The Variation of Animals and Plants under Domestication Volume II Part 19

[Hardy varieties would alone be valued or noticed in Europe; whilst tender varieties, requiring more warmth, would generally be neglected; but such occasionally arise. Thus Loudon (24/71. 'Arboretum et Fruticetum' volume 3 page 1376.) describes a Cornish variety of the elm which is almost an evergreen, and of which the shoots are often killed by the autumnal frosts, so that its timber is of little value. Horticulturists know that some varieties are much more tender than others: thus all the varieties of the broccoli are more tender than cabbages; but there is much difference in this respect in the sub-varieties of the broccoli; the pink and purple kinds are a little hardier than the white Cape broccoli, "but they are not to be depended on after the thermometer falls below 24 deg Fahr.;" the Walcheren broccoli is less tender than the Cape, and there are several varieties which will stand much severer cold than the Walcheren. (24/72. Mr. Robson in 'Journal of Horticulture' 1861 page 23.) Cauliflowers seed more freely in India than cabbages. (24/73. Dr.

Bonavia 'Report of the Agri.-Hort. Soc. of Oudh' 1866.) To give one instance with flowers: eleven plants raised from a hollyhock, called the Queen of the Whites (24/74. 'Cottage Gardener' 1860 April 24 page 57.) were found to be much more tender than various other seedlings. It may be presumed that all tender varieties would succeed better under a climate warmer than ours. With fruit-trees, it is well known that certain varieties, for instance of the peach, stand forcing in a hot-house better than others; and this shows either pliability of organisation or some const.i.tutional difference. The same individual cherry-tree, when forced, has been observed during successive years gradually to change its period of vegetation. (24/75. 'Gardener's Chronicle'

1841 page 291.) Few pelargoniums can resist the heat of a stove, but Alba Multiflora will, as a most skilful gardener a.s.serts, "stand pine-apple top and bottom heat the whole winter; without looking any more drawn than if it had stood in a common greenhouse; and Blanche Fleur seems as if it had been made on purpose for growing in winter, like many bulbs, and to rest all summer."

(24/76. Mr. Beaton in 'Cottage Gardener' March 20, 1860 page 377. Queen Mab will also stand stove heat. See 'Gardener's Chronicle' 1845 page 226.) There can hardly be a doubt that the Alba Multiflora pelargonium must have a widely different const.i.tution from that of most other varieties of this plant; it would probably withstand even an equatorial climate.

We have seen that according to Labat the vine and wheat require acclimatisation in order to succeed in the West Indies. Similar facts have been observed at Madras: "two parcels of mignonette-seed, one direct from Europe, the other saved at Bangalore (of which the mean temperature is much below that of Madras), were sown at the same time: they both vegetated equally favourably, but the former all died off a few days after they appeared above ground; the latter still survive, and are vigorous, healthy plants." "So again, turnip and carrot seed saved at Hyderabad are found to answer better at Madras than seed from Europe or from the Cape of Good Hope." (24/77.

'Gardener's Chronicle' 1841 page 439.) Mr. J. Scott of the Calcutta Botanic Gardens, informs me that seeds of the sweet-pea (Lathyrus odoratus) imported from England produce plants, with thick, rigid stems and small leaves, which rarely blossom and never yield seed; plants raised from French seed blossom sparingly, but all the flowers are sterile; on the other hand, plants raised from sweet-peas grown near Darjeeling in Upper India, but originally derived from England, can be successfully cultivated on the plains of India; for they flower and seed profusely, and their stems are lax and scandent. In some of the foregoing cases, as Dr. Hooker has remarked to me, the greater success may perhaps be attributed to the seeds having been more fully ripened under a more favourable climate; but this view can hardly be extended to so many cases, including plants, which, from being cultivated under a climate hotter than their native one, become fitted for a still hotter climate. We may therefore safely conclude that plants can to a certain extent become accustomed to a climate either hotter or colder than their own; although the latter cases have been more frequently observed.]

We will now consider the means by which acclimatisation may be effected, namely, through the appearance of varieties having a different const.i.tution, and through the effects of habit. In regard to new varieties, there is no evidence that a change in the const.i.tution of the offspring necessarily stands in any direct relation with the nature of the climate inhabited by the parents. On the contrary, it is certain that hardy and tender varieties of the same species appear in the same country. New varieties thus spontaneously arising become fitted to slightly different climates in two different ways; firstly, they may have the power, either as seedlings or when full-grown, of resisting intense cold, as with the Moscow pear, or of resisting intense heat, as with some kinds of Pelargonium, or the flowers may withstand severe frost, as with the Forelle pear. Secondly, plants may become adapted to climates widely different from their own, from flowering and fruiting either earlier or later in the season. In both these cases the power of acclimatisation by man consists simply in the selection and preservation of new varieties. But without any direct intention on his part of securing a hardier variety, acclimatisation may be unconsciously effected by merely raising tender plants from seed, and by occasionally attempting their cultivation further and further northwards, as in the case of maize, the orange and the peach.

How much influence ought to be attributed to inherited habit or custom in the acclimatisation of animals and plants is a much more difficult question. In many cases natural selection can hardly have failed to have come into play and complicated the result. It is notorious that mountain sheep resist severe weather and storms of snow which would destroy lowland breeds; but then mountain sheep have been thus exposed from time immemorial, and all delicate individuals will have been destroyed, and the hardiest preserved. So with the Arrindy silk-moths of China and India; who can tell how far natural selection may have taken a share in the formation of the two races, which are now fitted for such widely different climates? It seems at first probable that the many fruit-trees which are so well fitted for the hot summers and cold winters of North America, in contrast with their poor success under our climate, have become adapted through habit; but when we reflect on the mult.i.tude of seedlings annually raised in that country, and that none would succeed unless born with a fitting const.i.tution, it is possible that mere habit may have done nothing towards their acclimatisation. On the other hand, when we hear that Merino sheep, bred during no great number of generations at the Cape of Good Hope--that some European plants raised during only a few generations in the cooler parts of India, withstand the hotter parts of that country much better than the sheep or seeds imported directly from England, we must attribute some influence to habit. We are led to the same conclusion when we hear from Naudin (24/78. Quoted by Asa Gray in 'Am. Journ. of Sc.' 2nd series January 1865 page 106.) that the races of melons, squashes, and gourds, which have long been cultivated in Northern Europe, are comparatively more precocious, and need much less heat for maturing their fruit, than the varieties of the same species recently brought from tropical regions. In the reciprocal conversion of summer and winter wheat, barley, and vetches into each other, habit produces a marked effect in the course of a very few generations. The same thing apparently occurs with the varieties of maize, which, when carried from the Southern States of America, or into Germany, soon became accustomed to their new homes. With vine-plants taken to the West Indies from Madeira, which are said to succeed better than plants brought directly from France, we have some degree of acclimatisation in the individual, independently of the production of new varieties by seed.

The common experience of agriculturists is of some value, and they often advise persons to be cautious in trying the productions of one country in another. The ancient agricultural writers of China recommend the preservation and cultivation of the varieties peculiar to each country. During the cla.s.sical period, Columella wrote, "Vernaculum pecus peregrino longe praestantius est." (24/79. For China see 'Memoire sur les Chinois' tome 11 1786 page 60. Columella is quoted by Carlier in 'Journal de Physique' tome 24 1784.)

I am aware that the attempt to acclimatise either animals or plants has been called a vain chimera. No doubt the attempt in most cases deserves to be thus called, if made independently of the production of new varieties endowed with a different const.i.tution. With plants propagated by buds, habit rarely produces any effect; it apparently acts only through successive seminal generations. The laurel, bay, laurestinus, etc., and the Jerusalem artichoke, which are propagated by cuttings or tubers, are probably now as tender in England as when first introduced; and this appears to be the case with the potato, which until recently was seldom multiplied by seed. With plants propagated by seed, and with animals, there will be little or no acclimatisation unless the hardier individuals are either intentionally or unconsciously preserved. The kidney-bean has often been advanced as an instance of a plant which has not become hardier since its first introduction into Britain. We hear, however, on excellent authority (24/80. Messrs. Hardy and Son in 'Gardener's Chronicle' 1856 page 589.) that some very fine seed, imported from abroad, produced plants "which blossomed most profusely, but were nearly all but abortive, whilst plants grown alongside from English seed podded abundantly;" and this apparently shows some degree of acclimatisation in our English plants. We have also seen that seedlings of the kidney-bean occasionally appear with a marked power of resisting frost; but no one, as far as I can hear, has ever separated such hardy seedlings, so as to prevent accidental crossing, and then gathered their seed, and repeated the process year after year. It may, however, be objected with truth that natural selection ought to have had a decided effect on the hardiness of our kidney- beans; for the tenderest individuals must have been killed during every severe spring, and the hardier preserved. But it should be borne in mind that the result of increased hardiness would simply be that gardeners, who are always anxious for as early a crop as possible, would sow their seed a few days earlier than formerly. Now, as the period of sowing depends much on the soil and elevation of each district, and varies with the season; and as new varieties have often been imported from abroad, can we feel sure that our kidney-beans are not somewhat hardier? I have not been able, by searching old horticultural works, to answer this question satisfactorily.

On the whole the facts now given show that, though habit does something towards acclimatisation, yet that the appearance of const.i.tutionally different individuals is a far more effective agent. As no single instance has been recorded either with animals or plants of hardier individuals having been long and steadily selected, though such selection is admitted to be indispensable for the improvement of any other character, it is not surprising that man has done little in the acclimatisation of domesticated animals and cultivated plants. We need not, however, doubt that under nature new races and new species would become adapted to widely different climates, by variation, aided by habit, and regulated by natural selection.

[ARRESTS OF DEVELOPMENT: RUDIMENTARY AND ABORTED ORGANS.

Modifications of structure from arrested development, so great or so serious as to deserve to be called monstrosities, are not infrequent with domesticated animals, but, as they differ much from any normal structure, they require only a pa.s.sing notice. Thus the whole head may be represented by a soft nipple-like projection, and the limbs by mere papillae. These rudiments of limbs are sometimes inherited, as has been observed in a dog. (24/81. Isid. Geoffroy Saint-Hilaire 'Hist. Nat. des Anomalies' 1836 tome 2 pages 210, 223, 224, 395; 'Philosoph. Transact.' 1775 page 313.)

Many lesser anomalies appear to be due to arrested development. What the cause of the arrest may be, we seldom know, except in the case of direct injury to the embryo. That the cause does not generally act at an extremely early embryonic period we may infer from the affected organ seldom being wholly aborted,--a rudiment being generally preserved. The external ears are represented by mere vestiges in a Chinese breed of sheep; and in another breed, the tail is reduced "to a little b.u.t.ton, suffocated in a manner, by fat." (24/82. Pallas quoted by Youatt on 'Sheep' page 25.) In tailless dogs and cats a stump is left. In certain breeds of fowls the comb and wattles are reduced to rudiments; in the Cochin-China breed scarcely more than rudiments of spurs exist. With polled Suffolk cattle, "rudiments of horns can often be felt at an early age" (24/83. Youatt on 'Cattle' 1834 page 174.); and with species in a state of nature, the relatively great development of rudimentary organs at an early period of life is highly characteristic of such organs.

With hornless breeds of cattle and sheep, another and singular kind of rudiment has been observed, namely, minute dangling horns attached to the skin alone, and which are often shed and grow again. With hornless goats, according to Desmarest (24/84. 'Encyclop. Method.' 1820 page 483: see page 500, on the Indian zebu casting its horns. Similar cases in European cattle were given in the third chapter.), the bony protuberance which properly supports the horn exists as a mere rudiment.

With cultivated plants it is far from rare to find the petals, stamens, and pistils represented by rudiments, like those observed in natural species. So it is with the whole seed in many fruits; thus, near Astrakhan there is a grape with mere traces of seeds, "so small and lying so near the stalk that they are not perceived in eating the grape." (24/85. Pallas 'Travels' English Translat. volume 1 page 243.) In certain varieties of the gourd, the tendrils, according to Naudin, are represented by rudiments or by various monstrous growths. In the broccoli and cauliflower the greater number of the flowers are incapable of expansion, and include rudimentary organs. In the Feather hyacinth (Muscari comosum) in its natural state the upper and central flowers are brightly coloured but rudimentary; under cultivation the tendency to abortion travels downwards and outwards, and all the flowers become rudimentary; but the abortive stamens and pistils are not so small in the lower as in the upper flowers. In the Viburnum opulus, on the other hand, the outer flowers naturally have their organs of fructification in a rudimentary state, and the corolla is of large size; under cultivation, the change spreads to the centre, and all the flowers become affected. In the compositae, the so- called doubling of the flowers consists in the greater development of the corolla of the central florets, generally accompanied with some degree of sterility; and it has been observed (24/86. Mr. Beaton in 'Journal of Horticulture' May 21, 1861 page 133.) that the progressive doubling invariably spreads from the circ.u.mference to the centre,--that is, from the ray florets, which so often include rudimentary organs, to those of the disc. I may add, as bearing on this subject, that with Asters, seeds taken from the florets of the circ.u.mference have been found to yield the greatest number of double flowers.

(24/87. Lecoq 'De la Fecondation' 1862 page 233.) In the above cases we have a natural tendency in certain parts to be rudimentary, and this under culture spreads either to, or from, the axis of the plant. It deserves notice, as showing how the same laws govern the changes which natural species and artificial varieties undergo, that in the species of Carthamus, one of the Compositae, a tendency to the abortion of the pappus may be traced extending from the circ.u.mference to the centre of the disc as in the so-called doubling of the flowers in the members of the same family. Thus, according to A. de Jussieu (24/88. 'Annales du Museum' tome 6 page 319.), the abortion is only partial in Carthamus creticus, but more extended in C. lanatus; for in this species only two or three of the central seeds are furnished with a pappus, the surrounding seeds being either quite naked or furnished with a few hairs; and lastly in C. tinctorius, even the central seeds are dest.i.tute of pappus, and the abortion is complete.

With animals and plants under domestication, when an organ disappears, leaving only a rudiment, the loss has generally been sudden, as with hornless and tailless breeds; and such cases may be ranked as inherited monstrosities. But in some few cases the loss has been gradual, and has been effected partly by selection, as with the rudimentary combs and wattles of certain fowls. We have also seen that the wings of some domesticated birds have been slightly reduced by disuse, and the great reduction of the wings in certain silk-moths, with mere rudiments left, has probably been aided by disuse.]

With species in a state of nature, rudimentary organs are extremely common.

Such organs are generally variable, as several naturalists have observed; for, being useless, they are not regulated by natural selection, and they are more or less liable to reversion. The same rule certainly holds good with parts which have become rudimentary under domestication. We do not know through what steps under nature rudimentary organs have pa.s.sed in being reduced to their present condition; but we so incessantly see in species of the same group the finest gradations between an organ in a rudimentary and perfect state, that we are led to believe that the pa.s.sage must have been extremely gradual. It may be doubted whether a change of structure so abrupt as the sudden loss of an organ would ever be of service to a species in a state of nature; for the conditions to which all organisms are closely adapted usually change very slowly. Even if an organ did suddenly disappear in some one individual by an arrest of development, intercrossing with the other individuals of the same species would tend to cause its partial reappearance; so that its final reduction could only be effected by some other means. The most probable view is, that a part which is now rudimentary, was formerly, owing to changed habits of life, used less and less, being at the same time reduced in size by disuse, until at last it became quite useless and superfluous. But as most parts or organs are not brought into action during an early period of life, disuse or decreased action will not lead to their reduction until the organism arrives at a somewhat advanced age; and from the principle of inheritance at corresponding ages the reduction will be transmitted to the offspring at the same advanced stage of growth. The part or organ will thus retain its full size in the embryo, as we know to be the case with most rudiments. As soon as a part becomes useless, another principle, that of economy of growth, will come into play, as it would be an advantage to an organism exposed to severe compet.i.tion to save the development of any useless part; and individuals having the part less developed will have a slight advantage over others. But, as Mr. Mivart has justly remarked, as soon as a part is much reduced, the saving from its further reduction will be utterly insignificant; so that this cannot be effected by natural selection. This manifestly holds good if the part be formed of mere cellular tissue, entailing little expenditure of nutriment. How then can the further reduction of an already somewhat reduced part be effected? That this has occurred repeatedly under Nature is shown by the many gradations which exist between organs in a perfect state and the merest vestiges of them. Mr. Romanes (24/89. I suggested in 'Nature' (volume 8 pages 432, 505) that with organisms subjected to unfavourable conditions all the parts would tend towards reduction, and that under such circ.u.mstances any part which was not kept up to its standard size by natural selection would, owing to intercrossing, slowly but steadily decrease. In three subsequent communications to 'Nature' (March 12, April 9, and July 2, 1874), Mr. Romanes gives his improved view.) has, I think, thrown much light on this difficult problem. His view, as far as it can be given in a few words, is as follows: all parts are somewhat variable and fluctuate in size round an average point.

Now, when a part has already begun from any cause to decrease, it is very improbable that the variations should be as great in the direction of increase as of diminution; for the previous reduction shows that circ.u.mstances have not been favourable for its development; whilst there is nothing to check variations in the opposite direction. If this be so, the long continued crossing of many individuals furnished with an organ which fluctuates in a greater degree towards decrease than towards increase, will slowly but steadily lead to its diminution. With respect to the complete and absolute abortion of a part, a distinct principle, which will be discussed in the chapter on pangenesis, probably comes into action.

With animals and plants reared by man there is no severe or recurrent struggle for existence, and the principle of economy will not come into action, so that the reduction of an organ will not thus be aided. So far, indeed, is this from being the case, that in some few instances organs, which are naturally rudimentary in the parent-species, become partially redeveloped in the domesticated descendants. Thus cows, like most other ruminants, properly have four active and two rudimentary mamma; but in our domesticated animals, the latter occasionally become considerably developed and yield milk. The atrophied mammae, which, in male domesticated animals, including man, have in some rare cases grown to full size and secreted milk, perhaps offer an a.n.a.logous case. The hind feet of dogs naturally include rudiments of a fifth toe, and in certain large breeds these toes, though still rudimentary, become considerably developed and are furnished with claws. In the common Hen, the spurs and comb are rudimentary, but in certain breeds these become, independently of age or disease of the ovaria, well developed. The stallion has canine teeth, but the mare has only traces of the alveoli, which, as I am informed by the eminent veterinarian Mr. G.T. Brown, frequently contain minute irregular nodules of bone. These nodules, however, sometimes become developed into imperfect teeth, protruding through the gums and coated with enamel; and occasionally they grow to a fourth or even a third of the length of the canines in the stallion. With plants I do not know whether the redevelopment of rudimentary organs occurs more frequently under culture than under nature.

Perhaps the pear-tree may be a case in point, for when wild it bears thorns, which consist of branches in a rudimentary condition and serve as a protection, but, when the tree is cultivated, they are reconverted into branches.

CHAPTER 2.XXV.

LAWS OF VARIATION, continued.--CORRELATED VARIABILITY.

EXPLANATION OF TERM CORRELATION.

CONNECTED WITH DEVELOPMENT.

MODIFICATIONS CORRELATED WITH THE INCREASED OR DECREASED SIZE OF PARTS.

CORRELATED VARIATION OF h.o.m.oLOGOUS PARTS.

FEATHERED FEET IN BIRDS a.s.sUMING THE STRUCTURE OF THE WINGS.

CORRELATION BETWEEN THE HEAD AND THE EXTREMITIES.

BETWEEN THE SKIN AND DERMAL APPENDAGES.

BETWEEN THE ORGANS OF SIGHT AND HEARING.

CORRELATED MODIFICATIONS IN THE ORGANS OF PLANTS.

CORRELATED MONSTROSITIES.

CORRELATION BETWEEN THE SKULL AND EARS.

SKULL AND CREST OF FEATHERS.

SKULL AND HORNS.

CORRELATION OF GROWTH COMPLICATED BY THE ACc.u.mULATED EFFECTS OF NATURAL SELECTION.

COLOUR AS CORRELATED WITH CONSt.i.tUTIONAL PECULIARITIES.

All parts of the organisation are to a certain extent connected together; but the connection may be so slight that it hardly exists, as with compound animals or the buds on the same tree. Even in the higher animals various parts are not at all closely related; for one part may be wholly suppressed or rendered monstrous without any other part of the body being affected. But in some cases, when one part varies, certain other parts always, or nearly always, simultaneously vary; they are then subject to the law of correlated variation. The whole body is admirably co-ordinated for the peculiar habits of life of each organic being, and may be said, as the Duke of Argyll insists in his 'Reign of Law' to be correlated for this purpose. Again, in large groups of animals certain structures always co-exist: for instance, a peculiar form of stomach with teeth of peculiar form, and such structures may in one sense be said to be correlated. But these cases have no necessary connection with the law to be discussed in the present chapter; for we do not know that the initial or primary variations of the several parts were in any way related: slight modifications or individual differences may have been preserved, first in one and then in another part, until the final and perfectly co-adapted structure was acquired; but to this subject I shall presently recur. Again, in many groups of animals the males alone are furnished with weapons, or are ornamented with gay colours; and these characters manifestly stand in some sort of correlation with the male reproductive organs, for when the latter are destroyed these characters disappear. But it was shown in the twelfth chapter that the very same peculiarity may become attached at any age to either s.e.x, and afterwards be exclusively transmitted to the same s.e.x at a corresponding age. In these cases we have inheritance limited by both s.e.x and age; but we have no reason for supposing that the original cause of the variation was necessarily connected with the reproductive organs, or with the age of the affected being.

In cases of true correlated variation, we are sometimes able to see the nature of the connection; but in most cases it is hidden from us, and certainly differs in different cases. We can seldom say which of two correlated parts first varies, and induces a change in the other; or whether the two are the effects of some common cause. Correlated variation is an important subject for us; for when one part is modified through continued selection, either by man or under nature, other parts of the organisation will be unavoidably modified.

From this correlation it apparently follows that with our domesticated animals and plants, varieties rarely or never differ from one another by a single character alone.

One of the simplest cases of correlation is that a modification which arises during an early stage of growth tends to influence the subsequent development of the same part, as well as of other and intimately connected parts. Isidore Geoffroy Saint-Hilaire states (25/1. 'Hist. des Anomalies' tome 3 page 392.

Prof. Huxley applies the same principle in accounting for the remarkable, though normal, differences in the arrangement of the nervous system in the Mollusca, in his paper on the Morphology of the Cephalous Mollusca in 'Phil.

Transact.' 1853 page 56.) that this may constantly be observed with monstrosities in the animal kingdom; and Moquin-Tandon (25/2. 'Elements de Teratologie Veg.' 1841 page 13.) remarks, that, as with plants the axis cannot become monstrous without in some way affecting the organs subsequently produced from it, so axial anomalies are almost always accompanied by deviations of structure in the appended parts. We shall presently see that with short-muzzled races of the dog certain histological changes in the basal elements of the bones arrest their development and shorten them, and this affects the position of the subsequently developed molar teeth. It is probable that certain modifications in the larvae of insects would affect the structure of the mature insects. But we must be careful not to extend this view too far, for during the normal course of development, certain species pa.s.s through an extraordinary course of change, whilst other and closely allied species arrive at maturity with little change of structure.

Another simple case of correlation is that with the increased or decreased dimensions of the whole body, or of any particular part, certain organs are increased or diminished in number, or are otherwise modified. Thus pigeon fanciers have gone on selecting pouters for length of body, and we have seen that their vertebrae are generally increased not only in size but in number, and their ribs in breadth. Tumblers have been selected for their small bodies, and their ribs and primary wing-feathers are generally lessened in number.

Fantails have been selected for their large widely-expanded tails, with numerous tail-feathers, and the caudal vertebrae are increased in size and number. Carriers have been selected for length of beak, and their tongues have become longer, but not in strict accordance with the length of beak. In this latter breed and in others having large feet, the number of the scutellae on the toes is greater than in the breeds with small feet. Many similar cases could be given. In Germany it has been observed that the period of gestation is longer in large than in small breeds of cattle. With our highly-improved breeds of all kinds, the periods of maturity and of reproduction have advanced with respect to the age of the animal; and, in correspondence with this, the teeth are now developed earlier than formerly, so that, to the surprise of agriculturists, the ancient rules for judging of the age of an animal by the state of its teeth are no longer trustworthy. (25/3. Prof. J.B. Simonds on the Age of the Ox, Sheep, etc. quoted in 'Gardener's Chronicle' 1854 page 588.)

CORRELATED VARIATION OF h.o.m.oLOGOUS PARTS.

Parts which are h.o.m.ologous tend to vary in the same manner; and this is what might have been expected, for such parts are identical in form and structure during an early period of embryonic development, and are exposed in the egg or womb to similar conditions. The symmetry, in most kinds of animals, of the corresponding or h.o.m.ologous organs on the right and left sides of the body, is the simplest case in point; but this symmetry sometimes fails, as with rabbits having only one ear, or stags with one horn, or with many-horned sheep which sometimes carry an additional horn on one side of their heads. With flowers which have regular corollas, all the petals generally vary in the same manner, as we see in the complicated and symmetrical pattern, on the flowers, for instance, of the Chinese pink; but with irregular flowers, though the petals are of course h.o.m.ologous, this symmetry often fails, as with the varieties of the Antirrhinum or snapdragon, or that variety of the kidney-bean (Phaseolus) which has a white standard-petal.

In the Vertebrata the front and hind limbs are h.o.m.ologous, and they tend to vary in the same manner, as we see in long and short legged, or in thick and thin legged races of the horse and dog. Isidore Geoffroy (25/4. 'Hist. des Anomalies' tome 1 page 674.) has remarked on the tendency of supernumerary digits in man to appear, not only on the right and left sides, but on the upper and lower extremities. Meckel has insisted (25/5. Quoted by Isid.

Geoffroy ibid tome 1 page 635.) that, when the muscles of the arm depart in number or arrangement from their proper type, they almost always imitate those of the leg; and so conversely the varying muscles of the leg imitate the normal muscles of the arm.

In several distinct breeds of the pigeon and fowl, the legs and the two outer toes are heavily feathered, so that in the trumpeter pigeon they appear like little wings. In the feather-legged bantam the "boots" or feathers, which grow from the outside of the leg and generally from the two outer toes, have, according to the excellent authority of Mr. Hewitt (25/6. 'The Poultry Book'

by W.B. Tegetmeier 1866 page 250.), been seen to exceed the wing-feathers in length, and in one case were actually nine and a half inches long! As Mr.

Blyth has remarked to me, these leg-feathers resemble the primary wing- feathers, and are totally unlike the fine down which naturally grows on the legs of some birds, such as grouse and owls. Hence it may be suspected that excess of food has first given redundancy to the plumage, and then that the law of h.o.m.ologous variation has led to the development of feathers on the legs, in a position corresponding with those on the wing, namely, on the outside of the tarsi and toes. I am strengthened in this belief by the following curious case of correlation, which for a long time seemed to me utterly inexplicable, namely, that in pigeons of any breed, if the legs are feathered, the two outer toes are partially connected by skin. These two outer toes correspond with our third and fourth toes. (25/7. Naturalists differ with respect to the h.o.m.ologies of the digits of birds; but several uphold the view above advanced. See on this subject Dr. E.S. Morse in 'Annals of the Lyceum of Nat. Hist. of New York' volume 10 1872 page 16.) Now, in the wing of the pigeon or of any other bird, the first and fifth digits are aborted; the second is rudimentary and carries the so-called "b.a.s.t.a.r.d-wing;" whilst the third and fourth digits are completely united and enclosed by skin, together forming the extremity of the wing. So that in feather-footed pigeons, not only does the exterior surface support a row of long feathers, like wing-feathers, but the very same digits which in the wing are completely united by skin become partially united by skin in the feet; and thus by the law of the correlated variation of h.o.m.ologous parts we can understand the curious connection of feathered legs and membrane between the two outer toes.

Andrew Knight (24/8. A. Walker on 'Intermarriage' 1838 page 160.) has remarked that the face or head and the limbs usually vary together in general proportions. Compare, for instance, the limbs of a dray and race horse, or of a greyhound and mastiff. What a monster a greyhound would appear with the head of a mastiff! The modern bulldog, however, has fine limbs, but this is a recently-selected character. From the measurements given in the sixth chapter, we see that in several breeds of the pigeon the length of the beak and the size of the feet are correlated. The view which, as before explained, seems the most probable is, that disuse in all cases tends to diminish the feet, the beak becoming at the same time shorter through correlation; but that in some few breeds in which length of beak has been a selected point, the feet, notwithstanding disuse, have increased in size through correlation. In the following case some kind of correlation is seen to exist between the feet and beak: several specimens have been sent to Mr. Bartlett at different times, as hybrids between ducks and fowls, and I have seen one; these were, as might be expected, ordinary ducks in a semi-monstrous condition, and in all of them the swimming-web between the toes was quite deficient or much reduced, and in all the beak was narrow and ill-shaped.

With the increased length of the beak in pigeons, not only the tongue increases in length, but likewise the orifice of the nostrils. But the increased length of the orifice of the nostrils perhaps stands in closer correlation with the development of the corrugated skin or wattle at the base of the beak, for when there is much wattle round the eyes, the eyelids are greatly increased or even doubled in length.

There is apparently some correlation even in colour between the head and the extremities. Thus with horses a large white star or blaze on the forehead is generally accompanied by white feet. (25/9. 'The Farrier and Naturalist'

volume 1 1828 page 456. A gentleman who has attended to this point, tells me that about three-fourths of white-faced horses have white legs.) With white rabbits and cattle, dark marks often co-exist on the tips of the ears and on the feet. In black and tan dogs of different breeds, tan-coloured spots over the eyes and tan-coloured feet almost invariably go together. These latter cases of connected colouring may be due either to reversion or to a.n.a.logous variation,--subjects to which I shall hereafter return,--but this does not necessarily determine the question of their original correlation. Mr. H.W.

Jackson informs me that he has observed many hundred white-footed cats, and he finds that all are more or less conspicuously marked with white on the front of the neck or chest.

The lopping forwards and downwards of the immense ears of fancy rabbits seems partly due to the disuse of the muscles, and partly to the weight and length of the ears, which have been increased by selection during many generations.

Now, with the increased size and changed direction of the ears not only has the bony auditory meatus become changed in outline, direction, and greatly in size, but the whole skull has been slightly modified. This could be clearly seen in "half-lops"--that is, in rabbits with only one ear lopping forward-- for the opposite sides of their skulls were not strictly symmetrical. This seems to me a curious instance of correlation, between hard bones and organs so soft and flexible, as well as so unimportant under a physiological point of view, as the external ears. The result no doubt is largely due to mere mechanical action, that is, to the weight of the ears, on the same principle that the skull of a human infant is easily modified by pressure.

The skin and the appendages of hair, feathers, hoofs, horns, and teeth, are h.o.m.ologous over the whole body. Every one knows that the colour of the skin and that of the hair usually vary together; so that Virgil advises the shepherd to look whether the mouth and tongue of the ram are black, lest the lambs should not be purely white. The colour of the skin and hair, and the odour emitted by the glands of the skin, are said (25/10. G.o.dron 'Sur l'Espece' tome 2 page 217.) to be connected, even in the same race of men.

Generally the hair varies in the same way all over the body in length, fineness, and curliness. The same rule holds good with feathers, as we see with the laced and frizzled breeds both of fowls and pigeons. In the common c.o.c.k the feathers on the neck and loins are always of a particular shape, called hackles: now in the Polish breed, both s.e.xes are characterised by a tuft of feathers on the head, and through correlation these feathers in the male always a.s.sume the form of hackles. The wing and tail-feathers, though arising from parts not h.o.m.ologous, vary in length together; so that long or short winged pigeons generally have long or short tails. The case of the Jacobin-pigeon is more curious, for the wing and tail feathers are remarkably long; and this apparently has arisen in correlation with the elongated and reversed feathers on the back of the neck, which form the hood.

The hoofs and hair are h.o.m.ologous appendages; and a careful observer, namely Azara (25/11. 'Quadrupedes du Paraguay' tome 2 page 333.), states that in Paraguay horses of various colours are often born with their hair curled and twisted like that on the head of a negro. This peculiarity is strongly inherited. But what is remarkable is that the hoofs of these horses "are absolutely like those of a mule." The hair also of their manes and tails is invariably much shorter than usual, being only from four to twelve inches in length; so that curliness and shortness of the hair are here, as with the negro, apparently correlated.

With respect to the horns of sheep, Youatt (25/12. 'On Sheep' page 142.) remarks that "multiplicity of horns is not found in any breed of much value; it is generally accompanied by great length and coa.r.s.eness of the fleece."

Several tropical breeds of sheep which are clothed with hair instead of wool, have horns almost like those of a goat. Sturm (25/13. 'Ueber Racen, Kreuzungen' etc. 1825 s. 24.) expressly declares that in different races the more the wool is curled the more the horns are spirally twisted. We have seen in the third chapter, where other a.n.a.logous facts have been given, that the parent of the Mauchamp breed, so famous for its fleece, had peculiarly shaped horns. The inhabitants of Angora a.s.sert (25/14. Quoted from Conolly in 'The Indian Field' February 1859 volume 2 page 266.) that "only the white goats which have horns wear the fleece in the long curly locks that are so much admired; those which are not horned having a comparatively close coat." From these cases we may infer that the hair or wool and the horns tend to vary in a correlated manner. (25/15. In the third chapter I have said that "the hair and horns are so closely related to each other, that they are apt to vary together." Dr. Wilckens ("Darwin's Theorie" 'Jahrbuch der Deutschen Viehzucht'

1866 1. Heft) translates my words into "lang- und grobhaarige Thiere sollen geneigter sein, lange und viele Horner zu bekommen" and he then justly disputes this proposition; but what I have really said, in accordance with the authorities just quoted, may, I think, be trusted.) Those who have tried hydropathy are aware that the frequent application of cold water stimulates the skin; and whatever stimulates the skin tends to increase the growth of the hair, as is well shown in the abnormal growth of hair near old inflamed surfaces. Now, Professor Low (25/16. 'Domesticated Animals of the British Islands' pages 307, 368. Dr. Wilckens argues ('Landwirth. Wochenblatt' Nr. 10 1869) to the same effect with respect to domestic animals in Germany.) is convinced that with the different races of British cattle thick skin and long hair depend on the humidity of the climate which they inhabit. We can thus see how a humid climate might act on the horns--in the first place directly on the skin and hair, and secondly by correlation on the horns. The presence or absence of horns, moreover, both in the case of sheep and cattle, acts, as will presently be shown, by some sort of correlation on the skull.