HISTORICAL HYBRID MAMMALS TEXTS
In these historical texts, I have removed or paraphrased the sections on plant and bird hybrids. Darwin, and some of his commentators, wrote extensively on these, reflecting the great interests in horticulture and fancy birds (canaries, finches, pigeons) and poultry. A few cases are paraphrased or left in the text because the author refers back to the plant cases when discussing animal hybrids. By "sterility of the first cross" the authors mean that a hybrid could not be produced by crossing the parent. Some of the species described by early naturalists are now considered to be subspecies.
EXCERPTS FROM "ANIMAL KINGDOM"
CUVIER (1769 - 1832)
We have already had occasion to offer a few observations on the production of Mules [...] We have at this place to notice a similar reproduction between two specie of a more prominent character on the great theatre of life, between, in fact, the two tyrants of the world - the African Lion and the Asiatic Tiger. [...]
But to return to our Lion-Tiger cubs. Mr Atkins, an itinerant exhibitor, and dealer himself, bred the Lion, the father of these cubs. He was a very fine and very valuable beast, for his beauty and the docility of his disposition, the ferocity of which had never been entirely developed by natural habits. At the period in question, he was about four years old. The Tigress, the mother of the cubs, is supposed to be about four or five years old. She has been in Mr Atkins's possession for about two years, and was, probably, taken very young, as the gentleness of her disposition seems to evince.
These two animals, ever since the arrival of the Tigress, have been confined in one den, and have always agreed well together. From the beginning of their being so placed, there had been frequent possibility of issue, though the first, consisting of a litter of three cubs, was not born till the 17th of October, 1824, the result of a more particular intercourse, which lasted ten or twelve days, in the beginning of the previous July. They were born at Windsor, and were shortly afterwards honoured by a visit from his Majesty. The Lion, unfortunately, died about six weeks afterwards. The cubs were taken from the Tigress immediately after birth, and were fostered by several bitches and a Goat; they are all alive, and promise, at present, to attain maturity.
In regard to their personal appearance, we feel constrained, after what has already been said, to be very brief in our description, an omission, however, we hope to compensate by the figures. These show them at three months old, and we have added figures of the young, both of Lions and Tigers, that they may be compared with this singular breed.
The young of the Lion are calculated to deceive an inexperienced observer, from the fact of their being striped transversely, so as to induce the opinion, at first sight, that they belong to the Tiger, and, in this respect, the cubs in question agreed with those of the Lion. In the young Lions, however, these stripes soon become obliterated, but in those before us, they appear to be getting more decided and permanent, and, in fact, to be assuming the permanent Tigrine character. Our Mules, in common with ordinary Lions, were born without any traces of a mane, or of a tuft at the end of the tail. Their fur, in general, was rather woolly; the external ear was pendant toward the extremity; the nails were constantly out, and not cased in the sheath; and, in these particulars, they agreed with the common cubs of Lions. Their colour was dirty-yellow or blanket-colour; but from the nose over the head, along the back, and upper side of the tail, the colour was much darker, and, on these parts, the transverse stripes were stronger, and the forehead was covered with obscure spots, slighter indications of which appeared also on other parts of the body. The shape of the head, as appears by the figures, is assimilated to that of the father (the Lion); the superficies, of the body, on the other hand, is like that of the Tigress.
(Note: "Mule" referred to any hybrid, not just to equine hybrids)
EXCERPTS FROM CHARLES DARWIN'S "ORIGIN OF SPECIES" (1859)
From Chapter 5: Laws of Variation
Now let us turn to the effects of crossing the several species of the horse-genus. Rollin asserts, that the common mule from the ass and horse is particularly apt to have bars on its legs. I once saw a mule with its legs so much striped that any one at first would have thought that it must have been the product of a zebra; and Mr. W. C. Martin, in his excellent treatise on the horse, has given a figure of a similar mule. In four coloured drawings, which I have seen, of hybrids between the ass and zebra, the legs were much more plainly barred than the rest of the body; and in one of them there was a double shoulder-stripe. In Lord Moreton's famous hybrid from a chestnut mare and male quagga, the hybrid, and even the pure offspring subsequently produced from the mare by a black Arabian sire, were much more plainly barred across the legs than is even the pure quagga. Lastly, and this is another most remarkable case, a hybrid has been figured by Dr. Gray (and he informs me that he knows of a second case) from the ass and the hemionus; and this hybrid, though the ass seldom has stripes on its legs and the hemionus has none and has not even a shoulder-stripe, nevertheless had all four legs barred, and had three short shoulder-stripes, like those on the dun Welch pony, and even had some zebra-like stripes on the sides of its face. With respect to this last fact, I was so convinced that not even a stripe of colour appears from what would commonly be called an accident, that I was led solely from the occurrence of the face-stripes on this hybrid from the ass and hemionus, to ask Colonel Poole whether such face-stripes ever occur in the eminently striped Kattywar breed of horses, and was, as we have seen, answered in the affirmative.
From Chapter 8: Hybridism.
The view generally entertained by naturalists is that species, when intercrossed, have been specially endowed with the quality of sterility, in order to prevent the confusion of all organic forms. This view certainly seems at first probable, for species within the same country could hardly have kept distinct had they been capable of crossing freely. The importance of the fact that hybrids are very generally sterile, has, I think, been much underrated by some late writers. On the theory of natural selection the case is especially important, inasmuch as the sterility of hybrids could not possibly be of any advantage to them, and therefore could not have been acquired by the continued preservation of successive profitable degrees of sterility. I hope, however, to be able to show that sterility is not a specially acquired or endowed quality, but is incidental on other acquired differences.
In treating this subject, two classes of facts, to a large extent fundamentally different, have generally been confounded together; namely, the sterility of two species when first crossed, and the sterility of the hybrids produced from them.
Pure species have of course their organs of reproduction in a perfect condition, yet when intercrossed they produce either few or no offspring. Hybrids, on the other hand, have their reproductive organs functionally impotent, as may be clearly seen in the state of the male element in both plants and animals; though the organs themselves are perfect in structure, as far as the microscope reveals. In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed, or are imperfectly developed. This distinction is important, when the cause of the sterility, which is common to the two cases, has to be considered. The distinction has probably been slurred over, owing to the sterility in both cases being looked on as a special endowment, beyond the province of our reasoning powers.
The fertility of varieties, that is of the forms known or believed to have descended from common parents, when intercrossed, and likewise the fertility of their mongrel offspring, is, on my theory, of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species.
[Darwin describes the work of botanists, Kolreuter and Gartner, who almost devoted their lives to the subject of plant hybrids. Both found most hybrids to be sterile, but Gartner disputed those Kolreuter found to be fertile. Reverend W. Herbert and horticulturalist Mr C Noble also found some plant hybrids to be perfectly fertile.]
In regard to animals, much fewer experiments have been carefully tried than with plants. If our systematic arrangements can be trusted, that is if the genera of animals are as distinct from each other, as are the genera of plants, then we may infer that animals more widely separated in the scale of nature can be more easily crossed than in the case of plants; but the hybrids themselves are, I think, more sterile. I doubt whether any case of a perfectly fertile hybrid animal can be considered as thoroughly well authenticated. It should, however, be borne in mind that, owing to few animals breeding freely under confinement, few experiments have been fairly tried: for instance, the canary-bird has been crossed with nine other finches, but as not one of these nine species breeds freely in confinement, we have no right to expect that the first crosses between them and the canary, or that their hybrids, should be perfectly fertile. Again, with respect to the fertility in successive generations of the more fertile hybrid animals, I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents, so as to avoid the ill effects of close interbreeding [inbreeding]. On the contrary, brothers and sisters have usually been crossed in each successive generation, in opposition to the constantly repeated admonition of every breeder. And in this case, it is not at all surprising that the inherent sterility in the hybrids should have gone on increasing. If we were to act thus, and pair brothers and sisters in the case of any pure animal, which from any cause had the least tendency to sterility, the breed would assuredly be lost in a very few generations.
Although I do not know of any thoroughly well-authenticated cases of perfectly fertile hybrid animals, I have some reason to believe that the hybrids from Cervulus vaginalis [a muntjac] and Reevesii [muntjac] [...] are perfectly fertile. [Examples of fertile hybrid pheasants and geese are also given]
A doctrine which originated with Pallas, has been largely accepted by modern naturalists; namely, that most of our domestic animals have descended from two or more aboriginal species, since commingled by intercrossing. On this view, the aboriginal species must either at first have produced quite fertile hybrids, or the hybrids must have become in subsequent generations quite fertile under domestication. This latter alternative seems to me the most probable, and I am inclined to believe in its truth, although it rests on no direct evidence. I believe, for instance, that our dogs have descended from several wild stocks; yet, with perhaps the exception of certain indigenous domestic dogs of South America, all are quite fertile together; and analogy makes me greatly doubt, whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. So again there is reason to believe that our European and the humped Indian cattle are quite fertile together; but from facts communicated to me by Mr. Blyth, I think they must be considered as distinct species. On this view of the origin of many of our domestic animals, we must either give up the belief of the almost universal sterility of distinct species of animals when crossed; or we must look at sterility, not as an indelible characteristic, but as one capable of being removed by domestication.
Finally, looking to all the ascertained facts on the intercrossing of plants and animals, it may be concluded that some degree of sterility, both in first crosses and in hybrids, is an extremely general result; but that it cannot, under our present state of knowledge, be considered as absolutely universal.
LAWS GOVERNING THE STERILITY OF FIRST CROSSES AND OF HYBRIDS.
We will now consider a little more in detail the circumstances and rules governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not the rules indicate that species have specially been endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following rules and conclusions are chiefly drawn up from Gartner's admirable work on the hybridisation of plants. I have taken much pains to ascertain how far the rules apply to animals, and considering how scanty our knowledge is in regard to hybrid animals, I have been surprised to find how generally the same rules apply to both kingdoms.
It has been already remarked, that the degree of fertility, both of first crosses and of hybrids, graduates from zero to perfect fertility [the example given relates to plants].
Hybrids from two species which are very difficult to cross, and which rarely produce any offspring, are generally very sterile; but the parallelism between the difficulty of making a first cross, and the sterility of the hybrids thus produced - two classes of facts which are generally confounded together - is by no means strict. There are many cases, in which two pure species can be united with unusual facility, and produce numerous hybrid-offspring, yet these hybrids are remarkably sterile. On the other hand, there are species which can be crossed very rarely, or with extreme difficulty, but the hybrids, when at last produced, are very fertile. Even within the limits of the same genus, for instance in Dianthus [a plant], these two opposite cases occur.
The fertility, both of first crosses and of hybrids, is more easily affected by unfavourable conditions, than is the fertility of pure species. But the degree of fertility is likewise innately variable; for it is not always the same when the same two species are crossed under the same circumstances, but depends in part upon the constitution of the individuals which happen to have been chosen for the experiment. So it is with hybrids, for their degree of fertility is often found to differ greatly in the several individuals raised from seed out of the same capsule and exposed to exactly the same conditions.
By the term systematic affinity is meant, the resemblance between species in structure and in constitution, more especially in the structure of parts which are of high physiological importance and which differ little in the allied species. Now the fertility of first crosses between species, and of the hybrids produced from them, is largely governed by their systematic affinity. [The example given related to plants]
No one has been able to point out what kind, or what amount, of difference in any recognisable character is sufficient to prevent two species crossing. [This related to plants]
By a reciprocal cross between two species, I mean the case, for instance, of a stallion-horse being first crossed with a female-ass, and then a male-ass with a mare: these two species may then be said to have been reciprocally crossed. There is often the widest possible difference in the facility of making reciprocal crosses. Such cases are highly important, for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity, or of any recognisable difference in their whole organisation. On the other hand, these cases clearly show that the capacity for crossing is connected with constitutional differences imperceptible by us, and confined to the reproductive system. [...] It is also a remarkable fact, that hybrids raised from reciprocal crosses, though of course compounded of the very same two species, the one species having first been used as the father and then as the mother, generally differ in fertility in a small, and occasionally in a high degree.
Several other singular rules could be given from Gartner: for instance, some species have a remarkable power of crossing with other species; other species of the same genus have a remarkable power of impressing their likeness on their hybrid offspring; but these two powers do not at all necessarily go together. There are certain hybrids which instead of having, as is usual, an intermediate character between their two parents, always closely resemble one of them; and such hybrids, though externally so like one of their pure parent-species, are with rare exceptions extremely sterile. So again amongst hybrids which are usually intermediate in structure between their parents, exceptional and abnormal individuals sometimes are born, which closely resemble one of their pure parents; and these hybrids are almost always utterly sterile, even when the other hybrids raised from seed from the same capsule have a considerable degree of fertility. These facts show how completely fertility in the hybrid is independent of its external resemblance to either pure parent.
Considering the several rules now given, which govern the fertility of first crosses and of hybrids, we see that when forms, which must be considered as good and distinct species, are united, their fertility graduates from zero to perfect fertility, or even to fertility under certain conditions in excess. That their fertility, besides being eminently susceptible to favourable and unfavourable conditions, is innately variable. That it is by no means always the same in degree in the first cross and in the hybrids produced from this cross. That the fertility of hybrids is not related to the degree in which they resemble in external appearance either parent. And lastly, that the facility of making a first cross between any two species is not always governed by their systematic affinity or degree of resemblance to each other. This latter statement is clearly proved by reciprocal crosses between the same two species, for according as the one species or the other is used as the father or the mother, there is generally some difference, and occasionally the widest possible difference, in the facility of effecting an union. The hybrids, moreover, produced from reciprocal crosses often differ in fertility.
Now do these complex and singular rules indicate that species have been endowed with sterility simply to prevent their becoming confounded in nature? I think not. For why should the sterility be so extremely different in degree, when various species are crossed, all of which we must suppose it would be equally important to keep from blending together? Why should the degree of sterility be innately variable in the individuals of the same species? Why should some species cross with facility, and yet produce very sterile hybrids; and other species cross with extreme difficulty, and yet produce fairly fertile hybrids? Why should there often be so great a difference in the result of a reciprocal cross between the same two species? Why, it may even be asked, has the production of hybrids been permitted? to grant to species the special power of producing hybrids, and then to stop their further propagation by different degrees of sterility, not strictly related to the facility of the first union between their parents, seems to be a strange arrangement.
The foregoing rules and facts, on the other hand, appear to me clearly to indicate that the sterility both of first crosses and of hybrids is simply incidental or dependent on unknown differences, chiefly in the reproductive systems, of the species which are crossed. The differences being of so peculiar and limited a nature, that, in reciprocal crosses between two species the male sexual element of the one will often freely act on the female sexual element of the other, but not in a reversed direction. [A plant example was given and included grafting as a means of determining compatibility.]
CAUSES OF THE STERILITY OF FIRST CROSSES AND OF HYBRIDS.
We may now look a little closer at the probable causes of the sterility of first crosses and of hybrids. These two cases are fundamentally different, for, as just remarked, in the union of two pure species the male and female sexual elements are perfect, whereas in hybrids they are imperfect. Even in first crosses, the greater or lesser difficulty in effecting a union apparently depends on several distinct causes. [Plant examples are given: physical impossibility in the male penetrating the female, the male gamete being unable to penetrate the ovum, failure to fertilise the ovum, death of the embryo at an early stage] I was at first very unwilling to believe in this view [ the early death of the embryo]; as hybrids, when once born, are generally healthy and long-lived, as we see in the case of the common mule. Hybrids, however, are differently circumstanced before and after birth: when born and living in a country where their two parents can live, they are generally placed under suitable conditions of life. But a hybrid partakes of only half of the nature and constitution of its mother, and therefore before birth, as long as it is nourished within its mother's womb or within the egg or seed produced by the mother, it may be exposed to conditions in some degree unsuitable, and consequently be liable to perish at an early period; more especially as all very young beings seem eminently sensitive to injurious or unnatural conditions of life.
In regard to the sterility of hybrids, in which the sexual elements are imperfectly developed, the case is very different. I have more than once alluded to a large body of facts, which I have collected, showing that when animals and plants are removed from their natural conditions, they are extremely liable to have their reproductive systems seriously affected. This, in fact, is the great bar to the domestication of animals. Between the sterility thus superinduced and that of hybrids, there are many points of similarity. In both cases the sterility is independent of general health, and is often accompanied by excess of size or great luxuriance. In both cases, the sterility occurs in various degrees; in both, the male element is the most liable to be affected; but sometimes the female more than the male. In both, the tendency goes to a certain extent with systematic affinity, for whole groups of animals and plants are rendered impotent by the same unnatural conditions; and whole groups of species tend to produce sterile hybrids. On the other hand, one species in a group will sometimes resist great changes of conditions with unimpaired fertility; and certain species in a group will produce unusually fertile hybrids. No one can tell, till he tries, whether any particular animal will breed under confinement or any plant seed freely under culture; nor can he tell, till he tries, whether any two species of a genus will produce more or less sterile hybrids. Lastly, when organic beings are placed during several generations under conditions not natural to them, they are extremely liable to vary, which is due, as I believe, to their reproductive systems having been specially affected, though in a lesser degree than when sterility ensues. So it is with hybrids, for hybrids in successive generations are eminently liable to vary, as every experimentalist has observed.
Thus we see that when organic beings are placed under new and unnatural conditions, and when hybrids are produced by the unnatural crossing of two species, the reproductive system, independently of the general state of health, is affected by sterility in a very similar manner. In the one case, the conditions of life have been disturbed, though often in so slight a degree as to be inappreciable by us; in the other case, or that of hybrids, the external conditions have remained the same, but the organisation has been disturbed by two different structures and constitutions having been blended into one. For it is scarcely possible that two organisations should be compounded into one, without some disturbance occurring in the development, or periodical action, or mutual relation of the different parts and organs one to another, or to the conditions of life. When hybrids are able to breed inter se [among themselves], they transmit to their offspring from generation to generation the same compounded organisation, and hence we need not be surprised that their sterility, though in some degree variable, rarely diminishes.
It must, however, be confessed that we cannot understand, excepting on vague hypotheses, several facts with respect to the sterility of hybrids; for instance, the unequal fertility of hybrids produced from reciprocal crosses; or the increased sterility in those hybrids which occasionally and exceptionally resemble closely either pure parent. Nor do I pretend that the foregoing remarks go to the root of the matter: no explanation is offered why an organism, when placed under unnatural conditions, is rendered sterile. All that I have attempted to show, is that in two cases, in some respects allied, sterility is the common result, - in the one case from the conditions of life having been disturbed, in the other case from the organisation having been disturbed by two organisations having been compounded into one.
It may seem fanciful, but I suspect that a similar parallelism extends to an allied yet very different class of facts. It is an old and almost universal belief, founded, I think, on a considerable body of evidence, that slight changes in the conditions of life are beneficial to all living things. We see this acted on by farmers and gardeners in their frequent exchanges of seed, tubers, etc., from one soil or climate to another, and back again. During the convalescence of animals, we plainly see that great benefit is derived from almost any change in the habits of life. Again, both with plants and animals, there is abundant evidence, that a cross between very distinct individuals of the same species, that is between members of different strains or sub-breeds, gives vigour and fertility to the offspring. I believe, indeed, from the facts alluded to in our fourth chapter, that a certain amount of crossing is indispensable even with hermaphrodites; and that close interbreeding continued during several generations between the nearest relations, especially if these be kept under the same conditions of life, always induces weakness and sterility in the progeny.
Hence it seems that, on the one hand, slight changes in the conditions of life benefit all organic beings, and on the other hand, that slight crosses, that is crosses between the males and females of the same species which have varied and become slightly different, give vigour and fertility to the offspring. But we have seen that greater changes, or changes of a particular nature, often render organic beings in some degree sterile; and that greater crosses, that is crosses between males and females which have become widely or specifically different, produce hybrids which are generally sterile in some degree. I cannot persuade myself that this parallelism is an accident or an illusion. Both series of facts seem to be connected together by some common but unknown bond, which is essentially related to the principle of life.
FERTILITY OF VARIETIES WHEN CROSSED, AND OF THEIR MONGREL OFFSPRING.
It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. I fully admit that this is almost invariably the case. But if we look to varieties produced under nature, we are immediately involved in hopeless difficulties; for if two hitherto reputed varieties be found in any degree sterile together, they are at once ranked by most naturalists as species.
If we turn to varieties, produced, or supposed to have been produced, under domestication, we are still involved in doubt. For when it is stated, for instance, that the German Spitz dog unites more easily than other dogs with foxes, or that certain South American indigenous domestic dogs do not readily cross with European dogs, the explanation which will occur to everyone, and probably the true one, is that these dogs have descended from several aboriginally distinct species. Nevertheless the perfect fertility of so many domestic varieties, differing widely from each other in appearance, for instance of the pigeon or of the cabbage, is a remarkable fact; more especially when we reflect how many species there are, which, though resembling each other most closely, are utterly sterile when intercrossed. Several considerations, however, render the fertility of domestic varieties less remarkable than at first appears. It can, in the first place, be clearly shown that mere external dissimilarity between two species does not determine their greater or lesser degree of sterility when crossed; and we may apply the same rule to domestic varieties. In the second place, some eminent naturalists believe that a long course of domestication tends to eliminate sterility in the successive generations of hybrids, which were at first only slightly sterile; and if this be so, we surely ought not to expect to find sterility both appearing and disappearing under nearly the same conditions of life. Lastly, and this seems to me by far the most important consideration, new races of animals and plants are produced under domestication by man's methodical and unconscious power of selection, for his own use and pleasure: he neither wishes to select, nor could select, slight differences in the reproductive system, or other constitutional differences correlated with the reproductive system. He supplies his several varieties with the same food; treats them in nearly the same manner, and does not wish to alter their general habits of life. Nature acts uniformly and slowly during vast periods of time on the whole organisation, in any way which may be for each creature's own good; and thus she may, either directly, or more probably indirectly, through correlation, modify the reproductive system in the several descendants from any one species. Seeing this difference in the process of selection, as carried on by man and nature, we need not be surprised at some difference in the result.
I have as yet spoken as if the varieties of the same species were invariably fertile when intercrossed. But it seems to me impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases, which I will briefly abstract. The evidence is at least as good as that from which we believe in the sterility of a multitude of species. The evidence is, also, derived from hostile witnesses, who in all other cases consider fertility and sterility as safe criterions of specific distinction. [Plant examples are given where supposed species are interfertile and their hybrids are fertile]
From these facts; from the great difficulty of ascertaining the infertility of varieties in a state of nature, for a supposed variety if infertile in any degree would generally be ranked as species; from man selecting only external characters in the production of the most distinct domestic varieties, and from not wishing or being able to produce recondite and functional differences in the reproductive system; from these several considerations and facts, I do not think that the very general fertility of varieties can be proved to be of universal occurrence, or to form a fundamental distinction between varieties and species. The general fertility of varieties does not seem to me sufficient to overthrow the view which I have taken with respect to the very general, but not invariable, sterility of first crosses and of hybrids, namely, that it is not a special endowment, but is incidental on slowly acquired modifications, more especially in the reproductive systems of the forms which are crossed.
HYBRIDS AND MONGRELS COMPARED, INDEPENDENTLY OF THEIR FERTILITY.
Independently of the question of fertility, the offspring of species when crossed and of varieties when crossed may be compared in several other respects. Gartner, whose strong wish was to draw a marked line of distinction between species and varieties, could find very few and, as it seems to me, quite unimportant differences between the so-called hybrid offspring of species, and the so-called mongrel offspring of varieties. And, on the other hand, they agree most closely in very many important respects.
I shall here discuss this subject with extreme brevity. The most important distinction is, that in the first generation mongrels are more variable than hybrids; but Gartner admits that hybrids from species which have long been cultivated are often variable in the first generation; and I have myself seen striking instances of this fact. Gartner further admits that hybrids between very closely allied species are more variable than those from very distinct species; and this shows that the difference in the degree of variability graduates away. When mongrels and the more fertile hybrids are propagated for several generations an extreme amount of variability in their offspring is notorious; but some few cases both of hybrids and mongrels long retaining uniformity of character could be given. The variability, however, in the successive generations of mongrels is, perhaps, greater than in hybrids.
This greater variability of mongrels than of hybrids does not seem to me at all surprising. For the parents of mongrels are varieties, and mostly domestic varieties (very few experiments having been tried on natural varieties), and this implies in most cases that there has been recent variability; and therefore we might expect that such variability would often continue and be super-added to that arising from the mere act of crossing. The slight degree of variability in hybrids from the first cross or in the first generation, in contrast with their extreme variability in the succeeding generations, is a curious fact and deserves attention. For it bears on and corroborates the view which I have taken on the cause of ordinary variability; namely, that it is due to the reproductive system being eminently sensitive to any change in the conditions of life, being thus often rendered either impotent or at least incapable of its proper function of producing offspring identical with the parent-form. Now hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected, and they are not variable; but hybrids themselves have their reproductive systems seriously affected, and their descendants are highly variable.
But to return to our comparison of mongrels and hybrids: Gartner states that mongrels are more liable than hybrids to revert to either parent-form; but this, if it be true, is certainly only a difference in degree. Gartner further insists that when any two species, although most closely allied to each other, are crossed with a third species, the hybrids are widely different from each other; whereas if two very distinct varieties of one species are crossed with another species, the hybrids do not differ much. But this conclusion, as far as I can make out, is founded on a single experiment; and seems directly opposed to the results of several experiments made by Kolreuter.
These alone are the unimportant differences, which Gartner is able to point out, between hybrid and mongrel plants. On the other hand, the resemblance in mongrels and in hybrids to their respective parents, more especially in hybrids produced from nearly related species, follows according to Gartner the same laws. When two species are crossed, one has sometimes a prepotent power of impressing its likeness on the hybrid; and so I believe it to be with varieties of plants. With animals one variety certainly often has this prepotent power over another variety. Hybrid plants produced from a reciprocal cross, generally resemble each other closely; and so it is with mongrels from a reciprocal cross. Both hybrids and mongrels can be reduced to either pure parent-form, by repeated crosses in successive generations with either parent.
These several remarks are apparently applicable to animals; but the subject is here excessively complicated, partly owing to the existence of secondary sexual characters; but more especially owing to prepotency in transmitting likeness running more strongly in one sex than in the other, both when one species is crossed with another, and when one variety is crossed with another variety. For instance, I think those authors are right, who maintain that the ass has a prepotent power over the horse, so that both the mule and the hinny more resemble the ass than the horse; but that the prepotency runs more strongly in the male-ass than in the female, so that the mule, which is the offspring of the male-ass and mare, is more like an ass, than is the hinny, which is the offspring of the female-ass and stallion.
Much stress has been laid by some authors on the supposed fact, that mongrel animals alone are born closely like one of their parents; but it can be shown that this does sometimes occur with hybrids; yet I grant much less frequently with hybrids than with mongrels. Looking to the cases which I have collected of cross-bred animals closely resembling one parent, the resemblances seem chiefly confined to characters almost monstrous in their nature, and which have suddenly appeared - such as albinism, melanism, deficiency of tail or horns, or additional fingers and toes; and do not relate to characters which have been slowly acquired by selection. Consequently, sudden reversions to the perfect character of either parent would be more likely to occur with mongrels, which are descended from varieties often suddenly produced and semi-monstrous in character, than with hybrids, which are descended from species slowly and naturally produced. On the whole I entirely agree with Dr. Prosper Lucas, who, after arranging an enormous body of facts with respect to animals, comes to the conclusion, that the laws of resemblance of the child to its parents are the same, whether the two parents differ much or little from each other, namely in the union of individuals of the same variety, or of different varieties, or of distinct species.
Laying aside the question of fertility and sterility, in all other respects there seems to be a general and close similarity in the offspring of crossed species, and of crossed varieties. If we look at species as having been specially created, and at varieties as having been produced by secondary laws, this similarity would be an astonishing fact. But it harmonises perfectly with the view that there is no essential distinction between species and varieties.
SUMMARY OF CHAPTER.
First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived, have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of sterility does not strictly follow systematic affinity, but is governed by several curious and complex laws. It is generally different, and sometimes widely different, in reciprocal crosses between the same two species. It is not always equal in degree in a first cross and in the hybrid produced from this cross.
The sterility of first crosses between pure species, which have their reproductive systems perfect, seems to depend on several circumstances; in some cases largely on the early death of the embryo. The sterility of hybrids, which have their reproductive systems imperfect, and which have had this system and their whole organisation disturbed by being compounded of two distinct species, seems closely allied to that sterility which so frequently affects pure species, when their natural conditions of life have been disturbed. This view is supported by a parallelism of another kind; - namely, that the crossing of forms only slightly different is favourable to the vigour and fertility of their offspring; and that slight changes in the conditions of life are apparently favourable to the vigour and fertility of all organic beings. It is not surprising that the degree of difficulty in uniting two species, and the degree of sterility of their hybrid-offspring should generally correspond, though due to distinct causes; for both depend on the amount of difference of some kind between the species which are crossed. Nor is it surprising that the facility of effecting a first cross, the fertility of the hybrids produced, [and ability to be grafted together in plants] should all run, to a certain extent, parallel with the systematic affinity of the forms which are subjected to experiment; for systematic affinity attempts to express all kinds of resemblance between all species.
First crosses between forms known to be varieties, or sufficiently alike to be considered as varieties, and their mongrel offspring, are very generally, but not quite universally, fertile. Nor is this nearly general and perfect fertility surprising, when we remember how liable we are to argue in a circle with respect to varieties in a state of nature; and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences, and not of differences in the reproductive system. In all other respects, excluding fertility, there is a close general resemblance between hybrids and mongrels. Finally, then, the facts briefly given in this chapter do not seem to me opposed to, but even rather to support the view, that there is no fundamental distinction between species and varieties.
EXCERPTS FROM "DARWINISM AN EXPOSITION OF THE THEORY OF NATURAL SELECTION WITH SOME OF ITS APPLICATIONS"
BY ALFRED RUSSEL WALLACE (1889)
One of the greatest, or perhaps we may say the greatest, of all the difficulties in the way of accepting the theory of natural selection as a complete explanation of the origin of species, has been the remarkable difference between varieties and species in respect of fertility when crossed. Generally speaking, it may be said that the varieties of any one species, however different they may be in external appearance, are perfectly fertile when crossed, and their mongrel offspring are equally fertile when bred among themselves; while distinct species, on the other hand, however closely they may resemble each other externally, are usually infertile when crossed, and their hybrid offspring absolutely sterile. This used to be considered a fixed law of nature, constituting the absolute test and criterion of a species as distinct from a variety; and so long as it was believed that species were separate creations, or at all events had an origin quite distinct from that of varieties, this law could have no exceptions, because, if any two species had been found to be fertile when crossed and their hybrid offspring to be also fertile, this fact would have been held to prove them to be not species but varieties. On the other hand, if two varieties had been found to be infertile, or their mongrel offspring to be sterile, then it would have been said: These are not varieties but true species. Thus the old theory led to inevitable reasoning in a circle; and what might be only a rather common fact was elevated into a law which had no exceptions.
The elaborate and careful examination of the whole subject by Mr. Darwin, who has brought together a vast mass of evidence from the experience of agriculturists and horticulturists, as well as from scientific experimenters, has demonstrated that there is no such fixed law in nature as was formerly supposed. He shows us that crosses between some varieties are infertile or even sterile, while crosses between some species are quite fertile; and that there are besides a number of curious phenomena connected with the subject which render it impossible to believe that sterility is anything more than an incidental property of species, due to the extreme delicacy and susceptibility of the reproductive powers, and dependent on physiological causes we have not yet been able to trace. Nevertheless, the fact remains that most species which have hitherto been crossed produce sterile hybrids, as in the well-known case of the mule; while almost all domestic varieties, when crossed, produce offspring which are perfectly fertile among themselves. I will now endeavour to give such a sketch of the subject as may enable the reader to see something of the complexity of the problem, referring him to Mr. Darwin's works for fuller details.
EXTREME SUSCEPTIBILITY OF THE REPRODUCTIVE FUNCTIONS.
One of the most interesting facts, as showing how susceptible to changed conditions or to slight constitutional changes are the reproductive powers of animals, is the very general difficulty of getting those which are kept in confinement to breed; and this is frequently the only bar to domesticating wild species. Thus, elephants, bears, foxes, and numbers of species of rodents, very rarely breed in confinement; while other species do so more or less freely. [...] This inability to reproduce is not due to ill-health, since many of these creatures are perfectly vigorous and live very long.
With our true domestic animals, on the other hand, fertility is perfect, and is very little affected by changed conditions. [...] It therefore seems probable, that this facility for breeding under changed conditions was an original property of the species which man has domesticated - a property which, more than any other, enabled him to domesticate them. Yet, even with these, there is evidence that great changes of conditions affect the fertility. In the hot valleys of the Andes sheep are less fertile; while geese taken to the high plateau of Bogota were at first almost sterile, but after some generations recovered their fertility. These and many other facts seem to show that, with the majority of animals, even a slight change of conditions may produce infertility or sterility; and also that after a time, when the animal has become thoroughly acclimatised, as it were, to the new conditions, the infertility is in some cases diminished or altogether ceases. It is stated by Bechstein that the canary was long infertile, and it is only of late years that good breeding birds have become common; but in this case no doubt selection has aided the change.
Another indication of the extreme delicacy of the adjustment between the sexes, which is necessary to produce fertility, is afforded by the behaviour of many species and varieties when reciprocally crossed.
INDIVIDUAL DIFFERENCES IN RESPECT TO CROSS-FERTILISATION.
A still more remarkable illustration of the delicate balance of organisation needful for reproduction, is afforded by the individual differences of animals and plants, as regards both their power of intercrossing with other individuals or other species, and the fertility of the offspring thus produced. Among domestic animals, Darwin states that it is by no means rare to find certain males and females which will not breed together, though both are known to be perfectly fertile with other males and females. Cases of this kind have occurred among horses, cattle, pigs, dogs, and pigeons; and the experiment has been tried so frequently that there can be no doubt of the fact. Professor G.J. Romanes states that he has a number of additional cases of this individual incompatibility, or of absolute sterility, between two individuals, each of which is perfectly fertile with other individuals.
CASES OF THE FERTILITY OF HYBRIDS, AND OF THE INFERTILITY OF MONGRELS.
I now propose to adduce a few cases in which it has been proved, by experiment, that hybrids between two distinct species are fertile inter se; and then to consider why it is that such cases are so few in number. [Case 1 is the common domestic goose (Anser ferns) and the Chinese goose (A. cygnoides)]
Another equally striking case is that of the Indian humped and the common cattle, species which differ osteologically, and also in habits, form, voice, and constitution, so that they are by no means closely allied; yet Mr. Darwin assures us that he has received decisive evidence that the hybrids between these are perfectly fertile inter se.
Dogs have been frequently crossed with wolves and with jackals, and their hybrid offspring have been found to be fertile inter se [among themselves] to the third or fourth generation, and then usually to show some signs of sterility or of deterioration. The wolf and dog may be originally the same species, but the jackal is certainly distinct; and the appearance of infertility or of weakness is probably due to the fact that, in almost all these experiments, the offspring of a single pair - themselves usually from the same litter - - were bred in-and-in, and this alone sometimes produces the most deleterious effects. Thus, Mr. Low in his great work on the Domesticated Animals of Great Britain, says: "If we shall breed a pair of dogs from the same litter, and unite again the offspring of this pair, we shall produce at once a feeble race of creatures; and the process being repeated for one or two generations more, the family will die out, or be incapable of propagating their race. A gentleman of Scotland made the experiment on a large scale with certain foxhounds, and he found that the race actually became monstrous and perished utterly." The same writer tells us that hogs have been made the subject of similar experiments: "After a few generations the victims manifest the change induced in the system. They become of diminished size; the bristles are changed into hairs; the limbs become feeble and short; the litters diminish in frequency, and in the number of the young produced; the mother becomes unable to nourish them, and, if the experiment be carried as far as the case will allow, the feeble, and frequently monstrous offspring, will be incapable of being reared up, and the miserable race will utterly perish."
These precise statements, by one of the greatest authorities on our domesticated animals, are sufficient to show that the fact of infertility or degeneracy appearing in the offspring of hybrids after a few generations need not be imputed to the fact of the first parents being distinct species, since exactly the same phenomena appear when individuals of the same species are bred under similar adverse conditions. But in almost all the experiments that have hitherto been made in crossing distinct species, no care has been taken to avoid close interbreeding by securing several hybrids from quite distinct stocks to start with, and by having two or more sets of experiments carried on at once, so that crosses between the hybrids produced may be occasionally made. Till this is done no experiments, such as those hitherto tried, can be held to prove that hybrids are in all cases infertile inter se [among themselves].
It has, however, been denied by Mr. A.H. Huth, in his interesting work on The Marriage of Near Kin, that any amount of breeding in-and-in is in itself hurtful; and he quotes the evidence of numerous breeders whose choicest stocks have always been so bred, as well as cases like the Porto Santo rabbits, the goats of Juan Fernandez, and other cases in which animals allowed to run wild have increased prodigiously and continued in perfect health and vigour, although all derived from a single pair. But in all these cases there has been rigid selection by which the weak or the infertile have been eliminated, and with such selection there is no doubt that the ill effects of close interbreeding can be prevented for a long time; but this by no means proves that no ill effects are produced. Mr. Huth himself quotes M. Allié, M. Aubé, Stephens, Giblett, Sir John Sebright, Youatt, Druce, Lord Weston, and other eminent breeders, as finding from experience that close interbreeding does produce bad effects; and it cannot be supposed that there would be such a consensus of opinion on this point if the evil were altogether imaginary. Mr. Huth argues, that the evil results which do occur do not depend on the close interbreeding itself, but on the tendency it has to perpetuate any constitutional weakness or other hereditary taints; and he attempts to prove this by the argument that "if crosses act by virtue of being a cross, and not by virtue of removing an hereditary taint, then the greater the difference between the two animals crossed the more beneficial will that act be." He then shows that, the wider the difference the less is the benefit, and concludes that a cross, as such, has no beneficial effect. A parallel argument would be, that change of air, as from inland to the sea-coast, or from a low to an elevated site, is not beneficial in itself, because, if so, a change to the tropics or to the polar regions should be more beneficial. In both these cases it may well be that no benefit would accrue to a person in perfect health; but then there is no such thing as "perfect health" in man, and probably no such thing as absolute freedom from constitutional taint in animals. The experiments of Mr. Darwin, showing the great and immediate good effects of a cross between distinct strains in plants, cannot be explained away; neither can the innumerable arrangements to secure cross-fertilisation by insects, the real use and purport of which will be discussed in our eleventh chapter. On the whole, then, the evidence at our command proves that, whatever may be its ultimate cause, close interbreeding does usually produce bad results; and it is only by the most rigid selection, whether natural or artificial, that the danger can be altogether obviated.
FERTILE HYBRIDS AMONG ANIMALS.
One or two more cases of fertile hybrids may be given before we pass on to the corresponding experiments in plants. Professor Alfred Newton received from a friend a pair of hybrid ducks, bred from a common duck (Anas boschas), and a pintail (Dafila acuta). From these he obtained four ducklings, but these latter, when grown up, proved infertile, and did not breed again. In this case we have the results of close interbreeding, with too great a difference between the original species, combining to produce infertility, yet the fact of a hybrid from such a pair producing healthy offspring is itself noteworthy.
Still more extraordinary is the following statement of Mr. Low: "It has been long known to shepherds, though questioned by naturalists, that the progeny of the cross between the sheep and goat is fertile. Breeds of this mixed race are numerous in the north of Europe." Nothing appears to be known of such hybrids either in Scandinavia or in Italy; but Professor Giglioli of Florence has kindly given me some useful references to works in which they are described. The following extract from his letter is very interesting: "I need not tell you that there being such hybrids is now generally accepted as a fact. Buffon (Supplements, tom. iii. p. 7, 1756) obtained one such hybrid in 1751 and eight in 1752. Sanson (La Culture, vol. vi. p. 372, 1865) mentions a case observed in the Vosges, France. Geoff. St. Hilaire (Hist. Nat. Gén. des reg. org., vol. iii. p. 163) was the first to mention, I believe, that in different parts of South America the ram is more usually crossed with the she-goat than the sheep with the he-goat. The well-known 'pellones' of Chile are produced by the second and third generation of such hybrids (Gay, 'Hist, de Chile,' vol. i. p. 466, Agriculture, 1862). Hybrids bred from goat and sheep are called 'chabin' in French, and 'cabruno' in Spanish. In Chile such hybrids are called 'carneros lanudos'; their breeding inter se [among themselves] appears to be not always successful, and often the original cross has to be recommenced to obtain the proportion of three-eighths of he-goat and five-eighths of sheep, or of three-eighths of ram and five-eighths of she-goat; such being the reputed best hybrids."
With these numerous facts recorded by competent observers we can hardly doubt that races of hybrids between these very distinct species have been produced, and that such hybrids are fairly fertile inter se [among themselves]; and the analogous facts already given lead us to believe that whatever amount of infertility may at first exist could be eliminated by careful selection, if the crossed races were bred in large numbers and over a considerable area of country. This case is especially valuable, as showing how careful we should be in assuming the infertility of hybrids when experiments have been made with the progeny of a single pair, and have been continued only for one or two generations.
Among insects one case only appears to have been recorded. The hybrids of two moths (Bombyx cynthia and B. arrindia) were proved in Paris, according to M. Quatrefages, to be fertile inter se [among themselves] for eight generations.
CASES OF STERILITY OF MONGRELS.
The reverse phenomenon to the fertility of hybrids, the sterility of mongrels or of the crosses between varieties of the same species, is a comparatively rare one, yet some undoubted cases have occurred [plant examples were given].
No cases of this kind are recorded among animals; but this is not to be wondered at, when we consider how very few experiments have been made with natural varieties; while there is good reason for believing that domestic varieties are exceptionally fertile, partly because one of the conditions of domestication was fertility under changed conditions, and also because long continued domestication is believed to have the effect of increasing fertility and eliminating whatever sterility may exist. This is shown by the fact that, in many cases, domestic animals are descended from two or more distinct species. This is almost certainly the case with the dog, and probably with the hog, the ox, and the sheep; yet the various breeds are now all perfectly fertile, although we have every reason to suppose that there would be some degree of infertility if the several aboriginal species were crossed together for the first time.
REMARKS ON THE FACTS OF HYBRIDITY.
The facts that have now been adduced, though not very numerous, are sufficiently conclusive to prove that the old belief, of the universal sterility of hybrids and fertility of mongrels, is incorrect. The doctrine that such a universal law existed was never more than a plausible generalisation, founded on a few inconclusive facts derived from domesticated animals and cultivated plants. The facts were, and still are, inconclusive for several reasons. They are founded, primarily, on what occurs among animals in domestication; and it has been shown that domestication both tends to increase fertility, and was itself rendered possible by the fertility of those particular species being little affected by changed conditions. The exceptional fertility of all the varieties of domesticated animals does not prove that a similar fertility exists among natural varieties. In the next place, the generalisation is founded on too remote crosses, as in the case of the horse and the ass, the two most distinct and widely separated species of the genus Equus, so distinct indeed that they have been held by some naturalists to form distinct genera. Crosses between the two species of zebra, or even between the zebra and the quagga, or the quagga and the ass, might have led to a very different result. Again, in pre-Darwinian times it was so universally the practice to argue in a circle, and declare that the fertility of the offspring of a cross proved the identity of species of the parents, that experiments in hybridity were usually made between very remote species and even between species of different genera, to avoid the possibility of the reply: "They are both really the same species;" and the sterility of the hybrid offspring of such remote crosses of course served to strengthen the popular belief.
Now that we have arrived at a different standpoint, and look upon a species, not as a distinct entity due to special creation, but as an assemblage of individuals which have become somewhat modified in structure, form, and constitution so as to adapt them to slightly different conditions of life; which can be differentiated from other allied assemblages; which reproduce their like, and which usually breed together - we require a fresh set of experiments calculated to determine the matter of fact, - whether such species crossed with their near allies do always produce offspring which are more or less sterile inter se [among themselves]. Ample materials for such experiments exist, in the numerous "representative species" inhabiting distinct areas on a continent or different islands of a group; or even in those found in the same area but frequenting somewhat different stations.
To carry out these experiments with any satisfactory result, it will be necessary to avoid the evil effects of confinement and of too close interbreeding. If birds are experimented with, they should be allowed as much liberty as possible, a plot of ground with trees and bushes being enclosed with wire netting overhead so as to form a large open aviary. The species experimented with should be obtained in considerable numbers, and by two separate persons, each making the opposite reciprocal cross, as explained at p. 155. In the second generation these two stocks might be themselves crossed to prevent the evil effects of too close interbreeding. By such experiments, carefully carried out with different groups of animals and plants, we should obtain a body of facts of a character now sadly wanting, and without which it is hopeless to expect to arrive at a complete solution of this difficult problem. There are, however, some other aspects of the question that need to be considered, and some theoretical views which require to be carefully examined, having done which we shall be in a condition to state the general conclusions to which the facts and reasonings at our command seem to point.
STERILITY DUE TO CHANGED CONDITIONS AND USUALLY CORRELATED WITH OTHER CHARACTERS, ESPECIALLY WITH COLOUR.
The evidence already adduced as to the extreme susceptibility of the reproductive system, and the curious irregularity with which infertility or sterility appears in the crosses between some varieties or species while quite absent in those between others, seem to indicate that sterility is a characteristic which has a constant tendency to appear, either by itself or in correlation with other characters. It is known to be especially liable to occur under changed conditions of life; and, as such change is usually the starting-point and cause of the development of new species, we have already found a reason why it should so often appear when species become fully differentiated.
In almost all the cases of infertility or sterility between varieties or species, we have some external differences with which it is correlated; and though these differences are sometimes slight, and the amount of the infertility is not always, or even usually, proportionate to the external difference between the two forms crossed, we must believe that there is some connection between the two classes of facts. This is especially the case as regards colour; and Mr. Darwin has collected a body of facts which go far to prove that colour, instead of being an altogether trifling and unimportant character, as was supposed by the older naturalists, is really one of great significance, since it is undoubtedly often correlated with important constitutional differences [plant examples are given].
Similar phenomena have not been recorded among animals; but this is not to be wondered at when we consider that most of our pure and valued domestic breeds are characterised by definite colours which constitute one of their distinctive marks, and they are, therefore, seldom crossed with these of another colour; and even when they are so crossed, no notice would be taken of any slight diminution of fertility, since this is liable to occur from many causes. We have also reason to believe that fertility has been increased by long domestication, in addition to the fact of the original stocks being exceptionally fertile; and no experiments have been made on the differently coloured varieties of wild animals. There are, however, a number of very curious facts showing that colour in animals, as in plants, is often correlated with constitutional differences of a remarkable kind, and as these have a close relation to the subject we are discussing, a brief summary of them will be here given.
CORRELATION OF COLOUR WITH CONSTITUTIONAL PECULIARITIES.
The correlation of a white colour and blue eyes in male cats with deafness, and of the tortoise-shell marking with the female sex of the same animal, are two well-known but most extraordinary cases. Equally remarkable is the fact, communicated to Darwin by Mr. Tegetmeier, that white, yellow, pale blue, or dun pigeons, of all breeds, have the young birds born naked, while in all other colours they are well covered with down. Here we have a case in which colour seems of more physiological importance than all the varied structural differences between the varieties and breeds of pigeons. In Virginia there is a plant called the paint-root (Lachnanthes tinctoria), which, when eaten by pigs, colours their bones pink, and causes the hoofs of all but the black varieties to drop off; so that black pigs only can be kept in the district. Buckwheat in flower is also said to be injurious to white pigs but not to black. In the Tarentino, black sheep are not injured by eating the Hypericum crispum - a species of St. John's-wort - which kills white sheep. White terriers suffer most from distemper; white chickens from the gapes. White-haired horses or cattle are subject to cutaneous diseases from which the dark coloured are free; while, both in Thuringia and the West Indies, it has been noticed that white or pale coloured cattle are much more troubled by flies than are those which are brown or black.
These curious and inexplicable correlations of colour with constitutional peculiarities, both in animals and plants, render it probable that the correlation of colour with infertility, which has been detected in several cases in plants, may also extend to animals in a state of nature; and if so, the fact is of the highest importance as throwing light on the origin of the infertility of many allied species. This will be better understood after considering the facts which will be now described.
Footnote to section: In the Medico-Chirurgical Transactions, vol. liii. (1870), Dr. Ogle has adduced some curious physiological facts bearing on the presence or absence of white colours in the higher animals. He states that a dark pigment in the olfactory region of the nostrils is essential to perfect smell, and that this pigment is rarely deficient except when the whole animal is pure white, and the creature is then almost without smell or taste. He observes that there is no proof that, in any of the cases given above, the black animals actually eat the poisonous root or plant; and that the facts are readily understood if the senses of smell and taste are dependent on a pigment which is absent in the white animals, who therefore eat what those gifted with normal senses avoid. This explanation however hardly seems to cover the facts. We cannot suppose that almost all the sheep in the world (which are mostly white) are without smell or taste. The cutaneous disease on the white patches of hair on horses, the special liability of white terriers to distemper, of white chickens to the gapes, and of silkworms which produce yellow silk to the fungus, are not explained by it. The analogous facts in plants also indicate a real constitutional relation with colour, not an affection of the sense of smell and taste only.
THE ISOLATION OF VARIETIES BY SELECTIVE ASSOCIATION.
In the last chapter I have shown that the importance of geographical isolation for the formation of new species by natural selection has been greatly exaggerated, because the very change of conditions, which is the initial power in starting such new forms, leads also to a local or stational segregation of the forms acted upon. But there is also a very powerful cause of isolation in the mental nature - the likes and dislikes - of animals; and to this is probably due the fact of the comparative rarity of hybrids in a state of nature. The differently coloured herds of cattle in the Falkland Islands, each of which keeps separate, have been already mentioned; and it may be added, that the mouse-coloured variety seem to have already developed a physiological peculiarity in breeding a month earlier than the others. Similar facts occur, however, among our domestic animals and are well known to breeders. Professor Low, one of the greatest authorities on our domesticated animals, says: "The female of the dog, when not under restraint, makes selection of her mate, the mastiff selecting the mastiff, the terrier the terrier, and so on." And again: "The Merino sheep and Heath sheep of Scotland, if two flocks are mixed together, each will breed with its own variety." Mr. Darwin has collected many facts illustrating this point. One of the chief pigeon-fanciers in England informed him that, if free to choose, each breed would prefer pairing with its own kind. Among the wild horses in Paraguay those of the same colour and size associate together; while in Circassia there are three races of horses which have received special names, and which, when living a free life, almost always refuse to mingle and cross, and will even attack one another. On one of the Faroe Islands, not more than half a mile in diameter, the half-wild native black sheep do not readily mix with imported white sheep. In the Forest of Dean, and in the New Forest, the dark and pale coloured herds of fallow deer have never been known to mingle; and even the curious Ancon sheep of quite modern origin have been observed to keep together, separating themselves from the rest of the flock when put into enclosures with other sheep. The same rule applies to birds, for Darwin was informed by the Rev. W.D. Fox that his flocks of white and Chinese geese kept distinct. [Note: some of this is wishful thinking as dog breeds and horse breeds will both mate indiscriminately!]
This constant preference of animals for their like, even in the case of slightly different varieties of the same species, is evidently a fact of great importance in considering the origin of species by natural selection, since it shows us that, so soon as a slight differentiation of form or colour has been effected, isolation will at once arise by the selective association of the animals themselves; and thus the great stumbling-block of "the swamping effects of intercrossing," which has been so prominently brought forward by many naturalists, will be completely obviated.
If now we combine with this fact the correlation of colour with important constitutional peculiarities, and, in some cases, with infertility; and consider, further, the curious parallelism that has been shown to exist between the effects of changed conditions and the intercrossing of varieties in producing either an increase or a decrease of fertility, we shall have obtained, at all events, a starting-point for the production of that infertility which is so characteristic a feature of distinct species when intercrossed. All we need, now, is some means of increasing or accumulating this initial tendency; and to a discussion of this problem we will therefore address ourselves.
THE INFLUENCE OF NATURAL SELECTION UPON STERILITY AND FERTILITY.
It will occur to many persons that, as the infertility or sterility of incipient species would be useful to them when occupying the same or adjacent areas, by neutralising the effects of intercrossing, this infertility might have been increased by the action of natural selection; and this will be thought the more probable if we admit, as we have seen reason to do, that variations in fertility occur, perhaps as frequently as other variations. Mr. Darwin tells us that, at one time, this appeared to him probable, but he found the problem to be one of extreme complexity; and he was also influenced against the view by many considerations which seemed to render such an origin of the sterility or infertility of species when intercrossed very improbable. The fact that species which occupy distinct areas, and which nowhere come in contact with each other, are often sterile when crossed, is one of the difficulties; but this may perhaps be overcome by the consideration that, though now isolated, they may, and often must, have been in contact at their origination. More important is the objection that natural selection could not possibly have produced the difference that often occurs between reciprocal crosses, one of these being sometimes fertile, while the other is sterile. The extremely different amounts of infertility or sterility between different species of the same genus, the infertility often bearing no proportion to the difference between the species crossed, is also an important objection. But none of these objections would have much weight if it could be clearly shown that natural selection is able to increase the infertility variations of incipient species, as it is certainly able to increase and develop all useful variations of form, structure, instincts, or habits. Ample causes of infertility have been shown to exist, in the nature of the organism and the laws of correlation; the agency of natural selection is only needed to accumulate the effects produced by these causes, and to render their final results more uniform and more in accordance with the facts that exist.
About twenty years ago I had much correspondence and discussion with Mr. Darwin on this question. I then believed that I was able to demonstrate the action of natural selection in accumulating infertility; but I could not convince him, owing to the extreme complexity of the process under the conditions which he thought most probable. I have recently returned to the question; and, with the fuller knowledge of the facts of variation we now possess, I think it may be shown that natural selection is, in some probable cases at all events, able to accumulate variations in infertility between incipient species.
The simplest case to consider, will be that in which two forms or varieties of a species, occupying an extensive area, are in process of adaptation to somewhat different modes of life within the same area. If these two forms freely intercross with each other, and produce mongrel offspring which are quite fertile inter se [among themselves], then the further differentiation of the forms into two distinct species will be retarded, or perhaps entirely prevented; for the offspring of the crossed unions will be, perhaps, more vigorous on account of the cross, although less perfectly adapted to the conditions of existence than either of the pure breeds; and this would certainly establish a powerful antagonistic influence to the further differentiation of the two forms.
Now, let us suppose that a partial sterility of the hybrids between the two forms arises, in correlation with the different modes of life and the slight external or internal peculiarities that exist between them, both of which we have seen to be real causes of infertility. The result will be that, even if the hybrids between the two forms are still freely produced, these hybrids will not themselves increase so rapidly as the two pure forms; and as these latter are, by the terms of the problem, better suited to their conditions of life than are the hybrids between them, they will not only increase more rapidly, but will also tend to supplant the hybrids altogether whenever the struggle for existence becomes exceptionally severe. Thus, the more complete the sterility of the hybrids the more rapidly will they die out and leave the two parent forms pure. Hence it will follow that, if there is greater infertility between the two forms in one part of the area than the other, these forms will be kept more pure wherever this greater infertility prevails, will therefore have an advantage at each recurring period of severe struggle for existence, and will thus ultimately supplant the less infertile or completely fertile forms that may exist in other portions of the area. It thus appears that, in such a case as here supposed, natural selection would preserve those portions of the two breeds which were most infertile with each other, or whose hybrid offspring were most infertile; and would, therefore, if variations in fertility continued to arise, tend to increase that infertility. It must particularly be noted that this effect would result, not by the preservation of the infertile variations on account of their infertility, but by the inferiority of the hybrid offspring, both as being fewer in numbers, less able to continue their race, and less adapted to the conditions of existence than either of the pure forms. It is this inferiority of the hybrid offspring that is the essential point; and as the number of these hybrids will be permanently less where the infertility is greatest, therefore those portions of the two forms in which infertility is greatest will have the advantage, and will ultimately survive in the struggle for existence.
The differentiation of the two forms into distinct species, with the increase of infertility between them, would be greatly assisted by two other important factors in the problem. It has already been shown that, with each modification of form and habits, and especially with modifications of colour, there arises a disinclination of the two forms to pair together; and this would produce an amount of isolation which would greatly assist the specialisation of the forms in adaptation to their different conditions of life. Again, evidence has been adduced that change of conditions or of mode of life is a potent cause of disturbance of the reproductive system, and, consequently, of infertility. We may therefore assume that, as the two forms adopted more and more different modes of life, and perhaps acquired also decided peculiarities of form and coloration, the infertility between them would increase or become more general; and as we have seen that every such increase of infertility would give that portion of the species in which it arose an advantage over the remaining portions in which the two varieties were more fertile together, all this induced infertility would maintain itself, and still further increase the general infertility between the two forms of the species.
It follows, then, that specialisation to separate conditions of life, differentiation of external characters, disinclination to cross-unions, and the infertility of the hybrid produce of these unions, would all proceed pari passu [at an equal pace, side by side], and would ultimately lead to the production of two distinct forms having all the characteristics, physiological as well as structural, of true species.
In the case now discussed it has been supposed, that some amount of general infertility might arise in correlation with the different modes of life of two varieties or incipient species. A considerable body of facts already adduced renders it probable that this is the mode in which any widespread infertility would arise; and, if so, it has been shown that, by the influence of natural selection and the known laws which affect varieties, the infertility would be gradually increased. But, if we suppose the infertility to arise sporadically within the two forms, and to affect only a small proportion of the individuals in any area, it will be difficult, if not impossible, to show that such infertility would have any tendency to increase, or would produce any but a prejudicial effect. If, for example, five per cent of each form thus varied so as to be infertile with the other form, the result would be hardly perceptible, because the individuals which formed cross-unions and produced hybrids would constitute a very small portion of the whole species; and the hybrid offspring, being at a disadvantage in the struggle for existence and being themselves infertile, would soon die out, while the much more numerous fertile portion of the two forms would increase rapidly, and furnish a sufficient number of pure-bred offspring of each form to take the place of the somewhat inferior hybrids between them whenever the struggle for existence became severe. We must suppose that the normal fertile forms would transmit their fertility to their progeny, and the few infertile forms their infertility; but the latter would necessarily lose half their proper increase by the sterility of their hybrid offspring whenever they crossed with the other form, and when they bred with their own form the tendency to sterility would die out except in the very minute proportion of the five per cent (one-twentieth) that chance would lead to pair together. Under these circumstances the incipient sterility between the two forms would rapidly be eliminated, and could never rise much above the numbers which were produced by sporadic variation each year.
It was, probably, by a consideration of some such case as this that Mr. Darwin came to the conclusion that infertility arising between incipient species could not be increased by natural selection; and this is the more likely, as he was always disposed to minimise both the frequency and the amount even of structural variations.
We have yet to notice another mode of action of natural selection in favouring and perpetuating any infertility that may arise between two incipient species. If several distinct species are undergoing modification at the same time and in the same area, to adapt them to some new conditions that have arisen there, then any species in which the structural or colour differences that have arisen between it and its varieties or close allies were correlated with infertility of the crosses between them, would have an advantage over the corresponding varieties of other species in which there was no such physiological peculiarity. Thus, incipient species which were infertile together would have an advantage over other incipient species which were fertile, and, whenever the struggle for existence became severe, would prevail over them and take their place. Such infertility, being correlated with constitutional or structural differences, would probably, as already suggested, go on increasing as these differences increased; and thus, by the time the new species became fully differentiated from its parent form (or brother variety) the infertility might have become as well marked as we usually find it to be between distinct species.
This discussion has led us to some conclusions of the greatest importance as bearing on the difficult problem of the cause of the sterility of the hybrids between distinct species. Accepting, as highly probable, the fact of variations in fertility occurring in correlation with variations in habits, colour, or structure, we see, that so long as such variations occurred only sporadically, and affected but a small proportion of the individuals in any area, the infertility could not be increased by natural selection, but would tend to die out almost as fast as it was produced. If, however, it was so closely correlated with physical variations or diverse modes of life as to affect, even in a small degree, a considerable proportion of the individuals of the two forms in definite areas, it would be preserved by natural selection, and the portion of the varying species thus affected would increase at the expense of those portions which were more fertile when crossed. Each further variation towards infertility between the two forms would be again preserved, and thus the incipient infertility of the hybrid offspring might be increased till it became so great as almost to amount to sterility. Yet further, we have seen that if several competing species in the same area were being simultaneously modified, those between whose varieties infertility arose would have an advantage over those whose varieties remained fertile inter se [among themselves], and would ultimately supplant them.
The preceding argument, it will be seen, depends entirely upon the assumption that some amount of infertility characterises the distinct varieties which are in process of differentiation into species; and it may be objected that of such infertility there is no proof. This is admitted; but it is urged that facts have been adduced which render such infertility probable, at least in some cases, and this is all that is required. It is by no means necessary that all varieties should exhibit incipient infertility, but only, some varieties; for we know that, of the innumerable varieties that occur but few become developed into distinct species, and it may be that the absence of infertility, to obviate the effects of intercrossing, is one of the usual causes of their failure. All I have attempted to show is, that when incipient infertility does occur in correlation with other varietal differences, that infertility can be, and in fact must be, increased by natural selection; and this, it appears to me, is a decided step in advance in the solution of the problem.
Footnote to section: As this argument is a rather difficult one to follow, while its theoretical importance is very great, I add here the following briefer exposition of it, in a series of propositions; being, with a few verbal alterations, a copy of what I wrote on the subject about twenty years back. Some readers may find this easier to follow than the fuller discussion in the text: -
Can Sterility of Hybrids have been Produced by Natural Selection?
1. Let there be a species which has varied into two forms each adapted to certain existing conditions better than the parent form, which they soon supplant.
2. If these two forms, which are supposed to coexist in the same district, do not intercross, natural selection will accumulate all favourable variations till they become well suited to their conditions of life, and form two slightly differing species.
3. But if these two forms freely intercross with each other, and produce hybrids, which are also quite fertile inter se [among themselves], then the formation of the two distinct races or species will be retarded, or perhaps entirely prevented; for the offspring of the crossed unions will be more vigorous owing to the cross, although less adapted to their conditions of life than either of the pure breeds.
4. Now, let a partial sterility of the hybrids of some considerable proportion of these two forms arise; and, as this would probably be due to some special conditions of life, we may fairly suppose it to arise in some definite portion of the area occupied by the two forms.
5. The result will be that, in that area, the hybrids (although continually produced by first crosses almost as freely as before) will not themselves increase so rapidly as the two pure forms; and as the two pure forms are, by the terms of the problem, better suited to their several conditions of life than the hybrids, they will inevitably increase more rapidly, and will continually tend to supplant the hybrids altogether at every recurrent severe struggle for existence.
6. We may fairly suppose, also, that as soon as any sterility appears some disinclination to cross unions will appear, and this will further tend to the diminution of the production of hybrids.
7. In the other part of the area, however, where hybridism occurs with perfect freedom, hybrids of various degrees may increase till they equal or even exceed in number the pure species - that is, the incipient species will be liable to be swamped by intercrossing.
8. The first result, then, of a partial sterility of crosses appearing in one part of the area occupied by the two forms, will be - that the great majority of the individuals will there consist of the two pure forms only, while in the remaining part these will be in a minority, - which is the same as saying that the new physiological variety of the two forms will be better suited to the conditions of existence than the remaining portion which has not varied physiologically.
9. But when the struggle for existence becomes severe, that variety which is best adapted to the conditions of existence always supplants that which is imperfectly adapted; therefore, by natural selection the varieties which are sterile when crossed will become established as the only ones.
10. Now let variations in the amount of sterility and in the disinclination to crossed unions continue to occur - also in certain parts of the area: exactly the same result must recur, and the progeny of this new physiological variety will in time occupy the whole area.
11. There is yet another consideration that would facilitate the process. It seems probable that the sterility variations would, to some extent, concur with, and perhaps depend upon, the specific variations; so that, just in proportion as the two forms diverged and became better adapted to the conditions of existence, they would become more sterile when intercrossed. If this were the case, then natural selection would act with double strength; and those which were better adapted to survive both structurally and physiologically would certainly do so.]
Another form of infertility has been suggested by Professor G.J. Romanes as having aided in bringing about the characteristic infertility or sterility of hybrids. It is founded on the fact, already noticed, that certain individuals of some species possess what may be termed selective sterility - that is, while fertile with some individuals of the species they are sterile with others, and this altogether independently of any differences of form, colour, or structure. The phenomenon, in the only form in which it has been observed, is that of "infertility or absolute sterility between two individuals, each of which is perfectly fertile with all other individuals;" but Mr. Romanes thinks that "it would not be nearly so remarkable, or physiologically improbable, that such incompatibility should run through a whole race or strain."
[A footnote stated: Cases of this kind are referred to at p. 155. It must, however, be noted, that such sterility in first crosses appears to be equally rare between different species of the same genus as between individuals of the same species. Mules and other hybrids are freely produced between very distinct species, but are themselves infertile or quite sterile; and it is this infertility or sterility of the hybrids that is the characteristic - and was once thought to be the criterion - of species, not the sterility of their first crosses. Hence we should not expect to find any constant infertility in the first crosses between the distinct strains or varieties that formed the starting-point of new species, but only a slight amount of infertility in their mongrel offspring. It follows, that Mr. Romanes' theory of Physiological Selection - which assumes sterility or infertility between first crosses as the fundamental fact in the origin of species - does not accord with the general phenomena of hybridism in nature.]
Admitting that this may be so, though we have at present no evidence whatever in support of it, it remains to be considered whether such physiological varieties could maintain themselves, or whether, as in the cases of sporadic infertility already discussed, they would necessarily die out unless correlated with useful characters. Mr. Romanes thinks that they would persist, and urges that "whenever this one kind of variation occurs it cannot escape the preserving agency of physiological selection. Hence, even if it be granted that the variation which affects the reproductive system in this particular way is a variation of comparatively rare occurrence, still, as it must always be preserved whenever it does occur, its influence in the manufacture of specific types must be cumulative." The very positive statements which I have italicised would lead most readers to believe that the alleged fact had been demonstrated by a careful working out of the process in some definite supposed cases. This, however, has nowhere been done in Mr. Romanes' paper; and as it is the vital theoretical point on which any possible value of the new theory rests, and as it appears so opposed to the self-destructive effects of simple infertility, which we have already demonstrated when it occurs between the intermingled portion of two varieties, it must be carefully examined. In doing so, I will suppose that the required variation is not of "rare occurrence," but of considerable amount, and that it appears afresh each year to about the same extent, thus giving the theory every possible advantage.
et us then suppose that a given species consists of 100,000 individuals of each sex, with only the usual amount of fluctuating external variability. Let a physiological variation arise, so that 10 per cent of the whole number - 10,000 individuals of each sex - while remaining fertile inter se [among themselves] become quite sterile with the remaining 90,000. This peculiarity is not correlated with any external differences of form or colour, or with inherent peculiarities of likes or dislikes leading to any choice as to the pairing of the two sets of individuals. We have now to inquire, What would be the result?
Taking, first, the 10,000 pairs of the physiological or abnormal variety, we find that each male of these might pair with any one of the whole 100,000 of the opposite sex. If, therefore, there was nothing to limit their choice to particular individuals of either variety, the probabilities are that 9000 of them would pair with the opposite variety, and only 1000 with their own variety - that is, that 9000 would form sterile unions, and only one thousand would form fertile unions.
Taking, next, the 90,000 normal individuals of either sex, we find, that each male of these has also a choice of 100,000 to pair with. The probabilities are, therefore, that nine-tenths of them - that is, 81,000 - would pair with their normal fellows, while 9000 would pair with the opposite abnormal variety forming the above-mentioned sterile unions.
Now, as the number of individuals forming a species remains constant, generally speaking, from year to year, we shall have next year also 100,000 pairs, of which the two physiological varieties will be in the proportion of eighty-one to one, or 98,780 pairs of the normal variety to 1220 [The exact number is 1219.51, but the fractions were omitted for clearness in the main text] of the abnormal, that being the proportion of the fertile unions of each. In this year we shall find, by the same rule of probabilities, that only 15 males of the abnormal variety will pair with their like and be fertile, the remaining 1205 forming sterile unions with some of the normal variety. The following year the total 100,000 pairs will consist of 99,984 of the normal, and only 16 of the abnormal variety; and the probabilities, of course, are, that the whole of these latter will pair with some of the enormous preponderance of normal individuals, and, their unions being sterile, the physiological variety will become extinct in the third year.
If now in the second and each succeeding year a similar proportion as at first (10 per cent) of the physiological variety is produced afresh from the ranks of the normal variety, the same rate of diminution will go on, and it will be found that, on the most favourable estimate, the physiological variety can never exceed 12,000 to the 88,000 of the normal form of the species, as shown by the following table: -
1st Year. 10,000 of physiological variety to 90,000 of normal variety.
2nd Year. 1,220 + 10,000 again produced.
3rd Year. 16 + 1,220 + 10,000 do. = 11,236
4th Year. O + 16 + 1,220 + 10,000 do. = 11,236
5th Year. O + 16 + 1,220 + 10,000 = 11,236
and so on for any number of generations.
In the preceding discussion we have given the theory the advantage of the large proportion of 10 per cent of this very exceptional variety arising in its midst year by year, and we have seen that, even under these favourable conditions, it is unable to increase its numbers much above its starting-point, and that it remains wholly dependent on the continued renewal of the variety for its existence beyond a few years. It appears, then, that this form of inter-specific sterility cannot be increased by natural or any other known form of selection, but that it contains within itself its own principle of destruction. If it is proposed to get over the difficulty by postulating a larger percentage of the variety annually arising within the species, we shall not affect the law of decrease until we approach equality in the numbers of the two varieties. But with any such increase of the physiological variety the species itself would inevitably suffer by the large proportion of sterile unions in its midst, and would thus be at a great disadvantage in competition with other species which were fertile throughout. Thus, natural selection will always tend to weed out any species with too great a tendency to sterility among its own members, and will therefore prevent such sterility from becoming the general characteristic of varying species, which this theory demands should be the case.
On the whole, then, it appears clear that no form of infertility or sterility between the individuals of a species, can be increased by natural selection unless correlated with some useful variation, while all infertility not so correlated has a constant tendency to effect its own elimination. But the opposite property, fertility, is of vital importance to every species, and gives the offspring of the individuals which possess it, in consequence of their superior numbers, a greater chance of survival in the battle of life. It is, therefore, directly under the control of natural selection, which acts both by the self-preservation of fertile and the self-destruction of infertile stocks - except always where correlated as above, when they become useful, and therefore subject to be increased by natural selection.
SUMMARY AND CONCLUDING REMARKS ON HYBRIDITY.
The facts which are of the greatest importance to a comprehension of this very difficult subject are those which show the extreme susceptibility of the reproductive system both in plants and animals. We have seen how both these classes of organisms may be rendered infertile, by a change of conditions which does not affect their general health, by captivity, or by too close interbreeding. We have seen, also, that infertility is frequently correlated with a difference of colour, or with other characters; that it is not proportionate to divergence of structure; that it varies in reciprocal crosses between pairs of the same species; while in the cases of dimorphic and trimorphic plants the different crosses between the same pair of individuals may be fertile or sterile at the same time. It appears as if fertility depended on such a delicate adjustment of the male and female elements to each other, that, unless constantly kept up by the preservation of the most fertile individuals, sterility is always liable to arise. This preservation always occurs within the limits of each species, both because fertility is of the highest importance to the continuance of the race, and also because sterility (and to a less extent infertility) is self-destructive as well as injurious to the species.
So long therefore as a species remains undivided, and in occupation of a continuous area, its fertility is kept up by natural selection; but the moment it becomes separated, either by geographical or selective isolation, or by diversity of station or of habits, then, while each portion must be kept fertile inter se [among themselves], there is nothing to prevent infertility arising between the two separated portions. As the two portions will necessarily exist under somewhat different conditions of life, and will usually have acquired some diversity of form and colour - both which circumstances we know to be either the cause of infertility or to be correlated with it, - the fact of some degree of infertility usually appearing between closely allied but locally or physiologically segregated species is exactly what we should expect.
The reason why varieties do not usually exhibit a similar amount of infertility is not difficult to explain. The popular conclusions on this matter have been drawn chiefly from what occurs among domestic animals, and we have seen that the very first essential to their becoming domesticated was that they should continue fertile under changed conditions of life. During the slow process of the formation of new varieties by conscious or unconscious selection, fertility has always been an essential character, and has thus been invariably preserved or increased; while there is some evidence to show that domestication itself tends to increase fertility.
Among plants, wild species and varieties have been more frequently experimented on than among animals, and we accordingly find numerous cases in which distinct species of plants are perfectly fertile when crossed, their hybrid offspring being also fertile inter se [among themselves]. We also find some few examples of the converse fact - varieties of the same species which when crossed are infertile or even sterile.
The idea that either infertility or geographical isolation is absolutely essential to the formation of new species, in order to prevent the swamping effects of intercrossing, has been shown to be unsound, because the varieties or incipient species will, in most cases, be sufficiently isolated by having adopted different habits or by frequenting different stations; while selective association, which is known to be general among distinct varieties or breeds of the same species, will produce an effective isolation even when the two forms occupy the same area.
From the various considerations now adverted to, Mr. Darwin arrived at the conclusion that the sterility or infertility of species with each other, whether manifested in the difficulty of obtaining first crosses between them or in the sterility of the hybrids thus obtained, is not a constant or necessary result of specific difference, but is incidental on unknown peculiarities of the reproductive system. These peculiarities constantly tend to arise under changed conditions owing to the extreme susceptibility of that system, and they are usually correlated with variations of form or of colour. Hence, as fixed differences of form and colour, slowly gained by natural selection in adaptation to changed conditions, are what essentially characterise distinct species, some amount of infertility between species is the usual result.
Here the problem was left by Mr. Darwin; but we have shown that its solution may be carried a step further. If we accept the association of some degree of infertility, however slight, as a not unfrequent accompaniment of the external differences which always arise in a state of nature between varieties and incipient species, it has been shown that natural selection has power to increase that infertility just as it has power to increase other favourable variations. Such an increase of infertility will be beneficial, whenever new species arise in the same area with the parent form; and we thus see how, out of the fluctuating and very unequal amounts of infertility correlated with physical variations, there may have arisen that larger and more constant amount which appears usually to characterise well-marked species.
The great body of facts of which a condensed account has been given in the present chapter, although from an experimental point of view very insufficient, all point to the general conclusion we have now reached, and afford us a not unsatisfactory solution of the great problem of hybridism in relation to the origin of species by means of natural selection. Further experimental research is needed in order to complete the elucidation of the subject; but until these additional facts are forthcoming no new theory seems required for the explanation of the phenomena.
EXCERPTS FROM "THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION" BY CHARLES DARWIN
From Chapter 1.I "Domestic Dogs and Cats"
There is another and closely allied difficulty consequent on the doctrine of the descent of our domestic dogs from several wild species, namely, that they do not seem to be perfectly fertile with their supposed parents. But the experiment has not been quite fairly tried; the Hungarian dog, for instance, which in external appearance so closely resembles the European wolf, ought to be crossed with this wolf: and the pariah dogs of India with Indian wolves and jackals; and so in other cases. That the sterility is very slight between certain dogs and wolves and other Canidae is shown by savages taking the trouble to cross them. Buffon got four successive generations from the wolf and dog, and the mongrels were perfectly fertile together. (1/48. M. Broca has shown ('Journal de Physiologie' tome 2 page 353) that Buffon's experiments have been often misrepresented. Broca has collected (pages 390-395) many facts on the fertility of crossed dogs, wolves, and jackals.) But more lately M. Flourens states positively as the result of his numerous experiments that hybrids from the wolf and dog, crossed inter se [among themselves], become sterile at the third generation, and those from the jackal and dog at the fourth generation. (1/49. 'De la Longevite Humaine' par M. Flourens 1855 page 143. Mr. Blyth says ('Indian Sporting Review' volume 2 page 137) that he has seen in India several hybrids from the pariah-dog and jackal; and between one of these hybrids and a terrier. The experiments of Hunter on the jackal are well-known. See also Isid. Geoffroy St.-Hilaire, 'Hist. Nat. Gen.' tome 3 page 217, who speaks of the hybrid offspring of the jackal as perfectly fertile for three generations.) But these animals were closely confined; and many wild animals, as we shall see in a future chapter, are rendered by confinement in some degree or even utterly sterile. The Dingo, which breeds freely in Australia with our imported dogs, would not breed though repeatedly crossed in the Jardin des Plantes. (1/50. On authority of F. Cuvier quoted in Bronn's 'Geschichte der Natur' b. 2 s. 164.) Some hounds from Central Africa, brought home by Major Denham, never bred in the Town of London (1/51. W.C.L. Martin 'History of the Dog' 1845 page 203. Mr. Philip P. King, after ample opportunities of observation, informs me that the Dingo and European dogs often cross in Australia.); and a similar tendency to sterility might be transmitted to the hybrid offspring of a wild animal. Moreover, it appears that in M. Flourens' experiments the hybrids were closely bred in and in for three or four generations; and this circumstance would most certainly increase the tendency to sterility. Several years ago I saw confined in the Zoological Gardens of London a female hybrid from an English dog and jackal, which even in this the first generation was so sterile that, as I was assured by her keeper, she did not fully exhibit her proper periods; but this case was certainly exceptional, as numerous instances have occurred of fertile hybrids from these two animals. In almost all experiments on the crossing of animals there are so many causes of doubt, that it is extremely difficult to come to any positive conclusion. It would, however, appear, that those who believe that our dogs are descended from several species will have not only to admit that their offspring after a long course of domestication generally lose all tendency to sterility when crossed together; but that between certain breeds of dogs and some of their supposed aboriginal parents a certain degree of sterility has been retained or possibly even acquired.
We have already seen how often savages cross their dogs with wild native species; and Pennant gives a curious account (1/74. 'History of Quadrupeds' 1793 volume 1 page 238.) of the manner in which Fochabers, in Scotland, was stocked "with a multitude of curs of a most wolfish aspect" from a single hybrid-wolf brought into that district.
Several naturalists, as Pallas, Temminck, Blyth, believe that domestic cats are the descendants of several species commingled: it is certain that cats cross readily with various wild species, and it would appear that the character of the domestic breeds has, at least in some cases, been thus affected. Sir W. Jardine has no doubt that, "in the north of Scotland, there has been occasional crossing with our native species (F. sylvestris), and that the result of these crosses has been kept in our houses. I have seen," he adds, "many cats very closely resembling the wild cat, and one or two that could scarcely be distinguished from it." Mr. Blyth (1/89. Asiatic Soc. of Calcutta; Curator's Report, August 1856. The passage from Sir W. Jardine is quoted from this Report. Mr. Blyth, who has especially attended to the wild and domestic cats of India, has given in this Report a very interesting discussion on their origin.) remarks on this passage, "but such cats are never seen in the southern parts of England; still, as compared with any Indian tame cat, the affinity of the ordinary British cat to F. sylvestris is manifest; and due I suspect to frequent intermixture at a time when the tame cat was first introduced into Britain and continued rare, while the wild species was far more abundant than at present." In Hungary, Jeitteles (1/90. 'Fauna Hungariae Sup.' 1862 s. 12.) was assured on trustworthy authority that a wild male cat crossed with a female domestic cat, and that the hybrids long lived in a domesticated state. In Algiers the domestic cat has crossed with the wild cat (F. lybica) of that country. (1/91. Isid. Geoffroy Saint-Hilaire 'Hist. Nat. Gen.' tome 3 page 177.) In South Africa as Mr. E. Layard informs me, the domestic cat intermingles freely with the wild F. caffra; he has seen a pair of hybrids which were quite tame and particularly attached to the lady who brought them up; and Mr. Fry has found that these hybrids are fertile. In India the domestic cat, according to Mr. Blyth, has crossed with four Indian species. With respect to one of these species, F. chaus, an excellent observer, Sir W. Elliot, informs me that he once killed, near Madras, a wild brood, which were evidently hybrids from the domestic cat; these young animals had a thick lynx-like tail and the broad brown bar on the inside of the forearm characteristic of F. chaus. Sir W. Elliot adds that he has often observed this same mark on the forearms of domestic cats in India. Mr. Blyth states that domestic cats coloured nearly like F. chaus, but not resembling that species in shape, abound in Bengal; he adds, "such a colouration is utterly unknown in European cats, and the proper tabby markings (pale streaks on a black ground, peculiarly and symmetrically disposed), so common in English cats, are never seen in those of India." Dr. D. Short has assured Mr. Blyth (1/92. 'Proc. Zoolog. Soc.' 1863 page 184.) that, at Hansi, hybrids between the common cat and F. ornata (or torquata) occur, "and that many of the domestic cats of that part of India were undistinguishable from the wild F. ornata." Azara states, but only on the authority of the inhabitants, that in Paraguay the cat has crossed with two native species. From these several cases we see that in Europe, Asia, Africa, and America, the common cat, which lives a freer life than most other domesticated animals, has crossed with various wild species; and that in some instances the crossing has been sufficiently frequent to affect the character of the breed.
Whether domestic cats have descended from several distinct species, or have only been modified by occasional crosses, their fertility, as far as is known, is unimpaired. The large Angora or Persian cat is the most distinct in structure and habits of all the domestic breeds; and is believed by Pallas, but on no distinct evidence, to be descended from the F. manul of middle Asia; and I am assured by Mr. Blyth that the Angora cat breeds freely with Indian cats, which, as we have already seen, have apparently been much crossed with F. chaus. In England half-bred Angora cats are perfectly fertile with one another.
From Chapter 1.II "Horses and Asses"
I have seen, in the British Museum, a hybrid from the ass and zebra dappled on its hinder quarters.
From Chapter 1.III "Pigs, Cattle, Sheep, Goats"
Bos primigenius and longifrons have been ranked by nearly all palaeontologists as distinct species; and it would not be reasonable to take a different view simply because their domesticated descendants now intercross with the utmost freedom. All the European breeds have so often been crossed both intentionally and unintentionally, that, if any sterility had ensued from such unions, it would certainly have been detected. As zebus inhabit a distant and much hotter region, and as they differ in so many characters from our European cattle, I have taken pains to ascertain whether the two forms are fertile when crossed. The late Lord Powis imported some zebus and crossed them with common cattle in Shropshire; and I was assured by his steward that the cross-bred animals were perfectly fertile with both parent-stocks. Mr. Blyth informs me that in India hybrids, with various proportions of either blood, are quite fertile; and this can hardly fail to be known, for in some districts (3/50. Walther 'Das Rindvieh' 1817 s. 30.) the two species are allowed to breed freely together. Most of the cattle which were first introduced into Tasmania were humped, so that at one time thousands of crossed animals existed there; and Mr. B. O'Neile Wilson, M.A., writes to me from Tasmania that he has never heard of any sterility having been observed. He himself formerly possessed a herd of such crossed cattle, and all were perfectly fertile; so much so, that he cannot remember even a single cow failing to calve. These several facts afford an important confirmation of the Pallasian doctrine that the descendants of species which when first domesticated would if crossed have been in all probability in some degree sterile, become perfectly fertile after a long course of domestication. In a future chapter we shall see that this doctrine throws some light on the difficult subject of Hybridism.
From Chapter 1.IV "Domestic Rabbits"
But from what we hear of the marvellous success in France in rearing hybrids between the hare and rabbit (4/7. See Dr. P. Broca's interesting memoir on this subject in Brown-Sequard 'Journ. de. Phys.' volume 2 page 367.), it is possible, though not probable, from the great difficulty in making the first cross, that some of the larger races, which are coloured like the hare, may have been modified by crosses with this animal. Nevertheless, the chief differences in the skeletons of the several domestic breeds cannot, as we shall presently see, have been derived from a cross with the hare.
From Chapter 2.XVIII. "On The Advantages And Disadvantages Of Changed Conditions Of Life: Sterility From Various Causes"
A colour plate of the offspring of lion and tiger by Geoffrey St Hilaire (1772 - 1844)
Engraving of liger cubs born 1824 by G B Whittaker (engraving dated 1825)
Many species of Felidae have bred in various menageries, although imported from diverse climates and closely confined. Mr. Bartlett, the present superintendent of the Zoological Gardens (18/17. On the Breeding of the Larger Felidae 'Proc. Zoolog. Soc.' 1861 page 140.) remarks that the lion appears to breed more frequently and to bring forth more young at a birth than any other species of the family. He adds that the tiger has rarely bred; "but there are several well-authenticated instances of the female tiger breeding with the lion." Strange as the fact may appear, many animals under confinement unite with distinct species and produce hybrids quite as freely as, or even more freely than, with their own species.
From Chapter 2.XIII "Inheritance Continued - Reversion or Atavism [Throwbacks]"
Many years ago I saw in the Zoological Gardens a curious triple hybrid, from a bay mare, by a hybrid from a male ass and female zebra [this would have meant a fertile male hybrid]. This animal when old had hardly any stripes; but I was assured by the superintendent, that when young it had shoulder-stripes, and faint stripes on its flanks and legs. I mention this case more especially as an instance of the stripes being much plainer during youth than in old age.
As the zebra has such a conspicuously striped body and legs, it might have been expected that the hybrids from this animal and the common ass would have had their legs in some degree striped; but it appears from the figures given in Dr. Gray's 'Knowsley Gleanings' and still more plainly from that given by Geoffroy and F. Cuvier (13/34. 'Hist. Nat. des Mammiferes' 1820 tome 1), that the legs are much more conspicuously striped than the rest of the body; and this fact is intelligible only on the belief that the ass aids in giving, through the power of reversion, this character to its hybrid offspring.
The quagga is banded over the whole front part of its body like a zebra, but has no stripes on its legs, or mere traces of them. But in the famous hybrid bred by Lord Morton (13/35. 'Philosoph. Transact.' 1821 page 20.) from a chestnut, nearly purely-bred, Arabian mare, by a male quagga, the stripes were "more strongly defined and darker than those on the legs of "the quagga." The mare was subsequently put to a black Arabian horse, and bore two colts, both of which, as formerly stated, were plainly striped on the legs, and one of them likewise had stripes on the neck and body.
The Equus indicus (13/36. Sclater in 'Proc. Zoolog. Soc.' 1862 page 163: this species is the Ghor-Khur of N.W. India, and has often been called the Hemionus of Pallas. See also Mr. Blyth's excellent paper in 'Journal of Asiatic Soc. of Bengal' volume 28 1860 page 229.) is characterised by a spinal stripe, without shoulder or leg stripes; but traces of these latter stripes may occasionally be seen even in the adult (13/37. Another species of wild ass, the true E. hemionus or Kiang, which ordinarily has no shoulder-stripes, is said occasionally to have them; and these, as with the horse and ass, are sometimes double: see Mr. Blyth in the paper just quoted and in 'Indian Sporting Review' 1856 page 320: and Col. Hamilton Smith in 'Nat. Library, Horses' page 318; and 'Dict. Class. d'Hist. Nat.' tome 3 page 563.) and Colonel S. Poole, who has had ample opportunities for observation, informs me that in the foal, when first born, the head and legs are often striped, but the shoulder-stripe is not so distinct as in the domestic ass; all these stripes, excepting that along the spine, soon disappear. Now a hybrid, raised at Knowsley (13/38. Figured in the 'Gleanings from the Knowsley Menageries' by Dr. J.E. Gray.) from a female of this species by a male domestic ass, had all four legs transversely and conspicuously striped, had three short stripes on each shoulder and had even some zebra-like stripes on its face! Dr. Gray informs me that he has seen a second hybrid of the same parentage, similarly striped.
The parents of all our domesticated animals were of course aboriginally wild in disposition; and when a domesticated species is crossed with a distinct species, whether this is a domesticated or only a tamed animal, the hybrids are often wild to such a degree, that the fact is intelligible only on the principle that the cross has caused a partial return to a primitive disposition. Thus, the Earl of Powis formerly imported some thoroughly domesticated humped cattle from India, and crossed them with English breeds, which belong to a distinct species; and his agent remarked to me, without any question having been asked, how oddly wild the cross-bred animals were. The European wild boar and the Chinese domesticated pig are almost certainly specifically distinct: Sir F. Darwin crossed a sow of the latter breed with a wild Alpine boar which had become extremely tame, but the young, though having half-domesticated blood in their veins, were "extremely wild in confinement, and would not eat swill like common English pigs." Captain Hutton, in India, crossed a tame goat with a wild one from the Himalaya, and he remarked to me how surprisingly wild the offspring were.
These latter facts remind us of the statements, so frequently made by travellers in all parts of the world, on the degraded state and savage disposition of crossed races of man. That many excellent and kind-hearted mulattos have existed no one will dispute; and a more mild and gentle set of men could hardly be found than the inhabitants of the island of Chiloe, who consist of Indians commingled with Spaniards in various proportions. On the other hand, many years ago, long before I had thought of the present subject, I was struck with the fact that, in South America, men of complicated descent between Negroes, Indians, and Spaniards, seldom had, whatever the cause might be, a good expression. (13/47. 'Journal of Researches' 1845 page 71.) Livingstone - and a more unimpeachable authority cannot be quoted, - after speaking of a half-caste man on the Zambesi, described by the Portuguese as a rare monster of inhumanity, remarks, "It is unaccountable why half-castes, such as he, are so much more cruel than the Portuguese, but such is undoubtedly the case." An inhabitant remarked to Livingstone, "God made white men, and God made black men, but the Devil made halfcastes." (13/48. 'Expedition to the Zambesi' 1865 pages 25, 150.) When two races, both low in the scale, are crossed the progeny seems to be eminently bad. Thus the noble- hearted Humboldt, who felt no prejudice against the inferior races, speaks in strong terms of the bad and savage disposition of Zambos, or half-castes between Indians and Negroes; and this conclusion has been arrived at by various observers. (13/49. Dr. P. Broca on 'Hybridity in the Genus Homo' English translation 1864 page 39.) From these facts we may perhaps infer that the degraded state of so many half-castes is in part due to reversion to a primitive and savage condition, induced by the act of crossing, even if mainly due to the unfavourable moral conditions under which they are generally reared.
Another form of reversion is far commoner, indeed is almost universal with the offspring from a cross, namely, to the characters proper to either pure parent-form. As a general rule, crossed offspring in the first generation are nearly intermediate between their parents, but the grandchildren and succeeding generations continually revert, in a greater or lesser degree, to one or both of their progenitors. Several authors have maintained that hybrids and mongrels include all the characters of both parents, not fused together, but merely mingled in different proportions in different parts of the body; or, as Naudin (13/50. 'Nouvelles Archives du Museum' tome 1 page 151.) has expressed it, a hybrid is a living mosaic-work, in which the eye cannot distinguish the discordant elements, so completely are they intermingled. [...] Naudin further believes that the segregation of the two specific elements or essences is eminently liable to occur in the male and female reproductive matter; and he thus explains the almost universal tendency to reversion [to one or other parental type] in successive hybrid generations [...] But it would, as I suspect, be more correct to say that the elements of both parent-species exist in every hybrid in a double state, namely, blended together and completely separate. How this is possible, and what the term specific essence or element may be supposed to express, I shall attempt to show in the chapter on the hypothesis of pangenesis.
Naudin's view, as propounded by him, is not applicable to the reappearance of characters lost long ago by variation; and it is hardly applicable to races or species which, after having been crossed at some former period with a distinct form, and having since lost all traces of the cross, nevertheless occasionally yield an individual which reverts (as in the case of the great- great-grandchild of the pointer Sappho) to the crossing form. The most simple case of reversion, namely, of a hybrid or mongrel to its grandparents, is connected by an almost perfect series with the extreme case of a purely-bred race recovering characters which had been lost during many ages; and we are thus led to infer that all the cases must be related by some common bond.
From Chapter 2.XIV "Inheritance Continued. Fixedness Of Character, Prepotency, Sexual Limitation, Correspondence Of Age"
With animals, the jackal is prepotent over the dog, as is stated by Flourens, who made many crosses between these animals; and this was likewise the case with a hybrid which I once saw between a jackal and a terrier. I cannot doubt, from the observations of Colin and others, that the ass is prepotent over the horse; the prepotency in this instance running more strongly through the male than through the female ass; so that the mule resembles the ass more closely than does the hinny. (14/16. Flourens 'Longevite Humaine' page 144 on crossed jackals. With respect to the difference between the mule and the hinny I am aware that this has generally been attributed to the sire and dam transmitting their characters differently; but Colin, who has given in his 'Traite Phys. Comp.' tome 2 pages 537-539, the fullest description which I have met with of these reciprocal hybrids, is strongly of opinion that the ass preponderates in both crosses, but in an unequal degree. This is likewise the conclusion of Flourens, and of Bechstein in his 'Naturgeschichte Deutschlands' b. 1 s. 294. The tail of the hinny is much more like that of the horse than is the tail of the mule, and this is generally accounted for by the males of both species transmitting with greater power this part of their structure; but a compound hybrid which I saw in the Zoological Gardens, from a mare by a hybrid ass- zebra, closely resembled its mother in its tail.)
From Chapter 2.XV "On Crossing"
It is remarkable, as has been strongly insisted upon by Isidore Geoffroy St. Hilaire in regard to animals, that the transmission of characters without fusion occurs very rarely when species are crossed; I know of one exception alone, namely, with the hybrids naturally produced between the common and hooded crow (Corvus corone and cornix), which, however, are closely allied species, differing in nothing except colour. Nor have I met with any well- ascertained cases of transmission of this kind, even when one form is strongly prepotent over another, when two races are crossed which have been slowly formed by man's selection, and therefore resemble to a certain extent natural species. Such cases as puppies in the same litter closely resembling two distinct breeds, are probably due to superfoetation, - that is, to the influence of two fathers.
From Chapter 2.XVIII "On The Advantages And Disadvantages Of Changed Conditions Of Life: Sterility From Various Causes"
The Genetta has bred both here and in the Jardin des Plantes, and produced hybrids.
In the nine-year Report it is stated that the bears had been seen in the Zoological Gardens to couple freely, but previously to 1848 had most rarely conceived. In the Reports published since this date three species have produced young (hybrids in one case) [...]
The common hare when confined has, I believe, never bred in Europe; though, according to a recent statement, it has crossed with the rabbit. (18/21. Although the existence of the Leporides, as described by Dr. Broca ('Journal de Phys.' tome 2 page 370), has been positively denied, yet Dr. Pigeaux ('Annals and Mag. of Nat. Hist.' volume 20 1867 page 75) affirms that the hare and rabbit have produced hybrids.)
A Macacus, according to Flourens, bred in Paris; and more than one species of this genus has produced young in London, especially the Macacus rhesus, which everywhere shows a special capacity to breed under confinement. Hybrids have been produced both in Paris and London from this same genus.
Several members of the family of Lemurs have produced hybrids in the Zoological Gardens.
ANIMAL LIFE AND THE WORLD OF NATURE
Hybrids from the mating of lion and tiger are by no means common. In the wild state they are unheard of, although in parts of Asia these animals were, and still are, occasionally found in the same haunts; and even in captivity, where cubs from the coupling of the two species have been produced, these mixed offspring have been usually poor specimens, very difficult of rearing. It has remained for one of the most enterprising collectors and naturalists of our time, Mr Carl Hagenbeck, not only to breed, but to bring successfully to a healthy maturity, specimens of this rare alliance between those two great and formidable felidae, the lion and tiger. The illustrations will indicate sufficiently how fortunate Mr Hagenbeck has been in his efforts to produce these hybrids. The oldest and biggest of the animals shown is a hybrid born on the 11th May, 1897. This fine beast, now more than five years old, equals and even excels in his proportions a well-grown lion, measuring as he does from nose tip to tail 10 ft 2 inches in length, and standing only three inches less than 4 ft at the shoulder. A good big lion will weigh about 400 lbs - Mr Selous gives the length of a lion shot by himself as 9 ft 11 inches, its height as 3 ft 8 inches, and its weight as 410 lbs - and it is probably that in very exceptional cases lions may attain 450 and even 500 lbs. But the hybrid in question, weighing as it does no less than 467 lbs, is certainly the superior of most well-grown lions, whether wild-bred or born in a menagerie.
A male lion-tiger hybrid of a male lion and tigress.
A pair of young lion-tigers.
This animal shows faint striping and mottling, and, in its characteristics, exhibits strong traces of both its parents. It has a somewhat lion-like head, and the tail is more like that of a lion than of a tiger. On the other hand, it has little or no trace of mane. It is a huge and very powerful beast, but, like most of Mr Hagenbeck's feline pets, has been reduced to a state of comparative tameness, taking part in the shows given by the keeper, with other felidae, such as lions, tigers, leopards and pumas, not to mention dogs and various species of the genus Ursus.
The next hybrids are a pair born, like the bigger animal, of lion father and tigress mother. They are about seventeen month old, having been born on the 28th April, 1901. They show far more of the tiger striping than their elder relative, and are altogether much more tiger-like in appearance. At the age of twelve or thirteen months - when photographed - they were already as big as most full-grown lions and tigers, and they promise to develop into enormous specimens of the great cat family. They are, apparently, although reduced to a certain state of tameness, none too sweet tempered, judging from their photographs. The small photograph supplied by Mr Hagenbeck shows another pair of hybrid cubs - mere babies, born in April of the present year and photographed a few weeks later. They lie with their foster-mother, a fox terrier, whose services had to be called in for the purpose of rearing them. They too, show strong traces of striping; but as lion cubs in extreme infancy show a good deal of marking also, chiefly spots and blotches, it is possible that these stripes may to some extent disappear as they approach maturity.
A male lion-tiger of a male lion and tigress.
Two male lion-tiger, of a male lion and tigress.
These experiments of Mr Hagenbeck are extremely interesting, but, whether fortunately or otherwise, it is almost certain that the breeder of these strange crosses between lion and tiger will never succeed in perpetuating a race of feline mules. Hybrids are notoriously infertile, and although mares - the mules of horses and asses - have strong maternal instincts, and will even develop milk and suckle young foals, which they manage to decoy away from their true mothers, there is, I believe, no genuine recorded instance of such a mule bringing forth offspring of her own. Already, I understand, Mr Hagenbeck has mated the big lion-tiger hybrid with other pure-bred felines, but with no result.
Darwin long since pointed out in his "Origin of Species," that "hybrids raised from two species which are very difficult to cross, and which rarely produce any offspring, are generally very sterile." That assertion seems to be borne out strongly by the present case, and it seems altogether unlikely that any perpetuation of this new kind of fancy stock is to be looked for. Nor, indeed, is it desirable. In their own wild habitats, and after their own fashion, lions and tigers are necessary and very splendid creatures. A bastard strain from the crossing of these two species is not in the least likely to add to the beauty of the wildernesses of Africa or Asia or to the usefulness of two necessary forms in the scheme of nature. Nor in civilized countries, beyond the mere fact of producing a "sport," or curiosity, are these hybrids likely to be of interest or of use to anyone.
In this case, at all events, nature seems to have wisely set limits which even the ingenuity of man is not likely to be able to evade. The causes of the sterility of first crosses and their hybrid progeny were subjects that puzzled even that profound thinker and enquirer, Darwin. He arrived at the conclusion that such sterility had not been acquired by natural selection. "In the case of hybrids," he says, "it (sterility) apparently depends on their whole organisation having been disturbed by being compounded by two distinct forms; the sterility be closely allied to that which so frequently affects pure species, when exposed to new and unnatural conditions."
EXCERPT FROM "GREAT APES: A STUDY OF ANTHROPOID LIFE"
BY ROBERT M. YERKES & ADA WATTERSON YERKES (JUNE 1929)
“A somewhat embarrassing complication for the student of classification results from the existence of what seem to be forms intermediate between the chimpanzee the gorilla. The appearances naturally suggested to various observers hybridisation. Against the probability of the crossing of the genera is the prevalent belief that chimpanzees and gorillas do not intermingle, though, as I Geoffroy-Saint-Hilaire (1858-61, p 56) pointed out, they may inhabit neighbouring territory. To various authorities the widely differing morphological characters of social relations in the genera constitute negative argument. Nevertheless, the existence of forms which are with difficulty classifiable as either chimpanzee or gorilla is established and necessity of explanation is evident. The problem may best be exhibited by reference to specimens which have figured in the literature as classification puzzles.
There was brought to the Dresden Zoological Garden in 1874, from the Loango coast of Africa, an anthropoid ape which is known in the literature as “Mafuca”. The peculiarities of appearance (see fig 75, p 211) and behaviour of this ape aroused curiosity, and eventually various authorities were called to Dresden to examine her and pronounce judgment concerning classification. We shall list the opinions of several of these observers and give references to the more important literature on the controversy which lasted for several years.
Bolau (1876, 1877) pronounce Mafuca a chimpanzee; Meyer (1876, 1881, 1881a) agreed with Bolau, and expressed the opinion that she was neither a gorilla, nor a hybrid. By Hartmann (1876, 1877, 1880) Mafuca was classified as a young gorilla, and Nissle (1876) also believed her to be a gorilla. Brief account of this interesting controversy is given by Keith (1896, p 32).
A strong advocate of hybridisation as explanation of this classificatory puzzle is von Koppenfels, who as evidence of crossing offers personal observation of the association of gorillas with chimpanzees in their native habitat. He clearly indicates his conviction thus:
“I believe it is proved that there are crosses between the male Troglodytes gorilla and the female Troglodytes niger, but for reasons easily understood, there are none in the opposite direction. I have in my possession positive proof of this. This settles all the questions about the gorilla, chimpanzee, Kooloo Kamba, N’schigo, M’bouve, the Sokos, Baboos, etc (Von Koppenfels, 1881, p 448)”
By some authorities, Koppenfels’ evidence of hybridisation is considered unconvincing and his explanation of forms intermediate between chimpanzee and gorilla therefore unacceptable. For example, Garner, who unquestionably had excellent opportunity to gather information, thus opposes his position:
”The ape known as “Mafuka,” which was exhibited in Dresden in 1875, was also brought from the Loango coast, and it is possible that this is the ape to which the native name pongo really belonged. This specimen in many respects conforms to the description of the ntyii given, but the idea suggested by certain writers that “Mafuka” was a cross between the gorilla and chimpanezee is not, to my mind, a tenable supposition. It would be difficult to believe that two apes of different species in a wild state would cross, but to believe that two that belonged to different genera would do so is even more illogical. (Garner, 1896, p 255).
Pertinent also to this discussion of hybridisation is the case of the exhibition specimen Johanna, a well-grown female ape exhibited in America and England by the Barnum and Bailey Circus and subsequently lodged in the Zoological Garden at Lisbon. By Duckworth (1899) she is referred to as an unclassifiable ape which, although sometimes considered a gorilla, more likely is a form of chimpanzee, intermediate between gorilla and chimpanzee and similar to the “Kulu-Kamba” of Du Chaillu and the famous specimen Mafuca (p 157).
Peculiarly important is the comment of Keith:
“”Johanna” is of interest because she represents a variety of Chimpanzee which approaches the Gorilla in so many points that it is evident the characters which separate the two African anthropoids are not so well marked as many suppose. The difficulty of distinguishing the one from the other, as shown by a recent communication by Mr Duckworth to this society, is such that it has become necessary to sum up, from a much wider examination of material than has ever been at anyone’s disposal before, the structural and physiological differences which separate the Gorilla from the Chimpanzee, and at the same time to sum up the evidence of one or more species of Chimpanzee. Some five years ago on working minutely over all the anthropoid material in the collections of the Natural History Museum at South Kensington and the Museum of the Royal College of Surgeons, which contain the skulls of 31 Gorillas, 44 Chimpanzees, 73 Orangs, and 56 Gibbons, I was struck by the fact that nearly all the characters which had been used to differentiate species were points which varied in structure and form with age, sex, and the individual, but I have never had any difficulty in distinguishing between the skulls even of foetal Gorillas and Chimpanzees (Keith, 1899, p 266).
After critical examination of the entire literature we are of the opinion that hybridisation has not been definitely established. However, specimens of ape occur which so closely resemble the generic types that it is difficult to classify them. This we accept as evidence of the intimacy of genetic relation between the genera […] The problem of variability is considerably complicated at the present time by the possibility of hybridisation and the probability of dimorphism.
[…] The seemingly unique experience of von Koppenfels which is thus described:
“When beyond the Aschangolo Mountains in the neighbourhood of the Aschira land I shot a very strong male animal in a troop of chimpanzees which accidentally were eating cocoanuts peacefully with a family of gorillas. I privately suggested that I had shot the Kula Hamba discovered by Du Chaillu, and mentioned the possibility that since the two species of Troglodytes had been met peacefully together, there might be here a foundation of hybridisation. (1877, p 418)”
Even this is not necessarily a discrepant observation because the individual referred to by may very well have been a gorilla-like chimpanzee, as indeed the author admits in his mention of the “Kulu Hamba”.
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