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The Variation of Animals and Plants Under Domestication, Vol. I.
by Charles Darwin
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Mr. E. Trail stated in 1867, before the Botanical Society of Edinburgh (and has since given me fuller information), that several years ago he cut about sixty blue and white potatoes into halves through the eyes or buds, and then carefully joined them, destroying at the same time the other eyes. Some of these united tubers produced white, and others blue tubers; and it is probable that in these cases the one half alone of the bud grew. Some, however, produced tubers partly white and partly blue; and the tubers from about four or five were regularly mottled with the two colours. in these latter cases we may conclude that a stem had been formed by {396} the union of the bisected buds; and as tubers are produced by the enlargement of subterranean branches arising from the main stem, their mottled colour apparently affords clear evidence of the intimate commingling of the two varieties. I have repeated these experiments on the potato and on the hyacinth on a large scale, but with no success.

The most reliable instance known to me of the formation of a graft-hybrid is one, recorded by Mr. Poynter,[927] who assures me, in a letter of the entire accuracy of the statement, Rosa Devoniensis had been budded some years previously on a white Banksian rose; and from the much enlarged point of junction, whence the Devoniensis and Banksian still continued to grow, a third branch issued, which was neither pure Banksian nor pure Devoniensis, but partook of the character of both; the flowers resembled, but were superior in character to those of the variety called Lamarque (one of the Noisettes), while the shoots were similar in their manner of growth to those of the Banksian rose, with the exception that the longer and more robust shoots were furnished with prickles. This rose was exhibited before the Floral Committee of the Horticultural Society of London. Dr. Lindley examined it, and concluded that it had certainly been produced by the mingling of R. Banksiae with some rose like R. Devoniensis, "for while it was very greatly increased in vigour and in the size of all the parts, the leaves were half-way between a Banksian and Tea-scented rose." It appears that rose-growers were aware that the Banksian rose sometimes affects other roses. Had it not been for this latter statement, it might have been suspected that this new variety was simply due to bud-variation, and that it had occurred by a mere accident at the point of junction between the two old kinds.

To sum up the foregoing facts: the statement that Cytisus adami originated as a graft-hybrid is so precise that it can hardly be rejected, and, as we have just seen, some analogous facts render the statement to a certain extent probable. The peculiar, monstrous condition of the ovules, and the apparently sound condition of the pollen, favour the belief that it is not an ordinary or seminal hybrid. On the other hand, the fact that the same two species, viz. C. laburnum and purpureus, have spontaneously produced hybrids by seed, is a strong argument in support of the belief that C. adami originated in a similar manner. With respect to the extraordinary tendency which this tree exhibits to complete or partial reversion, we have seen that undoubted seminal hybrids and mongrels are similarly liable. On the whole, I am inclined to put trust in M. Adam's statement; and if it should ever be proved true, the same view would probably have {397} to be extended to the Bizzarria and Trifacial oranges and to the apples above described; but more evidence is requisite before the possibility of the production of graft-hybrids can be fully admitted. Although it is at present impossible to arrive at any certain conclusion with respect to the origin of these remarkable trees, the various facts above given appear to me to deserve attention under several points of view, more especially as showing that the power of reversion is inherent in Buds.

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On the direct or immediate action of the Male Element on the Mother Form.—Another remarkable class of facts must be here considered, because they have been supposed to account for some cases of bud-variation: I refer to the direct action of the male element, not in the ordinary way on the ovules, but on certain parts of the female plant, or in the case of animals on the subsequent progeny of the female by a second male. I may premise that with plants the ovarium and the coats of the ovules are obviously parts of the female, and it could not have been anticipated that they would be affected by the pollen of a foreign variety or species, although the development of the embryo, within the embryonic sack, within the ovule, within the ovarium, of course depends on the male element.

Even as long ago as 1729 it was observed[928] that white and blue varieties of the Pea, when planted near each other, mutually crossed, no doubt through the agency of bees, and in the autumn blue and white peas were found within the same pods. Wiegmann made an exactly similar observation in the present century. The same result has followed several times when a variety with peas of one colour has been artificially crossed by a differently-coloured variety.[929] These statements led Gaertner, who was highly sceptical on the subject, carefully to try a long series of experiments: he selected the most constant varieties, and the result conclusively showed that the colour of the skin of the pea is modified when pollen of a differently coloured variety is used. This conclusion has since been confirmed by experiments made by the Rev. J. M. Berkeley.[930]

Mr. Laxton of Stamford, whilst making experiments on peas for the express purpose of ascertaining the influence of foreign pollen on the mother-plant, has recently[931] observed an important additional fact. He fertilised the Tall Sugar pea, which bears very thin green pods, becoming {398} brownish-white when dry, with pollen of the Purple-podded pea, which, as its name expresses, has dark-purple pods with very thick skin, becoming pale reddish-purple when dry. Mr. Laxton has cultivated the tall sugar-pea during twenty years, and has never seen or heard of it producing a purple pod; nevertheless, a flower fertilised by pollen from the purple-pod yielded a pod clouded with purplish-red, which Mr. Laxton kindly gave to me. A space of about two inches in length towards the extremity of the pod, and a smaller space near the stalk, were thus coloured. On comparing the colour with that of the purple-pod, both pods having been first dried and then soaked in water, it was found to be identically the same; and in both the colour was confined to the cells lying immediately beneath the outer skin of the pod. The valves of the crossed pod were also decidedly thicker and stronger than those of the pods of the mother-plant, but this may have been an accidental circumstance, for I know not how far their thickness in the Tall Sugar-pea is a variable character.

The peas of the Tall Sugar-pea, when dry, are pale greenish-brown, thickly covered with dots of dark purple so minute as to be visible only through a lens, and Mr. Laxton has never seen or heard of this variety producing a purple pea; but in the crossed pod one of the peas was of a uniform beautiful violet-purple tint, and a second was irregularly clouded with pale purple. The colour lies in the outer of the two coats which surround the pea. As the peas of the purple-podded variety when dry are of a pale greenish-buff, it would at first appear that this remarkable change of colour in the peas in the crossed pod could not have been caused by the direct action of the pollen of the purple-pod: but when we bear in mind that this latter variety has purple flowers, purple marks on its stipules, and purple pods; and that the Tall sugar-pea likewise has purple flowers and stipules, and microscopically minute purple dots on the peas, we can hardly doubt that the tendency to the production of purple in both parents has in combination modified the colour of the peas in the crossed pod. After having examined these specimens, I crossed the same two varieties, and the peas in one pod, but not the pods themselves, were clouded and tinted with purplish-red in a much more conspicuous manner than the peas in the uncrossed pods produced at the same time by the same plants. I may notice as a caution that Mr. Laxton sent me various other crossed peas slightly, or even greatly, modified in colour; but the change in these cases was due, as had been suspected by Mr. Laxton, to the altered colour of the cotyledons, seen through the transparent coats of the peas; and as the cotyledons are parts of the embryo, these cases are not in any way remarkable.

Turning now to the genus Matthiola. The pollen of one kind of stock sometimes affects the colour of the seeds of another kind, used as the mother-plant. I give the following case the more readily, as Gaertner doubted similar statements with respect to the stock previously made by other observers. A well-known horticulturist, Major Trevor Clarke, informs me[932] that the seeds of the large red-flowered biennial stock {399} (Matthiola annua; Cocardeau of the French) are light brown, and those of the purple branching Queen stock (M. incana) are violet-black; and he found that, when flowers of the red stock were fertilised by pollen from the purple stock, they yielded about fifty per cent. of black seeds. He sent me four pods from a red-flowered plant, two of which had been fertilised by their own pollen, and they included pale brown seed; and two which had been crossed by pollen from the purple kind, and they included seeds all deeply tinged with black. These latter seeds yielded purple-flowered plants like their father; whilst the pale brown seeds yielded normal red-flowered plants; and Major Clarke, by sowing similar seeds, has observed on a greater scale the same result. The evidence in this case of the direct action of the pollen of one species on the colour of the seeds of another species appears to me conclusive.

In the foregoing cases, with the exception of that of the purple-podded pea, the coats of the seeds alone have been affected in colour. We shall now see that the ovarium itself, whether forming a large fleshy fruit or a mere thin envelope, may be modified by foreign pollen, in colour, flavour, texture, size, and shape.

The most remarkable instance, because carefully recorded by highly competent authorities, is one of which I have seen an account in a letter written, in 1867, by M. Naudin to Dr. Hooker. M. Naudin states that he has seen fruit growing on Chamaerops humilis, which had been fertilised by M. Denis with pollen from the Phoenix or date-palm. The fruit or drupe thus produced was twice as large as, and more elongated than, that proper to the Chamaerops; so that it was intermediate in these respects, as well as in texture, between the fruit of the two parents. These hybridised seeds germinated, and produced young plants likewise intermediate in character. This case is the more remarkable as the Chamaerops and Phoenix belong not only to distinct genera, but in the estimation of some botanists to distinct sections of the family.

Gallesio[933] fertilised the flowers of an orange with pollen from the lemon; and one fruit thus produced bore a longitudinal stripe of peel having the colour, flavour, and other characters of the lemon. Mr. Anderson[934] fertilised a green-fleshed melon with pollen from a scarlet-fleshed kind; in two of the fruits "a sensible change was perceptible; and four other fruits were somewhat altered both internally and externally." The seeds of the two first-mentioned fruits produced plants partaking of the good properties of both parents. In the United States, where Cucurbitaceae are largely cultivated, it is the popular belief[935] that the fruit is thus directly affected by foreign pollen; and I have received a similar statement with respect to {400} the cucumber in England. It is known that grapes have been thus affected in colour, size, and shape: in France a pale-coloured grape had its juice tinted by the pollen of the dark-coloured Teinturier; in Germany a variety bore berries which were affected by the pollen of two adjoining kinds; some of the berries being only partially affected or mottled.[936] As long ago as 1751[937] it was observed that, when differently coloured varieties of maize grow near each other, they mutually affect each other's seeds, and this is now a popular belief in the United States. Dr. Savi[938] tried the experiment with care: he sowed yellow and black-seeded maize together, and on the same ear some of the seeds were yellow, some black, and some mottled,[939] the differently coloured seeds being arranged in rows or irregularly. Mr. Sabine states[940] that he has seen the form of the nearly globular seed-capsule of Amaryllis vittata altered by the application of the pollen of another species, of which the capsule has gibbous angles. Mr. J. Anderson Henry[941] crossed Rhododendron Dalhousiae with the pollen of R. Nuttallii, which is one of the largest-flowered and noblest species of the genus. The largest pod produced by the former species, when fertilised with its own pollen, measured 1-2/8 inch in length and 11/2 in girth; whilst three of the pods which had been fertilised by pollen of R. Nuttallii measured 1-5/8 inch in length and no less than 2 inches in girth. Here we see the effect of foreign pollen apparently confined to increasing the size of the ovarium; but we must be cautious in assuming, as the following case shows, that in this instance size has been directly transferred from the male parent to the capsule of the female plant. Mr. Henry fertilised Arabis blepharophylla with pollen of A. Soyeri, and the pods thus produced, of which he was so kind as to send me detailed measurements and sketches, were much larger in all their dimensions than those naturally produced by either the male or female parent-species. In a future chapter we shall see {401} that the organs of vegetation in hybrid plants, independently of the character of either parent, are sometimes developed to a monstrous size; and the increased size of the pods in the foregoing cases may be an analogous fact.

No case of the direct action of the pollen of one variety on another is better authenticated or more remarkable than that of the common apple. The fruit here consists of the lower part of the calyx and of the upper part of the flower-peduncle[942] in a metamorphosed condition, so that the effect of the foreign pollen has extended even beyond the limits of the ovarium. Cases of apples thus affected were recorded by Bradley in the early part of the last century; and other cases are given in old volumes of the Philosophical Transactions;[943] in one of these a Russeting apple and an adjoining kind mutually affected each other's fruit; and in another case a smooth apple affected a rough-coated kind. Another instance has been given[944] of two very different apple-trees growing close to each other, which bore fruit resembling each other, but only on the adjoining branches. It is, however, almost superfluous to adduce these or other cases, after that of the St. Valery apple, which, from the abortion of the stamens, does not produce pollen, but, being annually fertilised by the girls of the neighbourhood with pollen of many kinds, bears fruit, "differing from each other in size, flavour, and colour, but resembling in character the hermaphrodite kinds by which they have been fertilised."[945]

I have now shown, on the authority of several excellent observers, in the case of plants belonging to widely different orders, that the pollen of one species or variety, when applied to a distinct form, occasionally causes the coats of the seeds and the ovarium or fruit, including even in one instance the calyx and upper part of the peduncle of the mother-plant, to become modified. Sometimes the whole of the ovarium or all the seeds are thus affected; sometimes only a certain number of the seeds, as in the case of the pea, or only a part of the ovarium, as with the striped orange, mottled grapes and maize, are thus affected. It must not be supposed that any direct or immediate effect invariably follows the use of foreign pollen: this is far from being the case; nor is it known on what conditions the result depends. Mr. Knight[946] expressly states that he has never seen {402} the fruit thus affected, though he has crossed thousands of apple and other fruit-trees. There is not the least reason to believe that a branch which has borne seed or fruit directly modified by foreign pollen is itself affected, so as subsequently to produce modified buds: such an occurrence, from the temporary connection of the flower with the stem, would be hardly possible. Hence but very few, if any, of the cases of sudden modifications in the fruit of trees, given in the early part of this chapter, can be accounted for by the action of foreign pollen; for such modified fruits have commonly been afterwards propagated by budding or grafting. It is also obvious that changes of colour in the flower which necessarily supervene long before it is ready for fertilisation, and changes in the shape or colour of the leaves, can have no relation to the action of foreign pollen: all such cases must be attributed to simple bud-variation.

The proofs of the action of foreign pollen on the mother-plant have been given in considerable detail, because this action, as we shall see in a future chapter, is of the highest theoretical importance, and because it is in itself a remarkable and apparently anomalous circumstance. That it is remarkable under a physiological point of view is clear, for the male element not only affects, in accordance with its proper function, the germ, but the surrounding tissues of the mother-plant. That the action is anomalous in appearance is true, but hardly so in reality, for apparently it plays the same part in the ordinary fertilisation of many flowers. Gaertner has shown,[947] by gradually increasing the number of pollen-grains until he succeeded in fertilising a Malva, that many grains are expended in the development, or, as he expresses it, in the satiation, of the pistil and ovarium. Again, when one plant is fertilised by a widely distinct species, it often happens that the ovarium is fully and quickly developed without any seeds being formed, or the coats of the seeds are developed without an embryo being formed within. Dr. Hildebrand also has lately shown in a valuable paper[948] that, with several Orchideae, the action of the plant's own {403} pollen is necessary for the development of the ovarium, and that this development takes place not only long before the pollen-tubes have reached the ovules, but even before the placentae and ovules have been formed; so that with these orchids the pollen apparently acts directly on the ovarium. On the other hand, we must not overrate the efficacy of pollen in this respect; for in the case of hybridised plants it might be argued that an embryo had been formed and had affected the surrounding tissues of the mother-plant before it perished at a very early age. Again, it is well known that with many plants the ovarium may be fully developed, though pollen be wholly excluded. And lastly, Mr. Smith, the late Curator at Kew (as I hear through Dr. Hooker), observed the singular fact with an orchid, the Bonatea speciosa, the development of the ovarium could be effected by mechanical irritation of the stigma. Nevertheless, from the number of the pollen-grains expended "in the satiation of the ovarium and pistil,"—from the generality of the formation of the ovarium and seed-coats in sterile hybridised plants,—and from Dr. Hildebrand's observations on orchids, we may admit that in most cases the swelling of the ovarium, and the formation of the seed-coats, are at least aided, if not wholly caused, by the direct action of the pollen, independently of the intervention of the fertilised germ. Therefore, in the previously-given cases we have only to add to our belief in the power of the plant's own pollen on the development of the ovarium and seed-coats, its further power, when applied to a distinct species or variety, of influencing the shape, size, colour, texture, &c., of these same parts.

* * * * *

Turning now to the animal kingdom. If we could imagine the same flower to yield seeds during successive years, then it would not be very surprising that a flower of which the ovarium had been modified by foreign pollen should next year produce, when self-fertilised, offspring modified by the previous male influence. Closely analogous cases have actually occurred with animals. In the case often quoted from Lord Morton,[949] a nearly purely-bred, Arabian, chesnut mare bore a hybrid to a quagga; she was subsequently sent to Sir Gore Ouseley, and produced {404} two colts by a black Arabian horse. These colts were partially dun-coloured, and were striped on the legs more plainly than the real hybrid, or even than the quagga. One of the two colts had its neck and some other parts of its body plainly marked with stripes. Stripes on the body, not to mention those on the legs, and the dun-colour, are extremely rare,—I speak after having long attended to the subject,—with horses of all kinds in Europe, and are unknown in the case of Arabians. But what makes the case still more striking is that the hair of the mane in these colts resembled that of the quagga, being short, stiff, and upright. Hence there can be no doubt that the quagga affected the character of the offspring subsequently begot by the black Arabian horse. With respect to the varieties of our domesticated animals, many similar and well-authenticated facts have been published,[950] and others have been communicated to me, plainly showing the influence of the first male on the progeny subsequently borne by the mother to other males. It will suffice to give a single instance, recorded in the 'Philosophical Transactions,' in a paper following that by Lord Morton: Mr. Giles put a sow of Lord Western's black and white Essex breed to a wild boar of a deep chesnut colour; and the "pigs produced partook in appearance of both boar and sow, but in some the chesnut colour of the boar strongly prevailed." After the boar had long been dead, the sow was put to a boar of her own black and white breed,—a kind which is well known to breed very true and never to show any chesnut colour,—yet from this union the sow produced some young pigs which were plainly marked with the same chesnut tint as in the first litter. Similar cases have so frequently occurred, that careful breeders avoid putting a choice female to an inferior male on account of the injury to her subsequent progeny which may be expected to follow.

{405}

Some physiologists have attempted to account for these remarkable results from a first impregnation by the close attachment and freely intercommunicating blood-vessels between the modified embryo and the mother. But it is a most improbable hypothesis that the mere blood of one individual should affect the reproductive organs of another individual in such a manner as to modify the subsequent offspring. The analogy from the direct action of foreign pollen on the ovarium and seed-coats of the mother-plant strongly supports the belief that the male element acts directly on the reproductive organs of the female, wonderful as is this action, and not through the intervention of the crossed embryo. With birds there is no such close connection between the embryo and mother as in the case of mammals: yet a careful observer, Dr. Chapuis, states[951] that with pigeons the influence of a first male sometimes makes itself perceived in the succeeding broods; but this statement, before it can be fully trusted, requires confirmation.

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Conclusion and Summary of the Chapter.—The facts given in the latter half of this chapter are well worthy of consideration, as they show us in how many extraordinary modes one organic form may lead to the modification of another, and often without the intervention of seminal reproduction. There is ample evidence, as we have just seen, that the male element may either directly affect the structure of the female, or in the case of animals lead to the modification of her offspring. There is a considerable but insufficient body of evidence showing that the tissues of two plants may unite and form a bud having a blended character; or again, that buds inserted into a stock may affect all the buds subsequently produced by this stock. Two embryos, differing from each other and contained in the same seed, may cohere and form a single plant. Offspring from a cross between two species or varieties may in the first or in a succeeding generation revert in various degrees by bud-variation to their parent-forms; and this reversion or segregation of character may affect the whole flower, fruit, or leaf-bud, or only the half or smaller segment, or a single organ. In some cases this segregation of character apparently depends on some {406} incapacity of union rather than on reversion, for the flowers or fruit which are first produced display by segments the characters of both parents. In the Cytisus adami and the Bizzarria orange, whatever their origin may have been, the two parent species occur blended together under the form of a sterile hybrid, or reappear with their characters perfect and their reproductive organs effective; and these trees, retaining the same sportive character, can be propagated by buds. These various facts ought to be well considered by any one who wishes to embrace under a single point of view the various modes of reproduction by gemmation, division, and sexual union, the reparation of lost parts, variation, inheritance, reversion, and other such phenomena. In a chapter towards the close of the following volume I shall attempt to connect these facts together by a provisional hypothesis.

In the early half of this chapter I have given a long list of plants in which through bud-variation, that is, independently of reproduction by seed, the fruit has suddenly become modified in size, colour, flavour, hairiness, shape, and time of maturity; flowers have similarly changed in shape, colour, and doubleness, and greatly in the character of the calyx; young branches or shoots have changed in colour, in bearing spines, and in habit of growth, as in climbing and weeping; leaves have changed in colour, variegation, shape, period of unfolding, and in their arrangement on the axis. Buds of all kinds, whether produced on ordinary branches or on subterranean stems, whether simple or, as in tubers and bulbs, much modified and supplied with a stock of nutriment, are all liable to sudden variations of the same general nature.

In the list, many of the cases are certainly due to reversion to characters not acquired from a cross, but which were formerly present, and have been lost for a longer or shorter period of time;—as when a bud on a variegated plant produces plain leaves, or when variously-coloured flowers on the Chrysanthemum revert to the aboriginal yellow tint. Many other cases included in the list are probably due to the plants being of crossed parentage, and to the buds reverting to one of the two parent-forms. In illustration of the origin of Cytisus adami, several cases were given of partial or complete reversion, both {407} with hybrid and mongrel plants; hence we may suspect that the strong tendency in the Chrysanthemum, for instance, to produce by bud-variation differently-coloured flowers, results from the varieties formerly having been intentionally or accidentally crossed; and that their descendants at the present day still occasionally revert by buds to the colours of the more persistent parent-varieties. This is almost certainly the case with Rollisson's Unique Pelargonium; and so it may be to a large extent with the bud-varieties of the Dahlia and with the "broken colours" of Tulips.

Many cases of bud-variation, however, cannot be attributed to reversion, but to spontaneous variability, such as so commonly occurs with cultivated plants when raised from seed. As a single variety of the Chrysanthemum has produced by buds six other varieties, and as one variety of the gooseberry has borne at the same time four distinct varieties of fruit, it is scarcely possible to believe that all these variations are reversions to former parents. We can hardly believe, as remarked in a previous chapter, that all the many peaches which have yielded nectarine-buds are of crossed parentage. Lastly, in such cases as that of the moss-rose with its peculiar calyx, and of the rose which bears opposite leaves, in that of the Imatophyllum, &c., there is no known natural species or seedling variety, from which the characters in question could have been derived by crossing. We must attribute all such cases to actual variability in the buds. The varieties which have thus arisen cannot be distinguished by any external character from seedlings; this is notoriously the case with the varieties of the Rose, Azalea, and many other plants. It deserves notice that all the plants which have yielded bud-variations have likewise varied greatly by seed.

These plants belong to so many orders that we may infer that almost every plant would be liable to bud-variation if placed under the proper exciting conditions. These conditions, as far as we can judge, mainly depend on long-continued and high cultivation; for almost all the plants in the foregoing lists are perennials, and have been largely propagated in many soils and under different climates, by cuttings, offsets, bulbs, tubers, and especially by budding or grafting. The instances of annuals varying by buds, or producing on the same plant {408} differently coloured flowers, are comparatively rare: Hopkirk[952] has seen this with Convolvulus tricolor; and it is not rare with the Balsam and annual Delphinium. According to Sir R. Schomburgk, plants from the warmer temperate regions, when cultivated under the hot climate of St. Domingo, are eminently liable to bud-variation; but change of climate is by no means a necessary contingent, as we see with the gooseberry, currant, and some others. Plants living under their natural conditions are very rarely subject to bud-variation: variegated and coloured leaves have, however, been occasionally observed; and I have given an instance of the variation of buds on an ash-tree; but it is doubtful whether any tree planted in ornamental grounds can be considered as living under strictly natural conditions. Gaertner has seen white and dark-red flowers produced from the same root of the wild Achillea millefolium; and Prof. Caspary has seen Viola lutea, in a completely wild condition, bearing flowers of different colours and sizes.[953]

As wild plants are so rarely liable to bud-variation, whilst highly cultivated plants long propagated by artificial means have yielded by this form of reproduction many varieties, we are led through a series such as the following,—namely, all the eyes in the same tuber of the potato varying in the same manner,—all the fruit on a purple plum-tree suddenly becoming yellow,—all the fruit on a double-flowered almond suddenly becoming peach-like,—all the buds on grafted trees being in some very slight degree affected by the stock on which they have been worked,—all the flowers on a transplanted heartsease changing for a time in colour, size, and shape,—we are led through such facts to look at every case of bud-variation as the direct result of the particular conditions of life to which the plant has been exposed. But if we turn to the other end of the series, namely, to such cases as that of a peach-tree which, after having been cultivated by tens of thousands during many years in many countries, and after having annually produced thousands of buds, all of which have apparently been exposed to precisely the same conditions, yet at last suddenly produces a single bud with its whole character greatly transformed, we are driven to an opposite {409} conclusion. In such cases as the latter it would appear that the transformation stands in no direct relation to the conditions of life.

We have seen that varieties produced from seeds and from buds resemble each other so closely in general appearance, that they cannot possibly be distinguished. Just as certain species and groups of species, when propagated by seed, are more variable than other species or genera, so it is in the case of certain bud-varieties. Thus the Queen of England Chrysanthemum has produced by this latter process no less than six, and Rollisson's Unique Pelargonium four distinct varieties; moss-roses have also produced several other moss-roses. The Rosaceae have varied by buds more than any other group of plants; but this may be in large part due to so many members having been long cultivated; but within this one group, the peach has often varied by buds, whilst the apple and pear, both grafted trees extensively cultivated, have afforded, as far as I can ascertain, extremely few instances of bud-variation.

The law of analogous variation holds good with varieties produced by buds, as with those produced from seed: more than one kind of rose has sported into a moss-rose; more than one kind of camellia has assumed an hexagonal form; and at least seven or eight varieties of the peach have produced nectarines.

The laws of inheritance seem to be nearly the same with seminal and bud-varieties. We know how commonly reversion comes into play with both, and it may affect the whole, or only segments, of a leaf, flower, or fruit. When the tendency to reversion affects many buds on the same tree, it becomes covered with different kinds of leaves, flowers, or fruit; but there is reason to believe that such fluctuating varieties have generally arisen from seed. It is well known that, out of a number of seedling varieties, some transmit their character much more truly by seed than others; so with bud-varieties some retain their character by successive buds more truly than others; of which instances have been given with two kinds of variegated Euonymus and with certain kinds of tulips. Notwithstanding the sudden production of bud-varieties, the characters thus acquired are sometimes capable of transmission by seminal reproduction: Mr. Rivers has found that moss-roses generally {410} reproduce themselves by seed; and the mossy character has been transferred by crossing, from one species of rose to another. The Boston nectarine, which appeared as a bud-variation, produced by seed a closely allied nectarine. We have however seen, on the authority of Mr. Salter, that seed taken from a branch with leaves variegated through bud-variation, transmits this character very feebly; whilst many plants, which became variegated as seedlings, transmit variegation to a large proportion of their progeny.

Although I have been able to collect a good many cases of bud-variation, as shown in the previous lists, and might probably, by searching foreign horticultural works, have collected more cases, yet their total number is as nothing in comparison with that of seminal varieties. With seedlings raised from the more variable cultivated plants, the variations are almost infinitely numerous, but their differences are generally slight: only at long intervals of time a strongly marked modification appears. On the other hand, it is a singular and inexplicable fact that, when plants vary by buds, the variations, though they occur with comparative rarity, are often, or even generally, strongly pronounced. It struck me that this might perhaps be a delusion, and that slight changes often occurred in buds, but from being of no value were overlooked or not recorded. Accordingly I applied to two great authorities on this subject, namely, to Mr. Rivers with respect to fruit-trees, and to Mr. Salter with respect to flowers. Mr. Rivers is doubtful, but does not remember having noticed very slight variations in fruit-buds. Mr. Salter informs me that with flowers such do occur, but, if propagated, they generally lose their new character in the following year; yet he concurs with me that bud-variations usually at once assume a decided and permanent character. We can hardly doubt that this is the rule, when we reflect on such cases as that of the peach, which has been so carefully observed and of which such trifling seminal varieties have been propagated, yet this tree has repeatedly produced by bud-variation nectarines, and only twice (as far as I can learn) any other variety, namely, the Early and Late Grosse Mignonne peaches; and these differ from the parent-tree in hardly any character except the period of maturity. {411}

To my surprise I hear from Mr. Salter that he brings the great principle of selection to bear on variegated plants propagated by buds, and has thus greatly improved and fixed several varieties. He informs me that at first a branch often produces variegated leaves on one side alone, and that the leaves are marked only with an irregular edging or with a few lines of white and yellow. To improve and fix such varieties, he finds it necessary to encourage the buds at the bases of the most distinctly marked leaves, and to propagate from them alone. By following with perseverance this plan during three or four successive seasons, a distinct and fixed variety can generally be secured.

Finally, the facts given in this chapter prove in how close and remarkable a manner the germ of a fertilised seed and the small cellular mass forming a bud resemble each other in function,—in their powers of inheritance with occasional reversion,—and in their capacity for variation of the same general nature, in obedience to the same laws. This resemblance, or rather identity, is rendered far more striking if the facts can be trusted which apparently render it probable that the cellular tissue of one species or variety, when budded or grafted on another, may give rise to a bud having an intermediate character. In this chapter we clearly see that variability is not necessarily contingent on sexual generation, though much more frequently its concomitant than on bud-reproduction. We see that bud-variability is not solely dependent on reversion or atavism to long-lost characters, or to those formerly acquired from a cross, but that it is often spontaneous. But when we ask ourselves what is the cause of any particular bud-variation, we are lost in doubt, being driven in some cases to look to the direct action of the external conditions of life as sufficient, and in other cases to feel a profound conviction that these have played a quite subordinate part, of not more importance than the nature of the spark which ignites a mass of combustible matter.

END OF VOL. I.

LONDON: PRINTED BY W. CLOWES AND SONS, STAMFORD STREET, AND CHARING CROSS.

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NOTES

[1] To any one who has attentively read my 'Origin of Species' this Introduction will be superfluous. As I stated that work that I should soon publish the facts on which the conclusions given in it were founded, I here beg permission to remark that the great delay in publishing this first work has been caused by continued ill-health.

[2] M. Pouchet has recently ('Plurality of Races,' Eng. Translat., 1864, p. 83, &c.) insisted that variation under domestication throws no light on the natural modification of species. I cannot perceive the force of his arguments, or, to speak more accurately, of his assertions to this effect.

[3] Leon Dufour in 'Annales des Scienc. Nat.' (3rd series, Zoolog.), tom. v. p. 6.

[4] In treating the several subjects included in the present and succeeding works I have continually been led to ask for information from many zoologists, botanists, geologists, breeders of animals, and horticulturists, and I have invariably received from them the most generous assistance. Without such aid I could have effected little. I have repeatedly applied for information and specimens to foreigners, and to British merchants and officers of the Government residing in distant lands, and, with the rarest exceptions, I have received prompt, open-handed, and valuable assistance. I cannot express too strongly my obligations to the many persons who have assisted me, and who, I am convinced, would be equally willing to assist others in any scientific investigation.

[5] Owen, 'British Fossil Mammals,' p. 123 to 133. Pictet's 'Traite de Pal.,' 1853, tom. i. p. 202. De Blainville, in his 'Osteographie, Canidae,' p. 142, has largely discussed the whole subject, and concludes that the extinct parent of all domesticated dogs came nearest to the wolf in organization, and to the jackal in habits.

[6] Pallas, I believe, originated this doctrine in 'Act. Acad. St. Petersburgh,' 1780, Part ii. Ehrenberg has advocated it, as may be seen in De Blainville's 'Osteographie,' p. 79. It has been carried to an extreme extent by Col. Hamilton Smith in the 'Naturalist Library,' vol. ix. and x. Mr. W. C. Martin adopts it in his excellent 'History of the Dog,' 1845; as does Dr. Morton, as well as Nott and Gliddon, in the United States. Prof. Low, in his 'Domesticated Animals,' 1845, p. 666, comes to this same conclusion. No one has argued on this side with more clearness and force than the late James Wilson, of Edinburgh, in various papers read before the Highland Agricultural and Wernerian Societies. Isidore Geoffroy Saint Hilaire ('Hist. Nat. Gen.,' 1860, tom. iii. 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), referring to the view that all the domestic races are the modified descendants of a single species, after a long discussion, says, "Cette opinion est, suivant nous du moins, la moins probable."

[7] Berjeau, 'The Varieties of the Dog; in old Sculptures and Pictures,' 1863. 'Der Hund,' von Dr. F. L. Walther, s. 48, Giessen, 1817: this author seems carefully to have studied all classical works on the subject. See also 'Volz, Beitraege zur Kultur-geschichte,' Leipzig, 1852, s. 115. 'Youatt on the Dog,' 1845, p. 6. A very full history is given by De Blainville in his 'Osteographie, Canidae.'

[8] I have seen drawings of this dog from the tomb of the son of Esar Haddon, and clay models in the British Museum. Nott and Gliddon, in their 'Types of Mankind,' 1854, p. 393, give a copy of these drawings. This dog has been called a Thibetan mastiff, but Mr. H. A. Oldfield, who is familiar with the so-called Thibet mastiff, and has examined the drawings in the British Museum, informs me that he considers them different.

[9] 'Proc. Zoolog. Soc.,' July 12th, 1831.

[10] 'Sporting in Algeria,' p. 51.

[11] Berjeau gives fac-similes of the Egyptian drawings. Mr. C. L. Martin, in his 'History of the Dog,' 1845, copies several figures from the Egyptian monuments, and speaks with much confidence with respect to their identity with still living dogs. Messrs. Nott and Gliddon ('Types of Mankind,' 1854, p. 388) give still more numerous figures. Mr. Gliddon asserts that a curl-tailed greyhound, like that represented on the most ancient monuments, is common in Borneo; but the Rajah, Sir J. Brooke, informs me that no such dog exists there.

[12] These, and the following facts on the Danish remains, are taken from M. Morlot's most interesting memoir in 'Soc. Vaudoise des Sc. Nat.,' tom. vi., 1860, pp. 281, 299, 320.

[13] 'Die Fauna der Pfahlbauten,' 1861, s. 117, 162.

[14] De Blainville, 'Osteographie, Canidae.'

[15] Sir R. Schomburgk has given me information on this head. See also 'Journal of R. Geograph. Soc.,' vol. xiii., 1843, p. 65.

[16] 'Domestication of Animals:' Ethnological Soc., Dec. 22nd, 1863.

[17] 'Journal of Researches,' &c., 1845, p. 393. With respect to Canis antarcticus, see p. 193. For the case of the antelope, see 'Journal Royal Geograph. Soc.,' vol. xxiii. p. 94.

[18] The authorities for the foregoing statements are as follow:—Richardson, in 'Fauna Boreali-Americana,' 1829, pp. 64, 75; Dr. Kane, 'Arctic Explorations,' 1856, vol. i. pp. 398, 455; Dr. Hayes, 'Arctic Boat Journey,' 1860, p. 167. Franklin's 'Narrative,' vol. i. p. 269, gives the case of three whelps of a black wolf being carried away by the Indians. Parry, Richardson, and others, give accounts of wolves and dogs naturally crossing in the eastern parts of North America. Seeman, in his 'Voyage of H.M.S. Herald,' 1853, vol. ii. p. 26, says the wolf is often caught by the Esqimaux for the purpose of crossing with their dogs, and thus adding to their size and strength. M. Lamare-Picquot, in 'Bull. de la Soc. d'Acclimat.,' tom. vii., 1860, p. 148, gives a good account of the half-bred Esquimaux dogs.

[19] 'Fauna Boreali-Americana,' 1829, pp. 73, 78, 80. Nott and Gliddon, 'Types of Mankind,' p. 383. The naturalist and traveller Bartram is quoted by Hamilton Smith, in 'Nat. Hist. Lib.,' vol. x. p. 156. A Mexican domestic dog seems also to resemble a wild dog of the same country; but this may be the prairie-wolf. Another capable judge, Mr. J. K. Lord ('The Naturalist in Vancouver Island,' 1866, vol. ii. p. 218), says that the Indian dog of the Spokans, near the Rocky Mountains, "is beyond all question nothing more than a tamed Cayote or prairie-wolf," or Canis latrans.

[20] I quote this from Mr. R. Hill's excellent account of the Alco or domestic dog of Mexico, in Gosse's 'Naturalist's Sojourn in Jamaica,' 1851, p. 329.

[21] 'Naturgeschichte der Saeugethiere von Paraguay,' 1830, s. 151.

[22] Quoted in Humboldt's 'Aspects of Nature' (Eng. transl.), vol. i. p. 108.

[23] Paget's 'Travels in Hungary and Transylvania,' vol. i. p. 501. Jeitteles, 'Fauna Hungariae Superioris,' 1862, s. 13. See Pliny, 'Hist. of the World' (Eng. transl.), 8th book, ch. xl., about the Gauls crossing their dogs. See also 'Hist. Animal.' lib. viii. c. 28. For good evidence about wolves and dogs naturally crossing near the Pyrenees, see M. Mauduyt, 'Du Loup et de ses Races,' Poitiers, 1851; also Pallas, in 'Acta Acad. St. Petersburgh,' 1780, part ii. p. 94.

[24] I give this on excellent authority, namely, Mr. Blyth (under the signature of Zoophilus), in the 'Indian Sporting Review,' Oct. 1856, p. 134. Mr. Blyth states that he was struck with the resemblance between a brush-tailed race of pariah-dogs, north-west of Cawnpore, and the Indian wolf. He gives corroborative evidence with respect to the dogs of the valley of the Nerbudda.

[25] For numerous and interesting details on the resemblance of dogs and jackals, see Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' 1860, tom. iii. p. 101. See also 'Hist. Nat. des Mammiferes,' par Prof. Gervais, 1855, tom. ii. p. 60.

[26] Gueldenstaedt, 'Nov. Comment. Acad. Petrop.,' tom. xx., pro anno 1775, p. 449.

[27] Quoted by De Blainville in his 'Osteographie, Canidae,' pp. 79, 98.

[28] See Pallas, in 'Act. Acad. St. Petersburgh,' 1780, part ii. p. 91. For Algeria, see Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 177. In both countries it is the male jackal which pairs with female domestic dogs.

[29] John Barbut's 'Description of the Coast of Guinea in 1746.'

[30] 'Travels in South Africa,' vol. ii. p. 272.

[31] Selwyn, Geology of Victoria; 'Journal of Geolog. Soc.,' vol. xiv., 1858, p. 536, and vol. xvi., 1860, p. 148; and Prof M^cCoy, in 'Annals and Mag. of Nat. Hist.' (3rd series), vol. ix., 1862, p. 147. The Dingo differs from the dogs of the central Polynesian islands. Dieffenbach remarks ('Travels,' vol. ii. p. 45) that the native New Zealand dog also differs from the Dingo.

[32] 'Proceedings Zoolog. Soc.,' 1833, p. 112. See, also, on the taming of the common wolf, L. Lloyd, 'Scandinavian Adventures,' vol. i. p. 460, 1854. With respect to the jackal, see Prof. Gervais, 'Hist. Nat. Mamm.,' tom. ii. p. 61. With respect to the aguara of Paraguay, see Rengger's work.

[33] Roulin, in 'Mem. present. par divers Savans,' tom. vi. p. 341.

[34] Martin, 'History of the Dog,' p. 14.

[35] Quoted by L. Lloyd in 'Field Sports of North of Europe,' vol. i. p. 387.

[36] Quatrefages, 'Soc. d'Acclimat.,' May 11th, 1863, p. 7.

[37] 'Annals and Mag. of Nat. Hist.,' vol. xv., 1845, p. 140.

[38] Azara, 'Voyages dans l'Amer. Merid.,' tom. i. p. 381; his account is fully confirmed by Rengger. Quatrefages gives an account of a bitch brought from Jerusalem to France which burrowed a hole and littered in it. See 'Discours, Exposition des Races Canines,' 1865, p. 3.

[39] With respect to wolves burrowing holes, see Richardson, Fauna Boreali-Americana,' p. 64; and Bechstein, 'Naturgesch. Deutschlands,' b. i. s. 617.

[40] See Poeppig, 'Reise in Chile,' b. i. s. 290; Mr. G. Clarke, as above; and Rengger, s. 155.

[41] Dogs, 'Nat. Library,' vol. x. p. 121: an endemic South American dog seems also to have become feral in this island. See Gosse's 'Jamaica,' p. 340.

[42] Low, 'Domesticated Animals,' p. 650.

[43] 'The Naturalist Library,' Dogs, vol. x. pp. 4, 19.

[44] Quoted by Prof. Gervais, 'Hist. Nat. Mamm.,' tom. ii. p. 66.

[45] J. Hunter shows that the long period of seventy-three days given by Buffon is easily explained by the bitch having received the dog many times during a period of sixteen days ('Phil. Transact.,' 1787, p. 253). Hunter found that the gestation of a mongrel from wolf and dog ('Phil. Transact.,' 1759, p. 160) apparently was sixty-three days, for she received the dog more than once. The period of a mongrel dog and jackal was fifty-nine days. Fred. Cuvier found the period of gestation of the wolf to be ('Dict. Class. d'Hist. Nat.,' tom. iv. p. 8) two months and a few days, which agrees with the dog. Isid. G. St. Hilaire, who has discussed the whole subject, and from whom I quote Bellingeri, states ('Hist. Nat. Gen.,' tom. iii. p. 112) that in the Jardin des Plantes the period of the jackal has been found to be from sixty to sixty-three days, exactly as with the dog.

[46] See Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 112, on the odour of jackals. Col. Ham. Smith, in 'Nat. Hist. Lib.,' vol. x. p. 289.

[47] Quoted by Quatrefages in 'Bull. Soc. d'Acclimat.,' May 11th, 1863.

[48] 'Journal de la Physiologie,' tom. ii. p. 385.

[49] See Mr. R. Hill's excellent account of this breed in Gosse's 'Jamaica,' p. 338; Rengger's 'Saeugethiere von Paraguay,' s. 153. With respect to Spitz dogs, see Bechstein's 'Naturgesch. Deutschlands,' 1801, b. i. s. 638. With respect to Dr. Hodgkin's statement made before Brit. Assoc., see 'The Zoologist,' vol. iv., for 1845-46, p. 1097.

[50] 'Acta Acad. St. Petersburgh,' 1780, part ii. pp. 84, 100.

[51] M. Broca has shown ('Journal de Physiologie,' tom. ii. p. 353) that Buffon's experiments have been often misrepresented. Broca has collected (pp. 390-395) many facts on the fertility of crossed dogs, wolves, and jackals.

[52] 'De la Longevite Humaine,' par M. Flourens, 1855, p. 143. Mr. Blyth says ('Indian Sporting Review,' vol. ii. p. 137) that he has seen in India several hybrids from the pariah-dog and jackal; and between one of these hybrids and a terrier. The experiments of Hunter on the jackal are well known. See also Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii, p. 217, who speaks of the hybrid offspring of the jackal as perfectly fertile for three generations.

[53] On authority of F. Cuvier, quoted in Bronn's 'Geschichte der Natur,' B. ii. s. 164.

[54] W. C. L. Martin, 'History of the Dog,' 1845, p. 203. Mr. Philip P. King, after ample opportunities of observation, informs me that the Dingo and European dogs often cross in Australia.

[55] Rueppel, 'Neue Wirbelthiere von Abyssinien,' 1835-40; 'Mammif.,' s. 39, pl. xiv. There is a specimen of this fine animal in the British Museum.

[56] Even Pallas admits this: see 'Act. Acad. St. Petersburgh,' 1780, p. 93.

[57] Quoted by I. Geoffroy, 'Hist. Nat. Gen.,' tom. iii. p. 453.

[58] F. Cuvier, in 'Annales du Museum,' tom. xviii. p. 337; Godron, 'De l'Espece,' tom. i. p. 342; and Col. Ham. Smith, in 'Naturalist's Library,' vol. ix. p. 101.

[59] Isid. Geoffroy Saint Hilaire, 'Hist. des Anomalies,' 1832, tom. i. p. 660. Gervais, 'Hist. Nat. des Mammiferes,' tom. ii., 1855, p. 66. De Blainville ('Osteographie, Canidae,' p. 137) has also seen an extra molar on both sides.

[60] 'Osteographie, Canidae,' p. 137.

[61] Wuerzburger, 'Medecin, Zeitschrift,' 1860, B. i. s. 265.

[62] Mr. Yarell, in 'Proc. Zoolog. Soc.,' Oct. 8th, 1833. Mr. Waterhouse showed me a skull of one of these dogs, which had only a single molar on each side and some imperfect incisors.

[63] Quoted in 'The Veterinary,' London, vol. viii. p. 415.

[64] 'Hist Nat. General,' tom. iii. p. 448.

[65] W. Scrope, 'Art of Deer-Stalking,' p. 354.

[66] Quoted by Col. Ham. Smith in 'Naturalist's Library,' vol. x. p. 79.

[67] De Blainville, 'Osteographie, Canidae,' p. 134. F. Cuvier, 'Annales du Museum,' tom. xviii. p. 342. In regard to mastiffs, see Col. Ham. Smith, 'Nat Lib.,' vol. x. p. 218. For the Thibet mastiff, see Mr. Hodgson in 'Journal of As. Soc. of Bengal,' vol. i., 1832, p. 342.

[68] 'The Dog,' 1845, p. 186. With respect to diseases, Youatt asserts (p. 167) that the Italian greyhound is "strongly subject" to polypi in the matrix or vagina. The spaniel and pug (p. 182) are most liable to bronchocele. The liability to distemper (p. 232) is extremely different in different breeds. On the distemper, see also Col. Hutchinson on 'Dog Breaking,' 1850, p. 279.

[69] See Youatt on the Dog, p. 15; 'The Veterinary,' London, vol. xi. p. 235.

[70] 'Journal of As. Soc. of Bengal,' vol. iii. p. 19.

[71] 'Travels,' vol. ii. p. 15.

[72] Hodgson, in 'Journal of As. Soc. of Bengal,' vol. i. p. 342.

[73] 'Field Sports of the North of Europe,' vol. ii. p. 165.

[74] 'Hist. Nat. des Mammif., 1855, tom. ii. pp. 66, 67.

[75] 'History of Quadrupeds,' 1793, vol. i. p. 238.

[76] 'Oriental Field Sports,' quoted by Youatt, 'The Dog,' p. 15.

[77] Quoted by Mr. Galton, 'Domestication of Animals,' p. 13.

[78] 'Hist. Nat. Gen.,' tom. iii. p. 450.

[79] Mr. Greenhow on the Canadian Dog, in Loudon's 'Mag. of Nat. Hist.,' vol. vi., 1833, p. 511.

[80] See Mr. C. O. Groom-Napier on the webbing of the hind feet of Otter-hounds, in 'Land and Water,' Oct. 13th, 1866, p. 270.

[81] 'Fauna Boreali-Americana,' 1829, p. 62.

[82] 'The Horse in all his Varieties,' &c., 1829, pp. 230, 234.

[83] 'The Dog,' 1845, pp. 31, 35; with respect to King Charles's spaniel, p. 45; for the setter, p. 90.

[84] In the 'Encyclop. of Rural Sports,' p. 557.

[85] 'The Farrier,' 1828, vol. i. p. 337.

[86] See Col. Hamilton Smith on the antiquity of the Pointer, in 'Naturalist's Library,' vol. x. p. 195.

[87] The Newfoundland dog is believed to have originated from a cross between the Esquimaux dog and a large French hound. See Dr. Hodgkin, 'Brit. Assoc.,' 1844; Bechstein's 'Naturgesch. Deutschlands,' Band i. s. 574; 'Naturalist's Library,' vol. x. p. 132; also Mr. Jukes' 'Excursion in and about Newfoundland.'

[88] De Blainville, 'Osteographie, Felis,' p. 65, on the character of F. caligulata; pp. 85, 89, 90, 175, on the other mummied species. He quotes Ehrenberg on F. maniculata being mummied.

[89] Asiatic Soc. of Calcutta; Curator's Report, Aug. 1856. The passage from Sir W. Jardine is quoted from this Report. Mr. Blyth, who has especially attended to the wild and domestic cats of India, has given in this Report a very interesting discussion on their origin.

[90] 'Fauna Hungariae Sup.,' 1862, s. 12.

[91] Isid. Geoffrey Saint Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 177.

[92] 'Proc. Zoolog. Soc.,' 1863, p. 184.

[93] 'Saeugethiere von Paraguay,' 1830, s. 212.

[94] 'Mem. presentes par divers Savans: Acad. Roy. des Sciences,' tom. vi. p. 346. Gomara first noticed this fact in 1554.

[95] 'Narrative of Voyages,' vol. ii. p. 180.

[96] J. Crawfurd, 'Descript. Dict. of the Indian Islands,' p. 255. The Madagascar cat is said to have a twisted tail: see Desmarest, in 'Encyclop. Nat. Mamm.,' 1820, p. 233, for some of the other breeds.

[97] Admiral Lutke's Voyage, vol. iii. p. 308.

[98] 'Zoology of the Voyage of the Beagle, Mammalia,' p. 20. Dieffenbach, 'Travels in New Zealand,' vol. ii. p. 185. Ch. St. John, 'Wild Sports of the Highlands,' 1846, p. 49.

[99] Quoted by Isid. Geoffroy, 'Hist. Nat. Gen.,' tom. iii. p. 427.

[100] Ruetimeyer, 'Fauna der Pfalbauten,' 1861, s. 122.

[101] See Youatt on the Horse: J. Lawrence on the Horse, 1829: W. C. L. Martin, 'History of the Horse,' 1845: Col. Ham. Smith, in 'Naturalist's Library, Horses,' 1841, vol. xii.: Prof. Veith, 'Die Naturgesch. Haussaeugethiere,' 1856.

[102] Crawfurd, 'Descript. Dict. of Indian Islands,' 1856, p. 153. "There are many different breeds, every island having at least one peculiar to it." Thus in Sumatra there are at least two breeds; in Achin and Batubara one; in Java several breeds; one in Bali, Lomboc, Sumbawa (one of the best breeds), Tambora, Bima, Gunung-api, Celebes, Sumba, and Philippines. Other breeds are specified by Zollinger in the 'Journal of the Indian Archipelago,' vol. v. p. 343, &c.

[103] 'The Horse,' &c., by John Lawrence, 1829, p. 14.

[104] 'The Veterinary,' London, vol. v. p. 543.

[105] Proc. Veterinary Assoc., in 'The Veterinary,' vol. xiii. p. 42.

[106] 'Bulletin de la Soc. Geolog.,' tom. xxii., 1866, p. 22.

[107] Mr. Percival, of the Enniskillen Dragoons, in 'The Veterinary,' vol. i. p. 224: see Azara, 'Des Quadrupedes du Paraguay,' tom. ii. p. 313. The French translator of Azara refers to other cases mentioned by Huzard as occurring in Spain.

[108] Godron, 'De l'Espece,' tom i. p. 378.

[109] 'Ueber die Eigenschaften,' &c., 1828, s. 10.

[110] 'Domesticated Animals of the British Islands,' pp. 527, 532. In all the veterinary treatises and papers which I have read, the writers insist in the strongest terms on the inheritance by the horse of all good and bad tendencies and qualities. Perhaps the principle of inheritance is not really stronger in the horse than in any other animal; but, from its value, the tendency has been more carefully observed.

[111] Andrew Knight crossed breeds so different in size as a dray-horse and Norwegian pony: see A. Walker on 'Intermarriage,' 1838, p. 205.

[112] 'Naturalist's Library,' Horses, vol. xii. p. 208.

[113] Gervais, 'Hist Nat. Mamm.,' tom. ii. p. 143. Owen, 'British Fossil Mammals,' p. 383.

[114] 'Kenntniss der fossilen Pferde,' 1863, s. 131.

[115] Mr. W. C. L. Martin ('The Horse,' 1845, p. 34), in arguing against the belief that the wild Eastern horses are merely feral, has remarked on the improbability of man in ancient times having extirpated a species in a region where it can now exist in numbers.

[116] 'Transact. Maryland Academy,' vol. i. part i. p. 28.

[117] Mr. Mackinnon on 'The Falkland Islands,' p. 25. The average height of the Falkland horses is said to be 14 hands 2 inches. See also my 'Journal of Researches.'

[118] Pallas, 'Act. Acad. St. Petersburgh,' 1777, part ii. p. 265. With respect to the tarpans scraping away the snow, see Col. Hamilton Smith in 'Nat. Lib.,' vol. xii. p. 165.

[119] Franklin's 'Narrative,' vol. i. p. 87; note by Sir J. Richardson.

[120] Mr. J. H. Moor, 'Notices of the Indian Archipelago:' Singapore, 1837, p. 189. A pony from Java was sent ('Athenaeum,' 1842, p. 718) to the Queen only 28 inches in height. For the Loo Choo Islands, see Beechey's 'Voyage,' 4th edit., vol. i. p. 499.

[121] J. Crawford, 'History of the Horse;' 'Journal of Royal United Service Institution,' vol. iv.

[122] 'Essays on Natural History,' 2nd series, p. 161.

[123] 'Quadrupedes du Paraguay,' tom. ii. p. 333.

[124] Prof. Low, 'Domesticated Animals,' p. 546. With respect to the writer in India, see 'India Sporting Review,' vol. ii. p. 181. As Lawrence has remarked ('The Horse,' p. 9), "perhaps no instance has ever occurred of a three-part bred horse (i.e. a horse, one of whose grand-parents was of impure blood) saving his distance in running two miles with thoroughbred racers." Some few instances are on record of seven-eighths racers having been successful.

[125] Prof. Gervais (in his 'Hist. Nat. Mamm.,' tom. ii. p. 144) has collected many facts on this head. For instance, Solomon (Kings, b. i. ch. x. v. 28) bought horses in Egypt at a high price.

[126] 'The Field,' July 13th, 1861, p. 42.

[127] E. Vernon Harcourt, 'Sporting in Algeria,' p. 26.

[128] I state this from my own observations made during several years on the colours of horses. I have seen cream-coloured, light-dun and mouse-dun horses dappled, which I mention because it has been stated (Martin, 'History of the Horse,' p. 134) that duns are never dappled. Martin (p. 205) refers to dappled asses. In 'The Farrier' (London, 1828, pp. 453, 455) there are some good remarks on the dappling of horses; and likewise in Col. Hamilton Smith on 'The Horse.'

[129] Some details are given in 'The Farrier,' 1828, pp. 452, 455. One of the least ponies I ever saw, of the colour of a mouse, had a conspicuous spinal stripe. A small Indian chesnut pony had the same stripe, as had a remarkably heavy chesnut cart-horse. Race-horses often have the spinal stripe.

[130] I have received information, through the kindness of the Consul-General, Mr. J. R. Crowe, from Prof. Boeck, Rasck, and Esmarck, on the colours of the Norwegian ponies. See, also, 'The Field,' 1861, p. 431.

[131] Col. Ham. Smith, 'Nat. Lib.,' vol. xii. p. 275.

[132] Mr. G. Clark, in 'Annal and Mag. of Nat. History,' 2nd series, vol. ii., 1848, p. 363. Mr. Wallace informs me that he saw in Java a dun and clay-coloured horse with spinal and leg stripes.

[133] See, also, on this point, 'The Field,' July 27th, 1861, p. 91.

[134] 'The Field,' 1861, pp. 431, 493, 545.

[135] 'Ueber die Eigenschaften,' &c, 1828, s. 13, 14.

[136] 'Naturalist's Library,' vol. xii. (1841), pp. 109, 156 to 163, 280, 281. Cream-colour, passing into Isabella (i.e. the colour of the dirty linen of Queen Isabella), seems to have been common in ancient times. See also Pallas's account of the wild horses of the East, who speaks of dun and brown as the prevalent colours.

[137] Azara, 'Quadrupedes du Paraguay,' tom. ii. p. 307; for the colour of mules, see p. 350. In North America, Catlin (vol. ii. p. 57) describes the wild horses, believed to have descended from the Spanish horses of Mexico, as of all colours, black, grey, roan, and roan pied with sorrel. F. Michaux ('Travels in North America,' Eng. translat., p. 235) describes two wild horses from Mexico as roan. In the Falkland Islands, where the horse has been feral only between 60 and 70 years, I was told that roans and iron-greys were the prevalent colours. These several facts show that horses do not generally revert to any uniform colour.

[138] Dr. Sclater, in 'Proc. Zoolog. Soc.,' 1862, p. 164.

[139] W. C. Martin, 'History of the Horse,' 1845, p. 207.

[140] Col. Sykes' Cat. of Mammalia, 'Proc. Zoolog. Soc.,' July 12th, 1831. Williamson, 'Oriental Field Sports,' vol. ii., quoted by Martin, p. 206.

[141] Blyth, in 'Charlesworth's Mag. of Nat. Hist.,' vol. iv., 1840, p. 83. I have also been assured by a breeder that this is the case.

[142] One case is given by Martin, 'The Horse,' p. 205.

[143] 'Journal As. Soc. of Bengal,' vol. xxviii. 1860, p. 231. Martin on the Horse, p. 205.

[144] Hermann von Nathusius, 'Die Racen des Schweines,' Berlin, 1860; and 'Vorstudien fur Geschichte,' &c., 'Schweineschaedel,' Berlin, 1864. Ruetimeyer, 'Die Fauna der Pfahlbauten,' Basel, 1861.

[145] Nathusius, 'Die Racen des Schweines,' Berlin, 1860. An excellent appendix is given with references to published and trustworthy drawings of the breeds of each country.

[146] For Europe, see Bechstein, 'Naturgesch. Deutschlands,' 1801, b. i., s. 505. Several accounts have been published on the fertility of the offspring from wild and tame swine. See Burdach's 'Physiology,' and Godron, 'De l'Espece,' tom. i. p. 370. For Africa, 'Bull. de la Soc. d'Acclimat.,' tom. iv. p. 389. For India, see Nathusius, 'Schweineschaedel,' s. 148.

[147] Sir W. Elliot, Catalogue of Mammalia, 'Madras Journal of Lit. and Science,' vol. x. p. 219.

[148] 'Pfahlbauten,' s. 163 et passim.

[149] See Ruetimeyer's Neue Beitrage, ... Torfschweine, Verh. Naturfor. Gesell. in Basel, iv. i., 1865, s. 139.

[150] Stan. Julien, quoted by De Blainville, 'Osteographie,' p. 163.

[151] Richardson, 'Pigs, their Origin,' &c., p. 26.

[152] 'Die Racen des Schweines,' s. 47, 64.

[153] 'Proc. Zoolog. Soc.,' 1861, p. 263.

[154] Sclater, in 'Proc. Zoolog. Soc.,' Feb. 26th, 1861.

[155] 'Proc. Zoolog. Soc.,' 1862, p. 13.

[156] 'Journal of Voyages and Travels from 1821 to 1829,' vol. i. p. 300.

[157] Rev. G. Low, 'Fauna Orcadensis,' p. 10. See also Dr. Hibbert's account of the pig of the Shetland Islands.

[158] 'Die Racen des Schweines,' s. 70.

[159] These woodcuts are copied from engravings given in Mr. S. Sidney's excellent edition of 'The Pig,' by Youatt, 1860. See pp. 1, 16, 19.

[160] 'Schweineschaedel,' s. 74, 135.

[161] Nathusius, 'Die Racen des Schweines,' s. 71.

[162] 'Die Racen des Schweines,' s. 47. 'Schweineschaedel,' s. 104. Compare, also, the figures of the old Irish and the improved Irish breeds in Richardson on 'The Pig,' 1847.

[163] Quoted by Isid. Geoffroy, 'Hist. Nat. Gen.,' tom. iii. p. 441.

[164] S. Sidney, 'The Pig,' p. 61.

[165] 'Schweineschaedel,' s. 2, 20.

[166] 'Proc. Zoolog. Soc.,' 1837, p. 23. I have not given the caudal vertebrae, as Mr. Eyton says some might possibly have been lost. I have added together the dorsal and lumbar vertebrae, owing to Prof. Owen's remarks ('Journal Linn. Soc.,' vol. ii. p. 28) on the difference between dorsal and lumbar vertebrae depending only on the development of the ribs. Nevertheless the difference in the number of the ribs in pigs deserves notice.

[167] 'Edinburgh New Philosoph. Journal,' April 1863. See also De Blainville's 'Osteographie,' p. 128, for various authorities on this subject.

[168] Eudes-Deslongchamps, 'Memoires de la Soc. Linn. de Normandie,' vol. vii., 1842, p. 41. Richardson, 'Pigs, their Origin, &c.,' 1847, p. 30. Nathusius, 'Die Racen des Schweines,' 1860, s. 54.

[169] 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.

[170] For Turkish pigs, see Desmarest, 'Mammalogie,' 1820, p. 391. For those of Westphalia, see Richardson's 'Pigs, their Origin,' &c., 1847, p. 41.

[171] With respect to the several foregoing and following statements on feral pigs, see Roulin, in 'Mem. presentes par divers Savans a l'Acad.,' &c., Paris, tom. vi., 1835, p. 326. It should be observed that his account does not apply to truly feral pigs; but to pigs long introduced into the country and living in a half-wild state. For the truly feral pigs of Jamaica, see Gosse's 'Sojourn in Jamaica,' 1851, p. 386; and Col. Hamilton Smith, in 'Nat. Library,' vol. ix. p. 93. 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); but this author attributes the state of these pigs to descent from a domestic stock which he saw in Spain. Admiral Sulivan, R.N., had ample opportunities of observing the wild pigs on Eagle Islet in the Falklands; and he informs me that they resembled wild boars with bristly ridged backs and large tusks. The pigs which have run wild in the province of Buenos Ayres (Rengger, 'Saeugethiere,' s. 331) have not reverted to the wild type. De Blainville ('Osteographie,' p. 132) refers to two skulls of domestic pigs sent from Patagonia by Al. d'Orbigny, and he states that they have the occipital elevation of the wild European boar, but that the head altogether is "plus courte et plus ramassee." He refers, also, to the skin of a feral pig from North America, and says, "il ressemble tout a fait a un petit sanglier, mais il est presque tout noir, et peut-etre un peu plus ramasse dans ses formes."

[172] Gosse's 'Jamaica,' p. 386, with a quotation from Williamson's 'Oriental Field Sports.' Also Col. Hamilton Smith, in 'Naturalist's Library,' vol. ix. p. 94.

[173] S. Sidney's edition of 'Youatt on the Pig,' 1860, pp. 7, 26, 27, 29, 30.

[174] 'Schweineschaedel,' s. 140.

[175] 'Die Fauna der Pfahlbauten,' 1861, s. 109, 149, 222. See also Geoffroy Saint Hilaire, in 'Mem. 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 the common ox five, sacral vertebrae. Mr. Hodgson found the ribs either thirteen or fourteen in number; see a note in 'Indian Field,' 1858, p. 62.

[176] '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 character of the grunt-like voice of the humped cattle.

[177] Mr. H. E. Marquand, in 'The Times,' June 23rd, 1856.

[178] Vasey, 'Delineations of the Ox-Tribe,' p. 124. Brace's 'Hungary,' 1851, p. 94. The Hungarian cattle descend, according to Ruetimeyer ('Zahmen. Europ. Rindes,' 1866, s. 13), from Bos primigenius.

[179] Moll and Gayot, 'La Connaissance Gen. du Boeuf,' Paris, 1860. Fig 82 is that of the Podolian breed.

[180] A translation appeared in three parts in the 'Annals and Mag. of Nat. Hist.,' 2nd series, vol. iv., 1849.

[181] See, also, Ruetimeyer's 'Beitrage pal. Gesch. der Wiederkauer,' Basel, 1865, s. 54.

[182] Pictet's 'Paleontologie,' tom. i. p. 365 (2nd edit.). With respect to B. trochoceros, see Ruetimeyer's 'Zahmen Europ. Rindes,' 1866, s. 26.

[183] Owen, 'British Fossil Mammals,' 1846, p. 510.

[184] 'British Pleistocene Mammalia,' by W. B. Dawkins and W. A. Sandford, 1866. p. xv.

[185] 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.

[186] 'Lecture: Royal Institution of G. Britain,' May 2nd, 1856, p. 4. 'British Fossil Mammals,' p. 513.

[187] Nilsson, in 'Annals and Mag. of Nat. Hist.,' 1849, vol. iv. p. 354.

[188] See W. R. Wilde, ut supra; and Mr. Blyth, in 'Proc. Irish Academy,' March 5th, 1864.

[189] Laing's 'Tour in Norway,' p. 110.

[190] Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 96.

[191] Idem, tom. iii. pp. 82, 91.

[192] 'Quadrupedes du Paraguay,' tom. ii. p. 360.

[193] Walther, 'Das Rindvieh,' 1817, s. 30.

[194] I am much indebted to the present Earl of Tankerville for information about his wild cattle; and for the skull which was sent to Prof. Ruetimeyer. The fullest account of the Chillingham cattle is given by Mr. Hindmarsh, together with a letter by the late Lord Tankerville, in 'Annals and Mag. of Nat. Hist.,' vol. ii., 1839, p. 274. See Bewick, 'Quadrupeds,' 2nd edit., 1791, p. 35, note. With respect to those of Duke of Queensberry, see Pennant's 'Tour in Scotland,' p. 109. For those of Chartley, see Low's 'Domesticated Animals of Britain,' 1845, p. 238. For those of Gisburne, see Bewick's 'Quadrupeds, and Encyclop. of Rural Sports,' p. 101.

[195] Boethius was born in 1470; 'Annals and Mag. of Nat. Hist.,' vol. ii., 1839, p. 281; and vol. iv. 1849, p. 424.

[196] Youatt on Cattle, 1834, p. 48: See also p. 242, on short-horn cattle. Bell, in his 'British Quadrupeds,' p. 423, states that, after long attending to the subject, he has found that white cattle invariably have coloured ears.

[197] Azara, 'Des Quadrupedes du Paraguay,' tom. ii. p. 361. Azara quotes Buffon for the feral cattle of Africa. For Texas, see 'Times,' Feb. 18th, 1846.

[198] Anson's Voyage. See Kerr and Porter's 'Collection,' vol. xii. p. 103.

[199] See also Mr. Mackinnon's pamphlet on the Falkland Islands, p. 24.

[200] 'The Age of the Ox, Sheep, Pig,' &c., by Prof. James Simonds, published by order of the Royal Agricult. Soc.

[201] 'Ann. Agricult. France,' April 1897. as quoted in 'The Veterinary,' vol. xii. p. 725. I quote Tessier's observations from Youatt on Cattle, p. 527.

[202] 'The Veterinary,' vol. viii. p. 681, and vol. x. p. 268. Low's 'Domest. Animals of Great Britain,' p. 297.

[203] Mr. Ogleby, in 'Proc. Zoolog. Soc.,' 1836, p. 138, and 1840, p. 4.

[204] Leguat's Voyage, quoted by Vasey in his 'Delineations of the Ox-tribe,' p. 132.

[205] 'Travels in South Africa,' pp. 317, 336.

[206] 'Mem. de l'Institut present. par divers Savans,' tom. vi., 1835, p. 333. For Brazil, see 'Comptes Rendus,' June 15th, 1846. See Azara, 'Quadrupedes du Paraguay,' tom. ii. pp. 359, 361.

[207] 'Schweineschaedel,' 1864, s. 104. Nathusius states that the form of skull characteristic of 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 the "bulldog cod." Prof. Wyman also concluded, after making numerous inquiries in La Plata, that the niata cattle transmit their peculiarities or form a race.

[208] Ueber Art des Zahmen Europ. Rindes, 1866, s. 28.

[209] '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. Ruetimeyer.

[210] Loudon's 'Magazine of Nat. Hist.,' vol. i., 1829, p. 113. Separate figures are given of the animal, its hoofs, eye, and dewlap.

[211] Low, 'Domesticated Animals of the British Isles,' p. 264.

[212] 'Mem. de l'Institut present. par divers Savans,' tom. vi., 1835, p. 332.

[213] Idem, pp. 304, 368, &c.

[214] Youatt on Cattle, p. 193. A full account of this bull is taken from Marshall.

[215] Youatt on Cattle, p. 116. Lord Spencer has written on this same subject.

[216] Blyth on the genus Ovis, in 'Annals and Mag. of Nat. History,' vol. vii., 1841, p. 261: with respect to the parentage of the breeds, see Mr. Blyth's excellent articles in 'Land and Water,' 1867, pp. 134, 156. Gervais, 'Hist. Nat. des Mammiferes,' 1855, tom. ii. p. 191.

[217] Dr. L. Fitzinger, 'Ueber die Racen des Zahmen Schafes,' 1860, s. 86.

[218] J. Anderson, 'Recreations in Agriculture and Natural History,' vol. ii. p. 164.

[219] 'Pfahlbauten,' s. 127, 193.

[220] Youatt on Sheep, p. 120.

[221] 'Journal of the Asiatic Soc. of Bengal,' vol. xvi. pp. 1007, 1016.

[222] Youatt on Sheep, pp. 142-169.

[223] 'Journal Asiat. Soc. of Bengal,' vol. xvi., 1847, p. 1015.

[224] 'Hist. Nat. Gen.,' tom. iii. p. 435.

[225] Youatt on Sheep, p. 138.

[226] 'Journal Asiat. Soc. of Bengal,' vol. xvi., 1847, pp. 1015, 1016.

[227] 'Racen des Zahmen Schafes,' s. 77.

[228] 'Rural Economy of Norfolk,' vol. ii. p. 136.

[229] Youatt on Sheep, p. 312. On same subject, see excellent remarks in 'Gardener's Chronicle,' 1858, p. 868. For experiments in crossing Cheviot sheep with Leicesters, see Youatt, p. 325.

[230] Youatt on Sheep, note, p. 491.

[231] 'The Veterinary,' vol. x. p. 217.

[232] A translation of his paper is given in 'Bull. Soc. Imp. d'Acclimat.,' tom. ix., 1862, p. 723.

[233] Erman's 'Travels in Siberia' (Eng. trans.), vol. i. p. 228. For Pallas on the fat-tailed sheep, I quote from Anderson's account of the 'Sheep of Russia,' 1794, p. 34. With respect to the Crimean sheep, see Pallas' 'Travels' (Eng. trans.), vol. ii. p. 454. For the Karakool sheep, see Burnes' 'Travels in Bokhara,' vol. iii. p. 151.

[234] See Report of the Directors of the Sierra Leone Company, as quoted in White's 'Gradation of Man,' p. 95. With respect to the change which sheep undergo in the West Indies, see also Dr. Davy, in 'Edin. New. Phil. Journal,' Jan. 1852. For the statement made by Roulin, see 'Mem. de l'Institut present. par divers Savans,' tom. vi., 1835, p. 347.

[235] Youatt on Sheep, p. 69, where Lord Somerville is quoted. See p. 117, on the presence of wool under the hair. With respect to the fleeces of Australian sheep, p. 185. On selection counteracting any tendency to change, see pp. 70, 117, 120, 168.

[236] Audubon and Bachman, 'The Quadrupeds of North America,' 1846, vol. v. p. 365.

[237] 'Journal of R. Agricult. Soc. of England,' vol. xx., part ii. W. C. Spooner on Cross-Breeding.

[238] 'Philosoph. Transactions,' London, 1813, p. 88.

[239] Isidore Geoffroy St. Hilaire, 'Hist. Nat. Generale,' tom. iii. p. 87. Mr. Blyth ('Land and Water,' 1867, p. 37) has arrived at a similar conclusion, but he thinks that certain Eastern races may perhaps be in part descended from the Asiatic markhor.

[240] Ruetimeyer, 'Pfahlbauten,' s. 127.

[241] Godron, 'De l'Espece,' tom. i. p. 402.

[242] 'Annals and Mag. of Nat. History,' vol. ii. (2nd series), 1848, p. 363.

[243] 'De l'Espece,' tom. i. p. 406. Mr. Clark also refers to differences in the shape of the mammae. Godron states that in the Nubian race the scrotum is divided into two lobes; and Mr. Clark gives a ludicrous proof of this fact, for he saw in the Mauritius a male goat of the Muscat breed purchased at a high price for a female in full milk. These differences in the scrotum are probably not due to descent from distinct species; for Mr. Clark states that this part varies much in form.

[244] Mr. Clark, 'Annals and Mag. of Nat. Hist.,' vol. ii. (2nd series), 1848, p. 361.

[245] Desmarest, 'Encyclop. Method. Mammalogie,' p. 480.

[246] 'Journal of Asiatic Soc. of Bengal,' vol. xvi., 1847, pp. 1020, 1025.

[247] M. P. Gervais, 'Hist. Nat. des Mammiferes, tom. i., 1854, p. 288.

[248] U. Aldrovandi, 'De Quadrupedibus digitatis,' 1637, p. 383. For Confucius and G. Markham, see a writer who has studied the subject, in 'Cottage Gardener,' Jan. 22nd, 1861, p. 250.

[249] Owen, 'British Fossil Mammals,' p. 212.

[250] Bechstein, 'Naturgesch. Deutschlands,' 1801, b. i. p. 1133. I have received similar accounts with respect to England and Scotland.

[251] 'Pigeons and Rabbits,' by E. S. Delamer, 1854, p. 133. Sir J. Sebright ('Observations on Instinct,' 1836, p. 10) speaks most strongly on the difficulty. But this difficulty is not invariable, as I have received two accounts of perfect success in taming and breeding from the wild rabbit. See also Dr. P. Broca, in 'Journal de la Physiologie' tom. ii. p. 368.

Transcriber's Note: this note and the previous one were interchanged; corrected by Errata page.

[252] Gervais, 'Hist. Nat. des Mammiferes,' tom. i. p. 292.

[253] See Dr. P. Broca's interesting memoir on this subject in Brown-Sequard's 'Journ. de Phys.' vol. ii. p. 367.

[254] They are briefly described in the 'Journal of Horticulture,' May 7th, 1861, p. 108.

[255] 'Journal of Horticulture,' 1861, p. 380.

[256] 'Journal of Horticulture,' May 28th, 1861, p. 169.

[257] 'Journal of Horticulture,' 1861, p. 327. With respect to the ears, see Delamer on 'Pigeons and Rabbits,' 1854, p. 141; also 'Poultry Chronicle,' vol. ii. p. 499, and ditto for 1854, p. 586.

[258] Delamer, 'Pigeons and Rabbits,' p. 136. See also 'Journal of Horticulture,' 1861, p. 375.

[259] 'An Account of the different Kinds of Sheep in the Russian Dominions,' 1794, p. 39.

[260] 'Proc. Zoolog. Soc.,' June 23rd, 1857, p. 159.

[261] 'Cottage Gardener,' 1857, p. 141.

[262] 'Journal of Horticulture,' April 9th, 1861, p. 35.

[263] Mr. Bartlett, in 'Proc. Zoolog. Soc.,' 1861. p. 40.

[264] 'Phenomenon in Himalayan Rabbits,' in 'Journal of Horticulture,' 1865, Jan. 27th, p. 102.

[265] G. R. Waterhouse, 'Natural History of Mammalia: Rodents,' 1846, pp. 52, 60, 105.

[266] Delamer on 'Pigeons and Rabbits,' p. 114.

[267] Gosse's 'Sojourn in Jamaica,' 1851, p. 441, as described by an excellent observer, Mr. R. Hill. This is the only known case in which rabbits have become feral in a hot country. They can be kept, however, at Loanda (see Livingstone's 'Travels,' p. 407). In parts of India, as I am informed by Mr. Blyth, they breed well.

[268] Darwin's 'Journal of Researches,' p. 193; and 'Zoology of the Voyage of the Beagle: Mammalia,' p. 92.

[269] Kerr's 'Collection of Voyages,' vol. ii. p. 177; p. 205 for Cada Mosto. According to a work published in Lisbon in 1717, entitled 'Historia Insulana,' written by a Jesuit, the rabbits were turned out in 1420. Some authors believe that the island was discovered in 1413.

[270] Something of the same kind has occurred on the island of Lipari, where, according to Spallanzani ('Voyage dans les deux Siciles,' quoted by Godron sur l'Espece, p. 364), a countryman turned out some rabbits which multiplied prodigiously, but, says Spallanzani, "les lapins de l'ile de Lipari sont plus petits que ceux qu'on eleve en domesticite."

[271] Waterhouse, 'Nat. Hist. Mammalia,' vol. ii. p. 36.

[272] These rabbits have run wild for a considerable time in Sandon Park, and in other places in Staffordshire and Shropshire. They originated, as I have been informed by the gamekeeper, from variously-coloured domestic rabbits which had been turned out. They vary in colour; but many are symmetrically coloured, being white with a streak along the spine, and with the ears and certain marks about the head of a blackish-grey tint. They have rather longer bodies than common rabbits.

[273] See Prof. Owen's remarks on this subject in his paper on the 'Zoological Significance of the Brain, &c., of Man, &c.,' read before Brit. Association, 1862; with respect to Birds, see 'Proc. Zoolog. Soc.,' Jan. 11th, 1848, p. 8.

[274] This standard is apparently considerably too low, for Dr. Crisp ('Proc. Zoolog. Soc.,' 1861, p. 80) gives 210 grains as the actual weight of the brain of a hare which weighed 7lbs., 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.

[275] The Hon. C. Murray has sent me some very valuable specimens from Persia; and H.M. Consul, Mr. Keith Abbott, has given me information on the pigeons of the same country. I am deeply indebted to Sir Walter Elliot for an immense collection of skins from Madras, with much information regarding them. Mr. Blyth has freely communicated to me his stores of knowledge on this and all other related subjects. The Rajah Sir James Brooke sent me specimens from Borneo, as has H.M. Consul, Mr. Swinhoe, from Amoy in China, and Dr. Daniell from the west coast of Africa.

[276] Mr. B. P. Brent, well known for his various contributions to poultry literature, has aided me in every way during several years; so has Mr. Tegetmeier, with unwearied kindness. This latter gentleman, who is well known for his works on poultry, and who has largely bred pigeons, has looked over this and the following chapters. Mr. Bult formerly showed me his unrivalled collection of Pouters, and gave me specimens. I had access to Mr. Wicking's collection, which contained a greater assortment of many kinds than could anywhere else be seen; and he has always aided me with specimens and information given in the freest manner. Mr. Haynes and Mr. Corker have given me specimens of their magnificent Carriers. To Mr. Harrison Weir I am likewise indebted. Nor must I by any means pass over the assistance received from Mr. J. M. Eaton, Mr. Baker, Mr. Evans, and Mr. J. Baily, jun., of Mount-street—to the latter gentleman I have been indebted for some valuable specimens. To all these gentlemen I beg permission to return my sincere and cordial thanks.

[277] 'Les Pigeons de Voliere et de Colombier,' Paris, 1824. During forty-five years the sole occupation of M. Corbie was the care of the pigeons belonging to the Duchess of Berry.

[278] 'Coup d'Oeil sur l'Ordre des Pigeons,' par Prince C. L. Bonaparte, Paris, 1855. This author makes 288 species, ranked under 85 genera.

[279] As I so often refer to the size of the C. livia, or rock-pigeon, it may be convenient to give the mean between the measurements of two wild birds, kindly sent me by Dr. Edmondstone from the Shetland Islands:—

Inches. Length from feathered base of beak to end of tail 14.25 " " " " to oil-gland 9.5 " from tip of beak to end of tail 15.02 " of tail-feathers 4.62 " from tip to tip of wing 26.75 " of folded wing 9.25

Beak.—Length from tip of beak to feathered base .77 " Thickness, measured vertically at further end of nostrils .23 " Breadth, measured at same place .16

Feet.—Length from end of middle toe (without claw) to distal end of tibia 2.77 " Length from end of middle toe to end of hind toe (without claws) 2.02

Weight 141/4 ounces.

[280] This drawing was made from a dead bird. The six following figures were drawn with great care by Mr. Luke Wells from living birds selected by Mr. Tegetmeier. It may be confidently asserted that the characters of the six breeds which have been figured are not in the least exaggerated.

[281] 'Das Ganze der Taubenzucht:' Weimar, 1837, pl. 11 and 12.

[282] Boitard and Corbie, 'Les Pigeons,' &c., p. 177, pl. 6.

[283] 'Die Taubenzucht,' Ulm, 1824, s. 42.

[284] This treatise was written by Sayzid Mohammed Musari, who died in 1770: I owe to the great kindness of Sir W. Elliot a translation of this curious treatise.

[285] 'Poultry Chronicle,' vol. ii. p. 573.

[286] 'Annals and Mag. of Nat. History,' vol. xix., 1847, p. 105.

[287] 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 destitute of an oil-gland, have an unusual number of tail-feathers, namely 16, and in this respect resemble Fantails.

[288] See the two excellent editions published by Mr. J. M. Eaton in 1852 and 1858, entitled 'A Treatise on Fancy Pigeons.'

[289] English translation, by F. Gladwin, 4th edition, vol. i. The habit of the Lotan is also described in the Persian treatise before alluded to, published about 100 years ago: at this date the Lotans were generally white and crested as at present. Mr. Blyth describes these birds in 'Annals and Mag. of Nat. Hist.,' vol. xiv., 1847, p. 104: he says that they "may be seen at any of the Calcutta bird-dealers."

[290] 'Journal of Horticulture,' Oct. 22, 1861, p. 76.

[291] See the account of the House-tumblers kept at Glasgow, in the 'Cottage Gardener,' 1858, p. 285. Also Mr. Brent's paper, 'Journal of Horticulture,' 1861, p. 76.

[292] J. M. Eaton's 'Treatise on Pigeons,' 1852, p. 9.

[293] J. M. Eaton's Treatise, edit. 1858, p. 76.

[294] Neumeister,'Taubenzucht,' Tab. 4, fig. i.

[295] Riedel, 'Die Taubenzucht,' 1824, s. 26. Bechstein, 'Naturgeschichte Deutschlands,' Band iv. s. 36, 1795.

[296] Willoughby's 'Ornithology,' edited by Ray.

[297] J. M. Eaton's edition (1858) of Moore, p. 98.

[298] Pigeon Patu Plongeur. 'Les Pigeons,' &c., p. 165.

[299] 'Naturgesch. Deutschlands,' Band iv. s. 47.

[300] Mr. W. B. Tegetmeier, 'Journal of Horticulture,' Jan. 20th, 1863, p. 58.

[301] 'Coup-d'oeil sur l'Ordre des Pigeons,' 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 number—twelve."

[302] Described and figured in the 'Poultry Chronicle,' vol. iii., 1855, p. 82.

[303] 'The Pigeon Book,' by Mr. B. P. Brent, 1859, p. 41.

[304] 'Die Staarhaelsige Taube, Das Ganze, &c.,' s. 21, tab. i. fig. 4.

[305] 'A Treatise on the Almond Tumbler,' by J. M. Eaton, 1852, p. 8, et passim.

[306] A Treatise, &c, p. 10.

[307] Boitard and Corbie, 'Les Pigeons,' &c. 1824, p. 173.

[308] 'Le Pigeon Voyageur Belge,' 1865, p. 87.

[309] Prof. A. Newton ('Proc. Zoolog. Soc.' 1865, p. 716) remarks that he knows no species which presents any remarkable sexual distinction; but it is stated ('Naturalist's Library, Birds,' vol. ix. p. 117) that the excrescence at the base of the beak in the Carpophaga oceanica is sexual: this, if correct, is an interesting point of analogy with the male Carrier, which has the wattle at the base of its beak so much more developed than in the female. Mr. Wallace informs me that in the sub-family of the Treronidae the sexes often differ in vividness of colour.

[310] I am not sure that I have designated the different kinds of vertebrae correctly: but I observe that different anatomists follow in this respect different rules, and, as I use the same terms in the comparison of all the skeletons, this, I hope, will not signify.

[311] J. M. Eaton's Treatise, edit. 1858, p. 78.

[312] In an analogous, but converse, manner, certain natural groups of the Columbidae, from being more terrestrial in their habits than other allied groups, have larger feet. See Prince Bonaparte's 'Coup-d'oeil sur l'Ordre des Pigeons.'

[313] It perhaps deserves notice that besides these five birds two of the eight were barbs, which, as I have shown, must be classed in the same group with the long-beaked carriers and runts. Barbs may properly be called short-beaked carriers. It would, therefore, appear as if, during the reduction of their beaks, their wings had retained a little of that excess of length which is characteristic of their nearest relations and progenitors.

[314] Temminck, 'Hist. Nat. Gen. des Pigeons et des Gallinaces,' tom. i., 1813, p. 170.

[315] This term was used by John Hunter for such differences in structure between the males and females, as are not directly connected with the act of reproduction, as the tail of the peacock, the horns of deer, &c.

[316] Temminck, 'Hist. Nat. Gen. des Pigeons,' &c., tom. i. p. 191.

[317] I have heard through Sir C. Lyell from Miss Buckley, that some half-bred carriers kept during many years near London regularly settled by day on some adjoining trees, and, after being disturbed in their loft by their young being taken, roosted on them at night.

[318] 'Annals and Mag. of Nat. Hist.,' 2nd ser., vol. xx., 1857, p. 509; and in a late volume of the Journal of the Asiatic Society.

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