the two first columns in 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.[433]

TABLE I.

+ + -+ -+ + Weight of Actual Actual Wingbones Weight Weight relatively to Names of Breeds. of of the Leg-bones, Femur Humerus in comparison and and with these Tibia. Ulna. same bones in Grains. Grains. G. bankiva. + + -+ -+ -+ + Gallus bankiva wild male 86 54 100 1 Cochin male 311 162 83 2 Dorking male 557 248 70 3 Spanish (Minorca) male 386 183 75 4 Gold Spangled Polish male 306 145 75 5 Game, black-breasted male 293 143 77 6 Malay female 231 116 80 7 Sultan male 189 94 79 8 Indian Frizzled male 206 88 67 9 Burmese Jumper female 53 36 108 10 Hamburgh (pencilled) male 157 104 106 11 Hamburgh (pencilled) female 114 77 108 12 Silk (black-boned) female 88 57 103 + + -+ -+ -+ +

{272}

In the eight first birds, belonging to distinct breeds, in this table, we see a decided reduction in the weight of the bones of the wing. In the Indian Frizzled fowl, which cannot fly, the reduction is carried to the greatest extent, namely, to thirty-three per cent. of their proper proportional weight. In the next four birds, including the Silk-hen, which is incapable of flight, we see that the wings, relatively to the legs, are slightly increased in weight; but it should be observed that, if in these birds the legs had become from any cause reduced in weight, this would give the false appearance of the wings having increased in relative weight. Now a reduction of this nature has certainly occurred with the Burmese Jumper, in which the legs are abnormally short, and in the two Hamburghs and Silk fowl, the legs, though not short, are formed of remarkably thin and light bones. I make these statements, not judging by mere eyesight, but after having calculated the weights of the leg-bones relatively to those of G. bankiva, according to the only two standards of comparison which I could use, namely, the relative lengths of the head and sternum; for I do not know the weight of the body in G. bankiva, which would have been a better standard. According to these standards, the leg-bones in these four fowls are in a marked manner far lighter than in any other breed. It may therefore be concluded that in all cases in which the legs have not been through some unknown cause much reduced in weight, the wing-bones have become reduced in weight relatively to the leg-bones, in comparison with those of G. bankiva. And this reduction of weight may, I apprehend, safely be attributed to disuse.

To make 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.[434] I am indeed inclined to suspect that the leg-bones in the Dorking, No. 2 in the table, are proportionally too heavy; but this bird was a very large one, weighing 7 lb. 2 oz., though very thin. Its leg-bones were more than ten times as heavy as those of the Burmese Jumper! I tried to ascertain the length both of the leg-bones and wing-bones relatively to other parts of the body and skeleton; but the whole organisation in these birds, which have been so long domesticated, has become so variable, that no certain conclusions could be reached. For instance, the legs of the above Dorking cock were nearly three-quarters of an inch too short relatively to the length of the sternum, and more than {273} 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.[435]

TABLE II.

+ -+ -+ + + Depth of Crest, relatively to Length Depth of the length of Names of Breeds. of Crest the Sternum in Sternum. of Sternum. comparison with G. bankiva. + + + -+ + + Inches. Inches. Gallus bankiva male 4.20 1.40 100 1 Cochin male 5.83 1.55 78 2 Dorking male 6.95 1.97 84 3 Spanish male 6.10 1.83 90 4 Polish male 5.07 1.50 87 5 Game male 5.55 1.55 81 6 Malay female 5.10 1.50 87 7 Sultan male 4.47 1.36 90 8 Frizzled hen male 4.25 1.20 84 9 Burmese Jumper female 3.06 0.85 81 10 Hamburgh male 5.08 1.40 81 11 Hamburgh female 4.55 1.26 81 12 Silk fowl female 4.49 1.01 66 + + + -+ + +

By looking to the third column we see that in every case the depth of the crest relatively to the length of the sternum, in comparison with G. bankiva, is diminished, generally between 10 and 20 per cent. But the degree of reduction varies much, partly in consequence of the frequently deformed state of the sternum. In the Silk-fowl, which cannot fly, the crest is 34 per cent. less deep than what it ought to have been. This reduction of the crest in all the breeds probably accounts for the great variability, before referred to, in the curvature of the furcula, and in the shape of its sternal extremity. Medical men believe that the abnormal form of the spine so commonly observed in women of the higher ranks results from the attached muscles not being fully exercised. So it is with our domestic fowls, for they use their pectoral muscles but little, and, out of twenty-five sternums examined by me, three alone were perfectly symmetrical, ten were moderately crooked, and twelve were deformed to an extreme degree.

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 {274} certainly become lighter relatively to the leg-bones in all the breeds in which these latter bones are not unnaturally short or delicate; and that the crest of the sternum, to which the pectoral muscles are attached, has invariably become less prominent, the whole sternum being also extremely liable to deformity. These results we may attribute to the lessened use of the wings.

Correlation of Growth.—I will here sum up the few facts which I have collected on this obscure, but important, subject. In Cochins and Game-fowls there is some relation between the colour of the plumage and the darkness of the egg-shell and even of the yolk. In Sultans the additional sickle-feathers in the tail are apparently related to the general redundancy of the plumage, as shown by the feathered legs, large crest, and beard. In two tailless fowls which I examined the oil-gland was aborted. A large crest of feathers, as Mr. Tegetmeier has remarked, seems always accompanied by a great diminution or almost entire absence of the comb. A large beard is similarly accompanied by diminished or absent wattles. These latter cases apparently come under the law of compensation or balancement of growth. A large beard beneath the lower jaw and a large top-knot on the skull often go together. The comb when of any peculiar shape, as with Horned, Spanish, and Hamburgh fowls, affects in a corresponding manner the underlying skull; and we have seen how wonderfully this is the case with Crested fowls when the crest is largely developed. With the protuberance of the frontal bones the shape of the internal surface of the skull and of the brain is greatly modified. The presence of a crest influences in some unknown way the development of the ascending branches of the premaxillary bone, and of the inner processes of the nasal bones; and likewise the shape of the external orifice of the nostrils. There is a plain and curious correlation between a crest of feathers and the imperfectly ossified condition of the skull. Not only does this hold good with nearly all crested fowls, but likewise with tufted ducks, and as Dr. Guenther informs me with tufted geese in Germany.

Lastly, the feathers composing the crest in male Polish fowls resemble hackles, and differ greatly in shape from those in the crest of the female. The neck, wing-coverts, and loins {275} in the male bird are properly covered with hackles, and it would appear that feathers of this shape have spread by correlation to the head of the male. This little fact is interesting; because, though both sexes of some wild gallinaceous birds have their heads similarly ornamented, yet there is often a difference in the size and shape of feathers forming their crests. Furthermore there is in some cases, as in the male Gold and in the male Amherst pheasants (P. pictus and Amherstiae), a close relation in colour, as well as in structure, between the plumes on the head and on the loins. Hence it would appear that the same law has regulated the state of the feathers on the head and body, both with species living under their natural conditions, and with birds which have varied under domestication.

* * * * *

{276}

CHAPTER VIII.

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

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

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

PEACOCK, ORIGIN OF BLACK-SHOULDERED BREED.

TURKEY, BREEDS OF—CROSSED WITH THE UNITED STATES SPECIES—EFFECTS OF CLIMATE ON.

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

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

I will, as in previous cases, first briefly describe the chief domestic breeds of the duck:—

BREED 1. Common Domestic Duck.—Varies much in colour and in proportions, and differs in instincts and disposition from the wild-duck. There are several sub-breeds:—(1) The Aylesbury, of great size, white, with pale-yellow beak and legs; abdominal sack largely developed. (2) The Rouen, of great size, coloured like the wild-duck, with green or mottled beak; abdominal sack largely developed. (3) Tufted Duck, with a large top-knot of fine downy feathers, supported on a fleshy mass, with the skull perforated beneath. The top-knot in a duck which I imported from Holland was two and a half inches in diameter. (4) Labrador (or Canadian, or Buenos Ayres, or East Indian); plumage entirely black; beak broader, relatively to its length, than in the wild-duck; eggs slightly tinted with black. This sub-breed perhaps ought to be ranked as a breed; it includes two sub-varieties, one as large as the common domestic duck, which I have kept alive, and the other smaller and often capable of flight.[436] I presume it is this latter sub-variety which has been described in France[437] as flying well, being rather wild, and when cooked having the flavour of the wild-duck; nevertheless this sub-variety is polygamous, like other domesticated ducks and unlike the wild duck. These black Labrador ducks breed true; {277} but a case is given by Dr. Turral of the French sub-variety producing young with some white feathers on the head and neck, and with an ochre-coloured patch on the breast.

BREED 2. Hook-billed Duck.—This bird presents an extraordinary appearance from the downward curvature of the beak. The head is often tufted. The common colour is white, but some are coloured like wild-ducks. It is an ancient breed, having been noticed in 1676.[438] It shows its prolonged domestication by almost incessantly laying eggs, like the fowls which are called everlasting layers.[439]

BREED 3. Call-Duck.—Remarkable from its small size, and from the extraordinary loquacity of the female. Beak short. These birds are either white, or coloured like the wild-duck.

BREED 4. Penguin Duck.—This is the most remarkable of all the breeds, and seems to have originated in the Malayan archipelago. It walks with its body extremely erect, and with its thin neck stretched straight upwards. Beak rather short. Tail upturned, including only 18 feathers. Femur and meta-tarsi elongated.

Almost all naturalists admit that the several breeds are descended from the common wild duck (Anas boschas); most fanciers, on the other hand, take as usual a very different view.[440] Unless we deny that domestication, prolonged during centuries, can affect even such unimportant characters as colour, size, and in a slight degree proportional dimensions and mental disposition, there is no reason whatever to doubt that the domestic duck is descended from the common wild species, for the one differs from the other in no important character. We have some historical evidence with respect to the period and progress of the domestication of the duck. It was unknown[441] to the ancient Egyptians, to the Jews of the Old Testament, and to the Greeks of the Homeric period. About eighteen centuries ago Columella[442] and Varro speak of the necessity of keeping ducks in netted enclosures like other wild fowl, so that at this period there was danger of their flying away. {278} Moreover, the plan recommended by Columella to those who might wish to increase their stock of ducks, namely, to collect the eggs of the wild bird and to place them under a hen, shows, as Mr. Dixon remarks, "that the duck had not at this time become a naturalised and prolific inmate of the Roman poultry-yard." The origin of the domestic duck from the wild species is recognised in nearly every language of Europe, as Aldrovandi long ago remarked, by the same name being applied to both. The wild duck has a wide range from the Himalayas to North America. It crosses readily with the domestic bird, and the crossed offspring are perfectly fertile.

Both in North America and Europe the wild duck has been found easy to tame and breed. In Sweden this experiment was carefully tried by Tiburtius; he succeeded in rearing wild ducks for three generations, but, though they were treated like common ducks, they did not vary even in a single feather. The young birds suffered from being allowed to swim about in cold water,[443] as is known to be the case, though the fact is a strange one, with the young of the common domestic duck. An accurate and well-known observer in England[444] has described in detail his often repeated and successful experiments in domesticating the wild duck. Young birds are easily reared from eggs hatched under a bantam; but to succeed it is indispensable not to place the eggs of both the wild and tame duck under the same hen, for in this case "the young wild ducks die off, leaving their more hardy brethren in undisturbed possession of their foster-mother's care. The difference of habit at the onset in the newly-hatched ducklings almost entails such a result to a certainty." The wild ducklings were from the first quite tame towards those who took care of them as long as they wore the same clothes, and likewise to the dogs and cats of the house. They would even snap with their beaks at the dogs, and drive them away from any spot which they coveted. But they were much alarmed at strange men and dogs. Differently from what {279} occurred in Sweden, Mr. Hewitt found that his young birds always changed and deteriorated in character in the course of two or three generations; notwithstanding that great care was taken to prevent any crossing with tame ducks. After the third generation his birds lost the elegant carriage of the wild species, and began to acquire the gait of the common duck. They increased in size in each generation, and their legs became less fine. The white collar round the neck of the mallard became broader and less regular, and some of the longer primary wing-feathers became more or less white. When this occurred, Mr. Hewitt always destroyed his old stock and procured fresh eggs from wild nests; so that he never bred the same family for more than five or six generations. His birds continued to pair together, and never became polygamous like the common domestic duck. I have given these details, because no other case, as far as I know, has been so carefully recorded by a competent observer of the progress of change in wild birds reared for several generations in a domestic condition.

From these considerations there can hardly be a doubt that the wild duck is the parent of the common domestic kind; nor need we look to distinct species for the parentage of the more distinct breeds, namely, Penguin, Call, Hook-billed, Tufted, and Labrador ducks. I will not repeat the arguments used in the previous chapters on the improbability of man having in ancient times domesticated several species since become unknown or extinct, though ducks are not readily exterminated in the wild state;—on some of the supposed parent-species having had abnormal characters in comparison with all the other species of the genus, as with hook-billed and penguin ducks;—on all the breeds, as far as is known, being fertile together;[445]—on all the breeds having the same general disposition, instinct, &c. But one fact bearing on this question may be noticed: in the great duck family, one species alone, namely, the male of {280} A. boschas, has its four middle tail-feathers curled upwardly; now in every one of the above-named domestic breeds these curled feathers exist, and on the supposition that they are descended from distinct species, we must assume that man formerly hit upon species all of which had this now unique character. Moreover, sub-varieties of each breed are coloured almost exactly like the wild duck, as I have seen with the largest and smallest breeds, namely Rouens and Call-ducks, and, as Mr. Brent states,[446] is the case with Hook-billed ducks. This gentleman, as he informs me, crossed a white Aylesbury drake and a black Labrador duck, and some of the ducklings as they grew up assumed the plumage of the wild duck.

With respect to Penguins, I have not seen many specimens, and none were coloured precisely like the wild duck; but Sir James Brooke sent me three skins from Lombok and Bali, in the Malayan archipelago; the two females were paler and more rufous than the wild duck, and the drake differed in having the whole under and upper surface (excepting the neck, tail-coverts, tail, and wings) silver-grey, finely pencilled with dark lines, closely like certain parts of the plumage of the wild mallard. But I found this drake to be identical in every feather with a variety of the common breed procured from a farm-yard in Kent, and I have occasionally elsewhere seen similar specimens. The occurrence of a duck bred under so peculiar a climate as that of the Malayan archipelago, where the wild species does not exist, with exactly the same plumage as may occasionally be seen in our farm-yards, is a fact worth notice. Nevertheless the climate of the Malayan archipelago apparently does tend to cause the duck to vary much, for Zollinger,[447] speaking of the Penguin breed, says that in Lombok "there is an unusual and very wonderful variety of ducks." One Penguin drake which I kept alive differed from those of which the skins were sent me from Lombok, in having its breast and back partially coloured with chestnut-brown, thus more closely resembling the Mallard.

From these several facts, more especially from the drakes of all the breeds having curled tail-feathers, and from certain sub-varieties in each breed occasionally resembling in general {281} plumage the wild duck, we may conclude with confidence that all the breeds are descended from A. boschas.

I will now notice some of the peculiarities characteristic of the several breeds. The eggs vary in colour; some common ducks laying pale-greenish and others quite white eggs. The eggs which are first laid during each season by the black Labrador duck, are tinted black, as if rubbed with ink. So that with ducks, as with poultry, some degree of correlation exists between the colour of the plumage and the egg-shell. A good observer assured me that one year his Labrador ducks laid almost perfectly white eggs, but that the yolks were this same season dirty olive-green, instead of as usual of a golden yellow, so that the black tint appeared to have passed inwards. Another curious case shows what singular variations sometimes occur and are inherited; Mr. Hansell[448] relates that he had a common duck which always laid eggs with the yolk of a dark-brown colour like melted glue; and the young ducks, hatched from these eggs, laid the same kind of eggs, so that the breed had to be destroyed.

The hook-billed duck has a most remarkable appearance (see fig. of skull, woodcut No. 39); and its peculiar beak has been inherited at least since the year 1676. This structure is evidently analogous with that described in the Bagadotten carrier pigeon. Mr. Brent[449] says that, when hook-billed ducks are crossed with common ducks, "many young ones are produced with the upper mandible shorter than the lower, which not unfrequently causes the death of the bird." A tuft of feathers on the head is by no means a rare occurrence; namely, in the true tufted breed, the hook-billed, the common farmyard duck, and in a duck having no other peculiarity which was sent to me from the Malayan archipelago. The tuft is only so far interesting as it affects the skull, which is thus rendered slightly more globular, and is perforated by numerous apertures. Call-ducks are remarkable from their extraordinary loquacity: the drake only hisses like common drakes; nevertheless, when paired with the common duck, he transmits to his female offspring a strong quacking tendency. This loquacity seems at first a surprising character to have been acquired under domestication. But the voice varies in the different breeds; Mr. Brent[450] says that hook-billed ducks are very loquacious, and that Rouens utter a "dull, loud, and monotonous cry, easily distinguishable by an experienced ear." As the loquacity of the Call-duck is highly serviceable, these birds being used in decoys, this quality may have been increased by selection. For instance, Colonel Hawker says, if young wild-ducks cannot be got for a decoy, "by way of make-shift, select tame birds which are the most clamorous, even if their colour should not be like that of wild ones."[451] It has been {282} falsely asserted that Call-ducks hatch their eggs in less time than common ducks.[452]

The Penguin duck is the most remarkable of all the breeds; the thin neck and body are carried erect; the wings are small; the tail is upturned; and the thigh-bones and metatarsi are considerably lengthened in proportion with the same bones in the wild duck. In five specimens examined by me there were only eighteen tail-feathers instead of twenty as in the wild duck; but I have also found only eighteen and nineteen tail-feathers in two Labrador ducks. On the middle toe, in three specimens, there were twenty-seven or twenty-eight scutellae, whereas in two wild ducks there were thirty-one and thirty-two. The Penguin when crossed transmits with much power its peculiar form of body and gait to its offspring; this was manifest with some hybrids raised in the Zoological Gardens between one of these birds and the Egyptian goose[453] (Anser AEgyptiacus), and likewise with some mongrels which I raised between the Penguin and Labrador duck. I am not much surprised that some writers have maintained that this breed must be descended from an unknown and distinct species; but from the reasons already assigned, it seems to me far more probable that it is the descendant, much modified by domestication under an unnatural climate, of Anas boschas.



Osteological Characters.—The skulls of the several breeds differ from each other and from the skull of the wild duck in very little except in the proportional length and curvature of the premaxillaries. These latter bones in the Call-duck are short, and a line drawn from their extremities to the summit of the skull is nearly straight, instead of being concave as in the {283} common duck; so that the skull resembles that of a small goose. In the hook-billed duck (fig. 39) these same bones as well as the lower jaw curve downwards in a most remarkable manner, as represented. In the Labrador duck the premaxillaries are rather broader than in the wild duck; and in two skulls of this breed the vertical ridges on each side of the supra-occipital bone are very prominent. In the Penguin the premaxillaries are relatively shorter than in the wild duck; and the inferior points of the paramastoids more prominent. In a Dutch tufted duck, the skull under the enormous tuft was slightly more globular and was perforated by two large apertures; in this skull the lachrymal bones were produced much further backwards, so as to have a different shape and to nearly touch the post. lat. processes of the frontal bones, thus almost completing the bony orbit of the eye. As the quadrate and pterygoid bones are of such complex shape and stand in relation with so many other bones, I carefully compared them in all the principal breeds; but excepting in size they presented no difference.



Vertebrae and Ribs.—In one skeleton of the Labrador duck there were the usual fifteen cervical vertebrae and the usual nine dorsal vertebrae bearing ribs; in the other skeleton there were fifteen cervical and ten dorsal vertebrae with ribs; nor, as far as could be judged, was this owing merely to a rib having been developed on the first lumbar vertebra; for in both skeletons the lumbar vertebrae agreed perfectly in number, shape, and size with those of the wild duck. In two skeletons of the Call-duck there were fifteen cervical and nine dorsal vertebrae; in a third skeleton small ribs were attached to the so-called fifteenth cervical vertebra, making ten pairs of ribs; but these ten ribs do not correspond, or arise from the same vertebrae, with the ten in the above-mentioned Labrador duck. In the Call-duck, which had small ribs attached to the fifteenth cervical vertebra, the haemal spines of the thirteenth and fourteenth (cervical) and of the seventeenth (dorsal) vertebrae corresponded with the spines on the fourteenth, fifteenth, and eighteenth vertebrae of the wild duck: so that each of these vertebrae had acquired a structure proper to one posterior to it in position. In the twelfth cervical vertebra of this same Call-duck (fig. 40, B), the two branches of the haemal spine stand much closer together than in the wild duck (A), and the descending haemal processes are much shortened. In the Penguin duck the neck from its thinness and erectness falsely appears (as ascertained by measurement) to be much elongated, but the cervical and dorsal vertebrae present no difference; the posterior dorsal vertebrae, however, are more completely anchylosed to {284} the pelvis than in the wild duck. The Aylesbury duck has fifteen cervical and ten dorsal vertebrae furnished with ribs, but the same number of lumbar, sacral, and caudal vertebrae, as far as could be traced, as in the wild duck. The cervical vertebrae in this same duck (fig. 40, D) were much broader and thicker relatively to their length than in the wild (C); so much so, that I have thought it worth while to give a sketch of the eighth cervical vertebra in these two birds. From the foregoing statements we see that the fifteenth cervical vertebra occasionally becomes modified into a dorsal vertebra, and when this occurs all the adjoining vertebrae are modified. We also see that an additional dorsal vertebra bearing a rib is occasionally developed, the number of the cervical and lumbar vertebrae apparently remaining the same as usual.

I examined the bony enlargement of the trachea in the males of the Penguin, Call, Hook-billed, Labrador, and Aylesbury breeds; and in all it was identical in shape.

The Pelvis is remarkably uniform; but in the skeleton of the Hook-billed duck the anterior part is much bowed inwards; in the Aylesbury and some other breeds the ischiadic foramen is less elongated. In the sternum, furcula, coracoids, and scapula, the differences are so slight and so variable as not to be worth notice, except that in two skeletons of the Penguin duck the terminal portion of the scapula was much attenuated.

In the bones of the leg and wing no modification in shape could be observed. But in Penguin and Hook-billed ducks, the terminal phalanges of the wing are a little shortened. In the former, the femur and metatarsus (but not the tibia) are considerably lengthened, relatively to the same bones in the wild duck, and to the wing-bones in both birds. This elongation of the leg-bones could be seen whilst the bird was alive, and is no doubt connected with its peculiar upright manner of walking. In a large Aylesbury duck, on the other hand, the tibia was the only bone of the leg which relatively to the other bones was slightly lengthened.

On the effects of the increased and decreased Use of the Limbs.—In all the breeds the bones of the wing (measured separately after having been cleaned) relatively to those of the leg have become slightly shortened, in comparison with the same bones in the wild duck, as may be seen in the following table:—

+ + -+ -+ -+ Length of Femur, Length of Humerus, Name of Breed. Tibia, and Radius, and Metatarsus Metacarpus Or as together. together. + + -+ -+ -+ Inches. Inches. Wild mallard 7.14 9.28 100 : 129 Aylesbury 8.64 10.43 100 : 120 Tufted (Dutch) 8.25 9.83 100 : 119 Penguin 7.12 8.78 100 : 123 Call 6.20 7.77 100 : 125 + + -+ -+ -+ Length of same Length of all the Bones. Bones of Wing. + -+ -+ Inches. Inches. Wild duck (another specimen) 6.85 10.07 100 : 147 Common domestic duck 8.15 11.26 100 : 138 + + -+ -+ -+

{285}

In the foregoing table we see that, in comparison with the wild duck, the reduction in the length of the bones of the wing, relatively to those of the legs, though slight, is universal. The reduction is least in the Call-duck, which has the power and the habit of frequently flying.

In weight there is a greater relative difference between the bones of the leg and wing, as may be seen in the following table:—

+ + -+ -+ -+ Weight of Femur, Weight of Name of Breed. Tibia, and Humerus, Radius, Metatarsus and Metacarpus Or as + + -+ -+ -+ Grains. Grains. Wild mallard 54 97 100 : 179 Aylesbury 164 204 100 : 124 Hooked-bill 107 160 100 : 149 Tufted (Dutch) 111 148 100 : 133 Penguin 75 90.5 100 : 120 Labrador 141 165 100 : 117 Call 57 93 100 : 163 + + -+ -+ -+ Weight of all the Weight of all the Bones of the Bones of the Leg and Foot. Wing. + -+ -+ Grains. Grains. Wild duck (another specimen) 66 115 100 : 173 Common domestic duck 127 158 100 : 124 + + -+ -+ -+

In these domesticated birds, the considerably lessened weight of the bones of the wing (i.e. on an average, twenty-five per cent. of their proper proportional weight), as well as their slightly lessened length, relatively to the leg-bones, might follow, not from any actual decrease in the wing-bones, but from the increased weight and length of the bones of the legs. The first of the two tables on the next page shows that the leg-bones relatively to the weight of the entire skeleton have really increased in weight; but the second table shows that according to the same standard the wing-bones have also really decreased in weight; so that the relative disproportion shown in the foregoing tables between the wing and leg bones, in comparison with those of the wild duck, is partly due to the increase in weight and length of the leg-bones, and partly to the decrease in weight and length of the wing-bones.

With respect to the two following tables, I may first state that I tested them by taking another skeleton of a wild duck and of a common domestic duck, and by comparing the weight of all the bones of the leg with all those of the wings, and the result was the same. In the first of these tables we see that the leg-bones in each case have increased in actual weight. It might have been expected that, with the increased or decreased weight of the entire skeleton, the leg-bones would have become proportionally heavier or lighter; but their greater weight in all the breeds relatively to the other bones can be accounted for only by these domestic birds having used their legs in walking and standing much more than the wild, for they never fly, and the more artificial breeds rarely swim. In the second {286} table we see, with the exception of one case, a plain reduction in the weight of the bones of the wing, and this no doubt has resulted from their lessened use. The one exceptional case, namely, in one of the Call-ducks, is in truth no exception, for this bird was constantly in the habit of flying about: and I have seen it day after day rise from my grounds, and fly for a long time in circles of more than a mile in diameter. In this Call-duck there is not only no decrease, but an actual increase in the weight of the wing-bones relatively to those of the wild duck; and this probably is consequent on the remarkable lightness and thinness of all the bones of the skeleton.

- - -+ Weight of entire Weight of Name of Breed. Skeleton. Femur, Tibia, (N.B. One and Metatarsus. Or as Metatarsus and Foot was removed from each skeleton, as it had been accidentally lost in two cases.) + - - -+ Grains. Grains. Wild mallard 839 54 1000 : 64 Aylesbury 1925 164 1000 : 85 Tufted (Dutch) 1404 111 1000 : 79 Penguin 871 75 1000 : 86 Call (from Mr. Fox) 717 57 1000 : 79 + - - - Weight of Skeleton Weight of as above. Humerus, Radius and Ulna, and Metacarpus. - -+ Grains. Grains. Wild mallard 839 97 1000 : 115 Aylesbury 1925 204 1000 : 105 Tufted (Dutch) 1404 148 1000 : 105 Penguin 871 90 1000 : 103 Call (from Mr. Baker) 914 100 1000 : 109 Call (from Mr. Fox) 717 92 1000 : 129 + - - -

Lastly, I weighed the furcula, coracoids, and scapula of a wild duck and of a common domestic duck, and I found that their weight, relatively to that of the whole skeleton, was as one hundred in the former to eighty-nine in the latter; this shows that these bones in the domestic duck have been reduced eleven per cent. of their due proportional weight. The prominence of the crest of the sternum, relatively to its length, is also much reduced in all the domestic breeds. These changes have evidently been caused by the lessened use of the wings.

It is well known that several birds, belonging to different Orders, and inhabiting oceanic islands, have their wings greatly reduced in size and are incapable of flight. I suggested in my 'Origin of Species' that, as these birds are not persecuted by any enemies, the reduction of their wings has probably been caused by gradual disuse. Hence, during the earlier stages of the {287} process of reduction, such birds might be expected to resemble in the state of their organs of flight our domesticated ducks. This is the case with the water-hen (Gallinula nesiotis) of Tristan d'Acunha, which "can flutter a little, but obviously uses its legs, and not its wings, as a mode of escape." Now Mr. Sclater[454] finds in this bird that the wings, sternum, and coracoids, are all reduced in length, and the crest of the sternum in depth, in comparison with the same bones in the European water-hen (G. chloropus). On the other hand, the thigh-bones and pelvis are increased in length, the former by four lines, relatively to the same bones in the common water-hen. Hence in the skeleton of this natural species nearly the same changes have occurred, only carried a little further, as with our domestic ducks, and in this latter case I presume no one will dispute that they have resulted from the lessened use of the wings and the increased use of the legs.

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