In the late 1800s and early 1900s, there were two foremost breeds of zebrules or zebroids. Unlike earlier hybrids, Professor James Cossar Ewart (Scotland) and the Baron de Parana (Brazil) experimented with zebra stallions mated to a variety of pure-blooded mares of different breeds. The docility, strength and hardiness of the offspring led to high hopes that they would replace the stubborn ass-mules that were prone to various diseases in exotic climes. In particular, the zebrules were considered for use in expeditions and war, or for carriage work. So why didn't they catch on? It s likely that zebra stallions were harder to keep and manage than jack donkeys/asses, making large scale production of zebra-mules more difficult.

JAMES COSSAR EWART S ZEBRULES

The Sketch, 7th October, 1896, gives a background into the interest in zebra-horse : THE HYBRID ZEBRA EXPERIMENT. The interesting experiments of Professor Cossar Ewart, of Edinburgh, in breeding a hybrid from a Burchell Zebra and a mare have been already dealt with in these pages. Herewith is a picture of a result of the experiment-- to wit, the pretty little hybrid when it was twenty-eight days old. A comparison of it with its parents is instructive. It will be seen that the hybrid zebra is even more banded than the sire, the stripes on the body, singularly enough, being more numerous than in the Burchell Zebra, and resembling to a great extent those of the animal which was formerly known as the Common Zebra, now much less abundant than the Burchell. The scientific importance of these experiments was spoken of at length in our last article the practical bearing has to be considered. Burchell Zebras have been used to some considerable extent in South Africa, having been driven in Cape carts and in the ten-horse coaches in the Transvaal. In this country they have also been employed, and may occasionally be seen in the West-End. The Hon. Walter Rothschild has been driving a four-in-hand team consisting of three Burchells and a pony. But the value of the mules or hybrids bred from the zebra is probably of the highest importance. Among those who have studied the subject of mule-breeding, and whose evidence will be found at length in Tegetmeier and Sutherland's work on "Horses, Zebras, and Mules," the Burchell Zebra is regarded as the wild equine which will produce the most valuable mules when mated to a mare, inasmuch as it is the most equine and least asinine of all the zebras. The little hybrid whose photograph is given is most promising in its character. It is better in its quarters, shorter in its ears, and more foal like in its form than a mule bred from an ordinary donkey, and altogether promises to be a most useful animal.

The value of mules is not properly appreciated in this country. They work harder, are infinitely more enduring, cost less to keep, and are at least double as long-lived as the horse. In every military campaign thousands have to be bought by our Government. Three thousand mules were employed as transport animals in the subjugation of Chitral. There is in India a department organised for the breeding of mules for the Indian Army. These are bought at great expense, mainly in Italy. Now it is suggested that the Burchell Zebra should be employed instead of the domestic ass for the purpose of breeding these animals, and hence the very great importance that attaches to the foal represented, which has been bred by Professor Ewart. There is no doubt of the fertility of the two animals, but hitherto it has been the Burchell Zebra mare which has been mated with the horse, the result being the production not of what are ordinarily known as mules, but as hinnies or jennets. It is singular that in almost every other civilised country mules are appreciated much more highly than they are in England. In America they are employed in agricultural work to a much greater extent than horses, and in the last return made the value of the agricultural mules was placed higher per head than that of the horses. In Poitou the mule-breeding industry is the great support of the country, many thousand mares being employed for that purpose. The value of the Spanish mule is well known. By the English, mules are rarely used except in a campaign. Those bought by the Government and no longer required are occasionally brought to this country and sold. A very interesting anecdote is told by the authors of the book above mentioned respecting the excessive endurance and value of mules as draught animals. Some years ago a number sold by the Government were bought by the South London Tramways Company. Though fitted for the Army, they were too small for the purpose to which they were put, and three of them had to be placed in each tram to do the work of two larger horses. The writer went to see these animals after they had been working for twelve years, and found two teams in active service and good health. He then asked to see any horses that had been bought at the same time, and met with the reply, "Horses, sir No horses last with us more than three years." No more convincing argument of the endurance and utility of hybrid animals could be adduced. If the zebra-mules, as may be anticipated, turn out a success, and are marked like the object of the photo, they will form a very ornamental as well as useful addition to our equine animals.

THE PENYCUIK EXPERIMENTS.
Nature, No. 1551, Vol 60, July 20, 1899, p 272 – 274
["The Penycuik Experiments." By]. C. Ewart M.D., F.R.S.. Regius Professor of Natural History, University of Edinburgh. Pp. xciii + 177 (London: Adam and Charles Black, 1899.)

THE well-devised breeding experiments now in progress at Penycuik under the direction of Prof. J. Cossar Ewart, are, it need scarcely be said, of the highest interest both theoretical and practical. To the general biologist the subject of hybridisation affords a wide field for the investigation of laws of heredity, and especially of such subsidiary factors, whether real or only imaginary, as reversion, prepotency, saturation and telegony; while the question of the sterility of hybrids has important bearings on the general theory of evolution. But besides the purely scientific aspects of the problems which are now being attacked by Prof. Ewart, there is also their practical application, which appeals with much force to the interests of the breeder of stock. It is of course true that the whole history of animals and plants under domestication may be said to provide a body of experiments in these and similar subjects on a very large scale; and it is undoubtedly the case that many of the questions referred to have been already answered, at least provisionally. The experience of many generations of breeders has led to the emergence of certain practical rules, which are seldom if ever disregarded by those whose interests are concerned in the rearing of animals with a definite object. But it still remains doubtful how far the widely accepted doctrines of fanciers and other breeders rest upon any firm scientific basis; and it is certainly most desirable that precise experiments should be undertaken with the sole object of arriving at the truth in such matters as prepotency, telegony and the effects of inbreeding. It cannot but be to the advantage of breeders if empiric methods founded on vague conjecture and imperfect generalisation can be made to give place to a rational system derived from exact knowledge of facts. Prof. Ewart's design ought therefore to meet with a warm welcome in scientific and practical quarters alike.

The volume before us contains an account of such results of the author's experiments as are now sufficiently mature for publication. It is not so much a book as a re-issue in book form of three papers that have already seen the light, together with a general introduction which, to some extent, summarises and supplements the information subsequently given. Prof. Ewart remarks that "as the problems under consideration are not of a kind that can be settled off-hand, and as one inquiry has begotten others, some years must elapse before a complete and systematic account is possible." Meantime, he thinks, the publication of his results in their present form "will indicate the lines along which the inquiries are proceeding, and also the kind of answers likely eventually to be made to some of the questions." Regarded in this way as an instalment, the volume well fulfils its purpose.

The first paper is chiefly devoted to a detailed description of "Romulus," a hybrid colt whose sire "Matopo" is a Burchell's zebra of the Chapmani form, and whose dam "Mulatto" is an Island of Rum pony. "Mulatto" was afterwards crossed with a grey Arab stallion "Benazrek," and a description of her second foal, which unfortunately only lived for a little over five months, is given in the last paper under the head of" Telegony and Reversion." The foal in question was naturally a centre of great interest, inasmuch as from its parental history it afforded an opportunity for the operation of the alleged principle of telegony. The answer returned by nature to this particular interrogation was, as is so often the case, ambiguous. The colt was certainly striped, but the stripes were not like those of Mulatto's first mate; nor, on the other hand, did they entirely resemble those stripes often obscurely visible in ordinary foals. The evidence so far, though in no way conclusive, seemed to be capable of interpretation in accordance with the theory of telegony. However, in the general introduction, after giving the results of a fresh comparison with pure-bred foals, and adding an account of three additional experiments of the same kind, the author concludes that in no one of these cases is it possible to maintain that infection, saturation, or telegony has taken place. The results of further trials with rabbits, dogs and pigeons have also at present been uniformly negative.

In the second paper, particulars are given of a further batch of hybrids sired by the same zebra stallion "Matopo"; the dams being respectively a Shetland and an Iceland pony, an Irish mare and a cross-bred Clydesdale mare. All presented points of interest, and the extent to which they resembled their sire or respective dams varied much, but it was found that even those which in several characters most distinctly suggested the zebra sire differed entirely from him in markings. According to Prof. Ewart, the markings of his hybrids accord fairly with those of the Somali zebra, which he regards as being the most ancestral in its colour-pattern of all recent Equidae. If this is really the case - and it is difficult to find any weak spot in the author's cautious yet cogent line of argument - the hybrids in question supply one more good illustration of Darwin's principle that the crossing of distinct species frequently leads to reversion. It may here be remarked that precisely the same conclusion seems to follow from the elaborate experiments in the hybridisation of insects recently conducted at Zurich by Dr. M. Standfuss. The latter investigator, it is true, speaks only of the "prepotency of the phylogenetically older'' of the two parent species; but, while he refrains from actually using the term "reversion" with regard to his hybrids, he records the fact that some of them exhibit characters which must have belonged to an ancestor of one of the parent species, though absent from the parents themselves.

With regard to the zebra hybrids now under discussion, the most striking point of difference in marking between "Romulus" and the other cross-bred foals on the one hand, and their common sire "Matopo" on the other, is the multiplication of in the former, and the tendency to the production of a gridiron pattern over the rump. The last·named of these characters resembles the condition seen in the mountain zebra, an earlier form, according to Prof. Ewart, than the Burchell group, while the former point recalls the still more ancestral pattern of the Somali zebra. In the Shetland pony's hybrid, "Norette," the pattern over the hindquarters from the first resembled that of the Somali zebra; in the other hybrids, the markings of the same region, indeterminate at first, finally settled down into a form suggestive rather of the less remote stage marked by the mountain zebra. The difference in the general system of striping between "Matopo" and his offspring is well brought out in the figures here reproduced, by the courtesy of the publishers, from Prof. Ewart's work. A more special point, but one of great interest, is that exemplified in the accompanying figures of the brow-stripes in "Matopo," in "Romulus," and in a Somali zebra. The numerous rounded arches shown on the forehead of "Romulus" are very different from the four or five acutely pointed arches of "Matopo," and clearly bear a much greater resemblance to the corresponding pattern of the Somali zebra. It should, however, be mentioned that a system of brow-striping not unlike that of "Romulus" occurs in Crawshay's zebra, a member of the Burchell group.

On one point of special importance the experiments have so far given results that, however interesting scientifically, are from the practical side disappointing. Following a suggestion of Captain Lugard, that zebra mules might possibly turn out to be immune to the disease communicated by the tsetse fly, and might thus help in solving some of the difficulties of African transport, Prof. Ewart, with great liberality, inoculated three of his hybrids with some of the tsetse organism at that time under investigation by Messrs. Blandford and Durham. The result of this experiment is not given in the present volume, but in the recently published Proceedings of the International Congress of Zoology at Cambridge it is stated that the inoculated animals, though apparently somewhat more resistant than horses, all died in about eight weeks.

The above-mentioned are a few only of the points of interest brought out by the remarkable series of experiments dealt with in the present volume, but enough has probably been said to show the importance of the problems which Prof. Ewart has set himself to solve, and the prospects of advance in knowledge which these researches hold out. It only remains to say a word in commendation of the general get-up of the book, and of the character and accuracy of the illustrations, which in many cases are reproduced from actual photographs. The absence of an index or detailed table of contents is a drawback, but this, like the frequent repetition of the same facts, is perhaps inseparable from the method of publication adopted. A tabular list of the hybrids, giving their parentage and the more important features of their aspect, might be a useful addition, as the reader finds it a little difficult at present to piece together the various details, scattered through many parts of the work, under their proper headings. But any small defects of this kind will, no doubt, be completely remedied in the connected and systematic account of the fruit of his researches which Prof. Ewart leads us to hope for at some future time. Meanwhile, the course of his experiments will be watched with keen interest by all those who realise the importance, both scientific and practical, of a right conception of the laws of heredity. - F. A. D.

ZEBRA-HYBRIDS AT THE AGRICULTURAL SHOW. REMARKABLE RESULTS OF CROSS-BREEDING. - Pall Mall Gazette, 22nd June 1900

A very remarkable section of the Royal Agricultural Show, now being held at York, is that comprising the Zebra-hybrids, sent by Professor Cossar Ewart, who holds the Chair of Zoology at Edinburgh University. It is interesting as involving one of the obscurest and most far-reaching problems of animal evolution, heredity and variation, and the distinction between hybrids and species. The really practical value of Professor Ewart s experiments, as first expounded in his book recently published, and in a little brochure to be obtained on the Sound, is that for all kinds of transport purposes, in almost every British colony, and especially in Africa, the mule is acknowledged to be far superior to the horse, or even the ox, and it is believed by high authorities that the transport animal of the future will be a mule with strain of zebra blood in it, if not actually a direct cross between the horse and the mule. In his introduction to his descriptive catalogue Professor Ewart says: " There are many possible equine hybrids. There are (1) Hybrids between the horse and various species of asses: for example, the common mule (a cross between a jackass and a mare), and the hinny (a cross between a horse and a she-ass ; (2) hybrids between different species of asses; (3) hybrids between asses and zebras ; (4) hybrids between different species of zebras. And (5) hybrids between the horse and the different species of zebras. Hybrids between the ass and the female zebra have often been bred, and frequently the horse and the female mountain and Burchell zebras have interbred, but, until recently, the male Burchell zebra appears never to have been crossed with ordinary mares. A number of zebra-norse hybrids (zebrules) are now on view, and, in addition, the skin of the zebrinny (a horse and female zebra hybrid), and also the skin of zebrette (an ass and female zebra hybrid).

A few of the more noted exhibits may be mentioned, such as Romulus, foaled in 1896, a hybrid between "Matopo, a male Burchell zebra, and "Mulatto, a West Highland pony, bred by Lord Arthur Cecil. This animal, in his markings, decidedly differs from his zebra sire, but agrees with Grevy s zebra of Shoa and Somaliland, probably the most primitive of living zebras. "Sheila is a black foal, half-sister to "Romulus, born in 1899, 'and in her colour and make she takes after her dam "Mulatto, supposed to be a relic of the Spanish Armada. "Matopo is the sire of all the zebra-hybrids on view, a Burchell zebra (Chapman variety) from the borders of the Transvaal. This is an extremely interesting animal and remarkably docile. The stripes, by blending, form a grey, or khakki colour, which render such animals practically invisible at a distance of three or four yards by starlight, and of forty or fifty yards by our moonlight. This is a most important feature of these animals, and it is doubly interesting as illustrating the operation of that mysterious and unsolved law of nature, protective colouration.

Note: Sir George Ousley performed an almost dentical experiment. From Bell's Weekly Messenger, 11th December 1871, page 11, there is a report on the meeting of the Central Farmer's Club where a paper by Reginald Orton was mentioned, containing this excerpt: Many you have perhaps heard of Lord Morton's curious experiment with the Quagga; he put a Quagga stallion to a thorough-bred chestnut mare, and the produce was a Quagga mule with stripes, and many characteristics of the male. The season he put the mare to a black Arab horse, and to the astonishment of his lordship, she produced a foal bearing strong marks of the Quagga, and this effect was visible for three generations from a blood stallion. There is a similar case recorded in the transactions of the Royal Society, where a mare of Sir George Ousley s was put to a Zebra, and the produce was striped like the sire; and she was next year served by a blood horse, and the foal had the zebra stripes, and the same result again followed the next year. It therefore would appear that a female once impregnated retains certain traces of the male, which remains by her for an undefined period.

This is one of the reasons for Cossar Ewart's breeding experiments. A COMMUNICATION OF A SINGULAR FACT IN NATURAL HISTORY. BY THE RIGHT HONOURABLE THE EARL OF MORTON, F. R. S. IN A LETTER ADDRESSED TO THE PRESIDENT Philosophical Transactions of the Royal Society, 31 December 1821, Volume 111
My Dear Sir, I yesterday had an opportunity of observing a singular fact in Natural History, which you may perhaps deem not unworthy of being communicated to the Royal Society, Some years ago, I was desirous of trying the experiment of domesticating the Quagga, and endeavoured to procure some individuals of that species. I obtained a male; but being disappointed of a female, I tried to breed from the male quagga and a young chesnut mare of seven-eighths Arabian blood, and which had never been bred from: the result was the production of a female hybrid, now five years old, and bearing, both in her form and in her colour, very decided indications of her mixed origin. I subsequently parted with the seven-eighths Arabian mare to Sir Gore Ouseley, who has bred from her by a very fine black Arabian horse. I yesterday morning examined the produce, namely, a two-years old filly, and a year-old colt. They have the character of the Arabian breed as decidedly as can be expected, where fifteen-sixteenths of the blood are Arabian; and they are fine specimens of that breed; but both in their colour, and in the hair of their manes, they have a striking resemblance to the quagga. Their colour is bay, marked more or less like the quagga in a darker tint. Both are distinguished by the dark line along the ridge of the back, the dark stripes across the fore-hand, and the dark bars across the back part of the legs. The stripes across the fore-hand of the colt are confined to the withers, and to the part of the neck next to them; those on the filly cover nearly the whole of the neck and the back, as far as the flanks. The colour of her coat on the neck adjoining to the mane is pale, and approaching to dun, rendering the stripes there more conspicuous than those on the colt. The same pale tint appears in a less degree on the rump; and in this circumstance of the dun tint also she resembles the quagga.

PROFESSOR EWART'S PENYCUIK EXPERIMENTS - Popular Science Monthly Volume 57 June 1900

THE views and works of Darwin have influenced in an unexpected way the nature of the work carried on by biological investigators during the latter end of this fast-dying century. To a great extent, while generally holding the doctrines he held, they have forsaken his methods of inquiry.

If animals and plants have arrived at their present state by descent with modification from simpler forms, it ought to be possible by careful searching to trace the line of ancestry; and it is this fascinating but frequently futile pursuit which has dominated the minds of many of our ablest zoologists for the last thirty years. To such an extent has this pedigree hunting been carried that there is scarcely a group of invertebrates from which the vertebrates have not been theoretically derived; and to-day one of the ablest of our physiologists is using his great powers in the attempt to trace the origin of the backboned animals from a spiderlike creature, and is exercising his ingenuity in a plausible but unconvincing effort to equate the organs of a king-crab with those of a lamprey. This appeal to comparative anatomy and the consequent neglect of living animals and their habits are no doubt partly due to the influence of Huxley, Darwin's most brilliant follower and exponent. He had the engineer's way of looking at the world, and his influence was paramount in many schools. The trend which biology has taken since Darwin's time is also partly due to a fervent belief in the recapitulation theory, according to which an animal in developing from the egg passes through phases which resemble certain stages in the past history of the ancestors of the animal. For example, there is no doubt that both birds and mammals are descended from some fishlike animal that lived in the water and breathed by gills borne on slits in the gullet, and every bird and mammal passes through a stage in which these gill-slits are present, though their function is lost and they soon close up and disappear. In the hope, which has been but partially realized, that a knowledge of the stages through which an animal passes on its path from the ovum to the adult would throw light on the origin of the race, the attention of zo logists has been largely concentrated on details of embryology, and a mass of facts has already been accumulated which threatens to overwhelm the worker.

The two chief factors which play a part in the origin of species are heredity and variation, and until we know more about the laws which govern these factors we cannot hope to arrive at any satisfactory criteria by which we can estimate the importance of the data accumulated for us by comparative anatomists and embryologists. Signs are not wanting that this view is beginning to be appreciated. The publication of 'Materials for the Study of Variation by Mr. Bateson, a few years ago, shows that there exists a small but active school of workers in this field; and the recent congress on hybridization, held in London under the auspices of the Royal Horticultural Society, is evidence that in America, on the Continent and in Great Britain one of the most important sides of heredity is being minutely and extensively explored. Prof. Cossar Ewart's experiments, which we shall attempt to summarize, deal with heredity and cognate matters, and, although they are so far from complete that the results hitherto obtained can not be regarded as final, they mark an important stage in the history of the subject.

Five years ago Professor Ewart began to collect material for the study of the embryology of the horse, about which, owing to the costliness of the necessary investigations, very little is at present known. At the same time he determined to inquire into certain theories of heredity which have for centuries influenced the breeders of horses and cattle, and the belief in which has played a large part in the production of our more highly bred domestic animals. Foremost among these is the view widely held among breeders that a sire influences all the later progeny of a dam which has once produced a foal to him. This belief in the 'infection of the germ,' or 'throwing-back' to a previous sire, is probably an old one possibly as old as the similar faith in maternal impressions which led Jacob to place peeled wands before the cattle and sheep of his father-in-law Laban. The phenomenon has recently been endowed with a new name Telegony. Since the publication of Lord Morton's letter to Dr. W. H. Wollaston, President of the Royal Society, in 1820, it has attracted the attention not only of practical breeders, but of theoretical men of science. The supporters of telegony, when pressed by opponents, having almost always fallen back on Lord Morton's mare, it will be well to recall the chief incidents in the history of this classic animal.

It appears that early in this century Lord Morton was desirous of domesticating the quagga. He succeeded in obtaining a male, but, failing to procure a female, he put him to a young chestnut mare, of seven eighths Arab blood, which had never been bred from before. The result was the production of a female hybrid apparently intermediate in character between the sire and the dam. A short time afterward Lord Morton sold his mare to Sir Gore Ouseley, who bred from her by a fine black Arabian horse. The offspring of this union, which were examined by Lord Morton, were a two-year-old filly and a year-old colt.

He describes them as having "the character of the Arabian breed as decidedly as can be expected where fifteen sixteenths of the blood are Arabian, and they are fine specimens of that breed; but both in their color and in the hair of their manes they have a striking resemblance to the quagga." The description of the stripes visible on their coats is careful and circumstantial, but the evidence of the nature of the mane is less convincing: "Both their manes are black; that of the filly is short, stiff, and stands upright, and Sir Gore Ouseley's stud groom alleged that it never was otherwise. That of the colt is long, but so stiff as to arch upward and to hang clear of the sides of the neck, in which circumstance it resembles that of the hybrid."

This is the classical, we might almost say the test, case of telegony: the offspring resembled not so much the sire as an earlier mate of the dam. The facts related tended to confirm the popular view, and that view is widely spread. Arab breeders act on the belief, and it is so strongly implanted in the minds of certain English breeders that they make a point of mating their mares first with stallions having a good pedigree, so that their subsequent progeny may benefit by its influence, even though poorly bred sires are subsequently resorted to.

The evidence of Lord Morton's mare convinced Darwin of the existence of telegony; after a careful review of the case he says "there can be no doubt that the quagga affected the character of the offspring subsequently got by the black Arabian horse." Darwin, however, latterly came to the conclusion that telegony only occurred rarely, and some years before his death expressed the opinion that it was "a very occasional phenomenon." Agassiz believed in telegony. He was strongly of the opinion "that the act of fecundation is not an act which is limited in its effects, but that it is an act which affects the whole system, the sexual system especially; and in the sexual system the ovary to be impregnated hereafter is so modified by the first act that later impregnations do not efface that first impression."

Romanes also believed that telegony occasionally occurred. He paid a good deal of attention to the matter, commenced experiments in the hope of settling the question, and corresponded at length on this subject with professional and amateur breeders and fanciers. The result of his investigations led him to the conclusion "that the phenomenon is of much less frequent occurrence than is generally supposed. Indeed, it is so rare that I doubt whether it takes place in more than one or two per cent of cases." A recent controversy in the Contemporary Review shows us that Mr. Herbert Spencer is a firm upholder of telegony, and that he has a theory of his own as to the mode in which it is brought about. He suggests that some 'germ-plasm' passes from the embryo into the mother and becomes a permanent part of her body, and that this is diffused throughout her whole structure until it affects, among other organs, the reproductive glands. This view, which in some respects recalls the pangenesis of Darwin, is intermediate between the saturation and the infection hypothesis. Professor Ewart refers to it as indirect infection.

Weismann, to whom we owe the term telegony, came to consider the facts for and against its existence in connection with his well- known inquiry into the inheritance of acquired characters. If telegony be true, there is no need to look further for a clear case of the in- heritance of a character which has been acquired during the lifetime of the parent. The quagga-ness if one may be permitted to use such an expression of Lord Morton's mare was acquired when she was put to the quagga or shortly afterward, and was transmitted to her foals. A clearer case of a character acquired during lifetime and transmitted to offspring could not be imagined. Weismann does not absolutely deny the possibility of the existence of telegony, but he would like more evidence. In the Contemporary Review he writes, "I must say that to this day, and in spite of the additional cases brought forward by Spencer and Romanes, I do not consider that telegony has been proved." And further: "I should accept a case like that of Lord Morton's mare as satisfactory evidence if it were quite certainly beyond doubt. But that is by no means the case, as Settegast has abundantly proved." He would, in fact, refer the case to reversion, and quotes Settegast to the effect that every horse breeder is well aware that the cases are not rare when colts are born with stripes which recall the markings of a quagga or zebra. We shall return to this point later.

A considerable number of German breeders support the contention of Weismann that telegony is as yet unproved, and it may be pointed out that in Germany, on the whole, breeders have had a more scientific education than in England, and that in that country science is regarded with less aversion or contempt than is usually the case among so-called practical men in England. We may mention one more case of an ex- perienced breeder who was equally skeptical the late Sir Everett Millais, who was, as is well known, an authority of great weight in the matter of dog-breeding. He writes as follows, in a lecture entitled Two Problems of Reproduction: "I may further adduce the fact that in a breeding experience of nearly thirty years' standing, during which I have made all sorts of experiments with pure-blood dams and wild sires, and returned them afterward to pure sires of their own breeds, T have never seen a case of telegony, nor has my breeding stock suffered. I may further adduce the fact that I have made over fifty experiments for Professor Romanes to induce a case of telegony in a variety of animals dogs, ducks, hens, pigeons, etc. but I have hopelessly failed, as has every single experimenter who has tried to produce the phenomenon."

It is thus evident that there is a considerable body of opinion, both practical and. theoretical, for and against telegony; and that a reinvestigation of the subject is urgently needed. Such a reinvestigation has been begun by Professor Ewart at Penycuik. Since the clearest and most definite evidence of this throwing back to a previous sire is derived from the crossing of different species of the Equidae, it was desirable to repeat the experiment of Lord Morton. This is now unfortunately impossible, because the quagga is extinct. The zebra is, however, still with us, and the mating of a zebra stallion with every variety of horse, pony, and ass, and subsequently putting the dam to pure-bred-sires, has been the more important part of the numerous experiments carried on in the Midlothian village some ten miles southwest of Edinburgh.

Before considering in detail the result of the experiments it will be necessary to say a few words on the question of the various species of zebra; and since, like Weismann, Professor Ewart explains certain of the phenomena attributed-to telegony by reversion, it will be as well to inquire how far reversion is known among the Equidae, and what evidence we have that the ancestor of the horse was striped.

Matopo, the zebra stallion from which Professor Ewart had up to last midsummer bred eleven zebra-hybrids from mares of various breeds and sizes, belongs to the widely distributed group of Burchell's zebras. Many subspecies or varieties are included in this group, which, as regards the pattern of the stripes, passes in certain varieties found in Nyassaland into the second species, the mountain zebra, once com- mon in South Africa. The third species is the Grevy's zebra of Shoa and Somaliland; it is probably this species which attracted so much at- tention in the Koman amphitheaters during the third century of our era. A pair of Somali zebras has recently been presented to the queen by the Emperor Menelik, and is now lodged in the Zoological Gardens, Kegent's Park. The species measures about fifteen hands high, is pro- fusely striped, and stands well apart from the other two groups. It is important to note that, in Professor Ewart's opinion, it is the most primitive of all the existing striped horses.

There is no direct evidence that the ancestors of horses were striped. Certain observers think that some of the scratches on the lifelike etchings on bone, left us by our palaeolithic cave-dwelling ancestors, indicate such stripes, but little reliance can be placed on this. On the other hand, there is much indirect evidence. Every one who has an eye for a horse, and who has traveled in Norway, is sure to have noticed the stripings, often quite conspicuous, on the dun-colored Norwegian ponies. Colonel Poole assured Darwin that the Kattiawar horses had frequently "stripes on the cheeks and sides of the nose." Breeders are well aware that foals are often born with stripes, usually on the shoulders or legs, less frequently on the face. Such stripes, as a rule, disappear as the colt grows up, but can often be detected in later life for a short time after the coat has been shed; they are sometimes only visible in certain lights, and then produce somewhat the same impression as a watered silk. From the facts that more or less striped horses are found all over the Old World; that in Mexico and other parts of America the descendants of horses which were introduced by the Spaniards and which afterward ran wild are frequently dun-colored and show stripes; that foals are frequently striped; and that mules not uncommonly have leg and shoulder stripes, the inference is largely justified that the ancestors of all our horses were striped.

We now pass to the experiments made at Penycuik in crossing the zebra Matopo with various mares of different breeds:

1. Matopo was first mated with Mulatto, one of Lord Arthur Cecil's black West Highland ponies. The result was the hybrid Romulus (see p. 132), which on the whole, both in mental disposition and bodily form, takes more after his father than his mother. His striping is even more marked than that of his sire. He has a semi-erect mane which has been shed annually. The pattern of the markings, on both body and face, resembles the stripes on a Somali zebra which, as we have seen, is regarded by Professor Ewart as the most primitive type more than they resemble that of any of Burchell's zebras. The profuse striping is a point of difference between this hybrid and Lord Morton's. The quagga-hybrid was less striped than many dun-colored horses.

The mother Mulatto was next mated with a highly bred gray Arab horse, Benazrek. The offspring agrees in all respects with ordinary foals; it had, however, a certain number of indistinct stripes, which could only be detected in certain lights. The stripes were not nearly so clear as in a foal bred by Mr. Darwin from a cross-bred bay mare and a thoroughbred, and they disappeared entirely in about five months.

Recently Mulatto has produced a third foal to Loch Corrie, a sire belonging to the Isle of Rum group of West Highland ponies, and closely resembling his mate. This foal was about as much striped as its immediate predecessor. In both cases the pattern of the stripe differed not only from that of Matopo, the previous sire, but from that of the hybrid Romulus. These two foals seem to lend some support to telegony; but the evidence which might be drawn from the second of them is destroyed by the fact that the sire Loch Corrie has produced foals from two West Highland mares, one brown and one black, and each of these foals has as many and as well marked stripes as the foal of Mulatto.

2. Four attempts were made to cross the zebra with Shetland ponies; only one succeeded. The hybrid was a smaller edition of Romulus. The dam Nora had been bred from before, and had produced, by a black Shetland pony, a foal of a dun color which was markedly striped. After the birth of the hybrid she was put to a bay Welsh pony; the resulting foal had only the faintest indication of stripes, which soon disappeared. It is a remarkable fact that Nora's foals were more striped before she had been mated with the zebra than afterward.

3. Five Icelandic ponies were mated with Matopo, of whom one produced, in 1897, a dark-colored hybrid. The dam, Tundra, was a yellow and white skewbald which had previously produced a light bay foal to a stallion of its own breed. Her third foal (1898) was fathered by a bay Shetland pony, and in coloration closely resembled its dam. There was no hint of infection in this case. This year Professor Ewart has bred from this mare, by Matopo, a zebra-hybrid of a creamy-fawn color, and so primitive in its markings that he believes it to stand in much the same relation to horses, zebras, and asses as the blue-rock does to the various breeds of pigeons (see illustration).

.Two Irish mares, both bays, produced hybrids by Matopo, and subsequently bore pure-bred foals. One of the latter was by a thoroughbred horse, the other by a hackney pony. The foals were without stripes, and showed no kind of indication that their mother had ever been mated with a zebra.

5. Although Professor Ewart experimented with seven English thoroughbred mares and an Arab, he only succeeded in one case. The mare produced twin hybrids, one of which unfortunately died immediately after birth. This summer the same mare has produced a foal to a thoroughbred chestnut; "neither in make, color, nor action" does it in any way resemble a zebra or a zebra-hybrid.

6. A bay mare which had been in foal to Matopo for some months miscarried. Here if there is anything in the direct infection theory the unused germ-cells of the zebra had a better chance than usual of reaching the ova from which future offspring are to arise, yet neither of the two foals which this mare subsequently produced to a thorough- bred horse "in any way suggests a zebra."

The above is the record of the successful experiments which have been tried at Penycuik, with a view of throwing light on the existence of telegony in the Equidae.

Throughout his account of his experiments Professor Ewart is extremely cautious in claiming to prove anything, but we think he has justified his claim to have shown that telegony by no means always occurs, as many breeders believe. His experiments so far support the view of Continental mule breeders, that telegony, if it takes place, occurs very seldom. But the experiments are not complete, and it is much to be hoped that they may be continued. If it should subsequently appear that out of fifty pure-bred foals from dams which have been previously mated with the zebra no single instance of telegony be found, the doctrine may surely be neglected by breeders; and if in the experiments which are now being carried out with various other mammals and birds telegony does not occur, the doctrine may be relegated to the 'dumping-ground' of old superstitions. The present state of the matter may be summed up in the professor's own words: "The experiments, as far as they have gone, afford no evidence in support of the telegony hypothesis." Nothing has occurred which is not explicable on the theory of reversion.

Wild animals are frequently thought to be prepotent over tame ones, but of the eleven zebra-hybrids bred at Penycuik only two took markedly after their sire, the zebra Matopo. There are other experiments recounted which tell the other way, and at present this matter remains in a state of considerable uncertainty. This article must not close without a word or two more about the zebra-hybrids. It is mentioned above that only two out of the eleven which have already been born took, strongly after their father.

Those who have seen the young hybrids playing about in the fields at Penycuik must agree that they are the most charming and compactly built little animals possible. Of Romulus, the eldest of the herd, Professor Ewart says: "When a few days old [he] was the most attractive little creature I have ever seen. He seemed to combine all the grace and beauty of an antelope and a well-bred Arab foal. . . . What has struck me from the first has been his alertness and the expedition with which he escapes from suspicious or unfamiliar objects. When quite young, if caught napping in the paddock, the facility with which he, as it were, rolled on to his feet and darted off was wonderful."

The writer can fully confirm all the praise Professor Ewart lavishes on his pets; in truth, Romulus has been well described as a "bonnie colt with rare quality of bone . . . and with the dainty step and the dignity of the zebra." Remus, the offspring of the Irish mare, has been from the first more friendly than his half-brother; he objected less to the process of weaning, and, if he survives, promises to be the handsomest and fleetest of the existing hybrids.

On the whole, the hybrids are unusually hardy; only two have been lost one, a twin, which died almost as soon as it was born, and another which lived some three months and then succumbed. It is only fair to say that the dam of the latter, who was only three years old when the hybrid was born, had been much weakened by attacks of the strongylus worm, and that she was the victim of close inbreeding. Both the zebras and the hybrids which have been under observation at Penycuik show a remarkable capacity for recovering from wounds. Accidental injuries heal with great rapidity. On one occasion the surviving twin was discovered with a flap of skin some five inches long hanging down over the front of the left fetlock. The skin was stitched into its place again, during which operation the little hybrid fought desperately and cried piteously; but it soon recovered, the wound healed, and now scarcely a scar remains. There was no lameness and no swelling either at the fetlock or above the knee. About a year ago four hybrid colts and three ordinary foals were attacked by that scourge of the stable, the strongylus worm. One of the latter died and another was reduced almost to a skeleton; the hybrids, though obviously affected, suffered much less than the others and soon recovered. It is further noticeable that the hybrids suffer less from colds and other slight ail- ments than the mares and horses among which they live.

There is no doubt that it is a comparatively easy matter to breed these hybrids, and that they are not only extremely attractive animals to the eye, but hardy and vigorous, possessed of great staying powers, and promising to be capable of severe work. From what we have said, it is evident that the Penycuik experiments are of the highest interest, both practical and theoretical, and the public spirit and self-devotion shown by the Edinburgh professor in carrying them out can not be too widely recognized.

ZEBRA HYBRIDS: AND WHAT TELEGONY MEANS. The Sphere, 15th December 1900

Dr. Cossar Ewart, Professor of Natural History in the University of Edinburgh, has for a number of years been conducting at Penicuik an interesting series of biological experiments, the more important of which has been associated with the crossing of the zebra and horse. The result of these has been the production of a number of pretty hybrids which were seen by many people in Edinburgh at the Highland and Agricultural Society's show, and in York at the Royal Agricultural Society's exhibition, and were much admired. What is it that the professor seeks to attain by these experiments The objects are varied. In the first place he has been investigating what physiologists call the "infection" or telegony problem, which is an article of faith with many breeders of horses, dogs, and other beasts and birds. This doctrine may best be stated by the recital of a case which is always quoted in its support. Lord Morton in the early years of the century was the possessor of a quagga stallion, a species of zebra now believed to be extinct, which he mated with an Arab mare. The foal of course was a hybrid, but it was put on record afterwards by Lord Morton in a letter in 1820 to the Royal Society that this mare had been so "infected or "saturated by its first prepotent mate, the quagga, that afterwards to a black Arabian stallion it had four foals in successive years with striped coats. Is such a thing possible or is it not? Dr. Cossar Ewart wants to settle this point. He owns a beautiful specimen of a Burchell zebra stallion named "Matopo." Ten mares have had an opportunity of being "infected by "Matopo," but it may be said at once that the professor considers there is not the slightest evidence that any of them have been so. Let us examine the evidence as shown in the particular hybrids given on this page.

Romulus. He was got in 1896 by crossing "Matopo" with the mare Mulatto," one of Lord Arthur Cecil's Island of Rum ponies. At York the Prince of Wales took so great a fancy to "Romulus that the professor was constrained to let him go to Sandringham, where he is occasionally driven in harness by His Royal Highness - a proof, by the way, that a zebra hybrid is not the wild creature it is sometimes represented to be. "Romulus," it may be said, is richly striped, extremely well formed, and is unlike a mule in shape and action. Next year "Mulatto" had a foal to a grey Arab horse, and this foal presented a number of subtle markings that could hardly be called stripes, but which suggested stripes. They all, however, disappeared with the foal-coat. Two years afterwards "Mulatto had a foal to a West Highland pony, and again the youngster was faintly striped. These markings also disappeared when the first coat was shed. These cases seemed to give colour to the infection theory, but the doctrine was knocked on the head when it transpired on investigation that two mares of the same breed as "Mulatto," which had never seen a zebra, had similar foals to the same West Highland pony. Other mares have been mated in the same way, one year with the zebra and another year with a horse, and the foals of the latter have never shown any markings whatever that would support the infection theory.

Norette. Was born June 8, 1897. Its sire was "Matopo," its dam "Nora," a black Shetland pony with so little character that "Norette," except in colour, is extremely like a zebra. As showing how careful people should be in dogmatising on this subject it may be mentioned that in 1895 "Nora," two years before she saw "Matopo," had to "Wallace," a Shetland pony stallion, a foal which was richly striped, while the year after she had her hybrid (1898) she had a cross-bred foal, "Skua," which in no way suggested a zebra.

Remus. This zebra hybrid was born in 1897, by "Matopo out of "Biddy," an Irish roadster mare, while the other animal on the left is "Kathleen," the daughter of "Biddy in the following year (1898) by "Tupgill," a thoroughbred chestnut horse. In no single point, as will be observed by a study of the picture, does "Kathleen suggest a zebra.

Bergus.-- The picture of "Valda," a chestnut polo pony, with her zebra hybrid foal, "Bergus," the sire of which was "Matopo," is rather interesting as Valda bore twin hybrids, one of which died, while the other, "Bergus," still survives.

Hector. By way of contrast a picture is given of "Hector," the foal of "Valda," the year after the twin hybrids were born. His sire was "Lockstitch," a thoroughbred, and as will be seen he has nothing of the zebra about him either.

As already stated Professor Cossar Ewart's decided opinion from his experiments is that he has failed to obtain any evidence that either mares or other female animals can be infected in the way suggested by Lord Morton. It should be mentioned, as great stress is laid on the point by some breeders, that as regards several of the mares "Matopo was their first mate; in other cases he was not, but the results were just the same. Further, it may be stated that Professor Cossar Ewart is in correspondence with Baron de Parana of Brazil, who has been carrying out similar experiments, and with precisely the same results. The Baron has had also under observation the mares at a great establishment where some 3,000 mules are bred every year. After taking three mules off a mare the animal is then used for breeding horses. In no instance has a case been observed in these horse foals of any suggestion of a throwing back to the first donkey mate of the dam.

The professor has also been endeavouring to find out whether the unborn young can be influenced by "mental impressions," another common belief; but in that respect the results of his experiments have been of a wholly negative character. His inquiry has also included the investigation of the problem of the relative influence on the offspring of male and female parents. It is an old theory that from the male parent is mainly derived external structure, configuration, and outward characteristics, as also the locomotive system or development, and from the female parent the internal structures, the vital organs, and, more than from the male, the constitution, temper, and habits. This, however, is a theory that cannot be maintained. Experiments at Penicuik have gone to show conclusively that it is the prepotent animal which always prevails.

The experiments have also a practical side, doubly interesting at the present moment. Professor Cossar Ewart believes that a zebra hybrid will be found to be the best substitute in South Africa for the horse, which in some districts of that continent cannot live on account of sickness and the tsetse fly. The professor hopes that when the country is settled the Government will set up a breeding establishment at Vreda in the Orange River Colony, where he thinks it would do well.

NEW ANIMAL. - Lincolnshire Chronicle , 30th June 1903 Professor Cossar has been experimenting for some time with zebras to see if he could obtain an animal superior to the ordinary male. During the last eight or nine years he has bred from his Burchell zebra, Matopo, and different classes of mares chiefly Shetland, Iceland and Clydesdale a number of hybrids, which have now reached marketable value. Within the last twelve months Hagenbeck, of Hamburg, has bought eight the zebrules, as the new animals are called, and in February last the Indian Government bought two, which have been sent to Qaetta to be tried for mountain battery work. Baron de Parana, of Brazil, has also been experimenting with zebras, and obtained his first zebra horse hybrid, and it his belief that the zebrule will be the mule of the twentieth century. For strength, intelligence, and alertness the hybrid is greatly indebted to its zebra sire, and it is more easily broken in than the ordinary mule. This addition to our animals for military use comes opportunely, the tremendous drafts we had to make on the resources of the mule-raising countries to supply our need in the late war, together with the heavy losses in South Africa, have seriously depleted the stock of this useful animal.

A letter written by Carl Hagenbeck on 26 October 1903 stated that Baron von Falz-Fein from Russia and the Baron de Parana both had eight to ten zebra hybrids, and agreed that if their usefulness and strength were more widely known, they would only breed zebroids rather than asses and horses. Hagenbeck asked for the price of Ewart's two zebra hybrids and his Shetland pony and offered his Korea stallion to him as a present, as he wished to be rid of it.

SOME NEW HYBRIDS. - HAROLD J. SHEPSTONE.

SCIENTIFIC AMERICAN, November 7, 1903.

The recent purchase by the Indian government of two zebra-horse hybrids from Prof. J. Cossar Ewart, of Edinburgh University, Scotland, has called attention to the work of this gentleman, and, incidentally, to the experiments in cross-breeding of Mr. Carl Hagenbeck, the animal dealer at Hamburg, who has for some seven years past given serious consideration to this subject. During this comparatively short period Mr. Hagenbeck has produced an entirely new variety of sheep, deer, and pheasant by crossing the stronger, wilder species with tame, domestic specimens. It should be stated at once, perhaps, that the one aim in conducting these experiments has been to obtain a stronger and better animal that is to say, better blood. That this has been accomplished in the case of the zebra-horse cross there is not the slightest doubt. The new animal, which has been named the zebrule, has been produced with the object of taking up the work of the ordinary military mule. It is a little larger than the average mule, and when once broken to harness is much more tractable and far more intelligent. Furthermore, the new animal does not possess that stubborn will and dangerous tricks often found in the common mule. It is also hardy, a good trotter, very sure of its footing, and capable of adapting itself to great changes of climate and temperature.

Prof. Ewart seriously commenced his experiments nine years ago. He obtained a fine, healthy zebra stallion, "Matopo," now the sire of many zebrules. The first hybrid was born in August, 1896, at Penicuik, in Midlothian, Scotland, the dam being a pony, selected from a first-class breed, the sire, of course, being "Matopo." The foal proved strong and hardy, very easily broken to saddle and harness, while its intelligence surprised those in charge of it. The animal was produced merely for scientific purposes, but it was such a success that Prof. Ewart came to the conclusion that the new hybrid would be an ideal animal for ordnance and commissariat work. The experiments were therefore continued, and a number of zebrules bred from thoroughbreds, half-Arabs and Clydesdales. Several of these were purchased by Carl Hagenbeck , of Hamburg. Indeed, he had already bought the two which have been dispatched to India, but willingly gave them up to the Indian government when they decided to take them, as he is convinced from his experience and handling of the zebrule that it will be the mule of the twentieth century. He is endeavoring to get the German government to take some of these animals for use in their army. He has several in his park at Stellingen, near Hamburg, and when in that city recently the writer went for a drive behind two of these interesting creatures. Full-grown they stand about 14 hands high, and have a girth measurement of about 63 inches. They are therefore somewhat larger than a mule. At first sight they strike one as strange, the zebra striping in some instances being very distinct, though, of course, by no means so conspicuous as in the wild zebra. They answer to the reins instantly, are not at all nervous of electric cars or traffic, and fast trotters.

Prof. Ewart, in speaking of the hybrid in his book, "A Guide to Zebra Hybrids," says: "Zebrules are usually better able to take care of themselves than pure-bred animals, are more alert, more active, and altogether more vigorous and intelligent. From the first, zebra hybrids are more friendly, more curious and confiding than ordinary foals. With time and care most of them can be trained to any kind of work. It is almost impossible and far from safe to break in a young mule by itself, but quite possible to break in by itself a zebra hybrid." It was hoped that the British War Office would take the new animal, but it stolidly refused to touch them. It is as well, perhaps, that the animals are to be tried first in India, where it is well known the natives manage their mules with more tact and patience than the British soldier. The animals have already arrived at their destination, Quetta, and are now undergoing a series of tests as to their suitability for mountain battery work. Although no official report has been received, it has been stated that the animals fulfill all the requirements set down by the Indian government. Indeed, so well have they behaved that a number of drives have been organized in the East Africa Protectorate for the capture of young zebra stallions for exportation to India, Jamaica, and elsewhere for breeding purposes.

A hybrid that has attracted the attention of zoologists is the lion and tiger cross, a number of which may now be seen at the Hagenbeck establishment in Hamburg. The oldest is four years of age, and is a fine animal, called "Prince." He was recently seen performing with several other animals in one of the Hagenbeck trained groups in New York. When only three years of age he weighed five hundredweight and measured 10 feet from the tip of his tail to the tip of his nose, and stood four feet high to the top of the shoulder. The peculiarity of this beast is that he has a tiger's body and a lion's head, the stripes, of course, not being so distinct as in the common tiger. Prince's father was a Senegal lion and his mother a Bengal tiger. This one animal is valued by its owner at 10,000.

"The first successful experiment I made in the crossing of animals," said Mr. Hagenbeck, in speaking of this side of his interesting work, "was about seven years ago, when I crossed a leopard and a puma. The only living animal of the litter, which, fortunately, was a very healthy one, I sold some time ago to the Berlin Zoological Garden, where it is still in the collection. I am now busy endeavoring to obtain a new variety of sheep by crossing the giant sheep of Central Asia with our common domestic animal."

Sportsmen will be particularly interested in the new pheasant which the Hamburg establishment has produced. It has been obtained by crossing pheasants from Central Asia with the European variety. The result is a larger bird, much stronger on the wing, and more prettily marked. The new pheasant very much resembles the English or common American pheasant in appearance. It is somewhat larger, however, and sportsmen have already discovered that it is very quick on the wing, and not so easy to bring to the ground. Mr. Hagenbeck has succeeded in securing several broods of these pheasants. The first of these were obtained in April of last year. They were quickly snapped up by the Duke of Bedford and Lord Rothschild, who own extensive shooting grounds in England, these two gentlemen taking the larger portion of them. Several broods were obtained again at the beginning of this year, and the birds are all doing well. For years the European pheasant has been gradually declining, but with the introduction of the Asiatic variety if has received a new lease of life. In the same way several new varieties of deer have been secured by crossing Persian fallow deer with ordinary European deer.

NEW DRAFT ANIMAL DOOMS ARMY MULE. CROSS BETWEEN HORSE AND ZEBRA IS SUPERIOR. DOES NOT CONTRACT DISEASE. More Lively Than the Animal That Provokes Cussing and Cannot Be Less Intelligent. (Various, August 1903)

The days of the mule are numbered. Within the next few centuries his melodious voice will have been stilled forever. This is the prophecy of United States Consul General Richard Guenther, at Frankfort, Germany, who sends a report to the state department on the chances of the zebrula, a cross between the horse and zebra, superceding the mule. He says of the qualifications of the zebrula:

"German papers contend that it has been demonstrated that the mule, the cross between the horse and donkey, is inferior to the cross between horse and zebra. Formerly the opinion prevailed that the zebra was almost extinct. The opening up of Africa, particularly the eastern part, reveals these fine animals in large numbers. Compared with horses and cattle, they possess peculiar advantages, as they are immune against the very dangerous horse disease of Africa, and also against the deadly "tztze. The question was therefore raised whether the zebra could not take the place of the mule, commonly used in the tropics. The greatest credit with reference to the solution of this problem is due to Professor Cossar Ewart, who has been trying since 1895 to produce crosses between horses and zebras, with a view to developing an animal superior in every respect to the mule.

"Professor Ewart produced crosses from mares of different breeds and zebra stallions of the burchell kind. The offspring is called zebrula, and on account of its form and general bodily condition, especially the hardness of the hoofs, is specially adapted for all transport work heretofore performed by mules. The zebrula is much livelier than the mule and at least as intelligent. The zebra stripes are often well preserved, while the undertone of the skin is generally that of the mother. A full grown zebrula is fourteen hands high and the girdle circumference about 160 centimeters (sixty-three inches). The experiments so far have been so successful that it is predicted that the zebrula during the present century will completely supercede the mule.

Other newspapers of the time wrote "The days of the mule are numbered, comes the word from Germany, where a new animal, the Zebrula, has been evolved to take the place of the gentle-eyed, melodious-voiced and hard- hoofed little animal. The new animat1 is the result of a cross between the horse and a zebra. The nomenclature of horseology would indicate that a better name for the new quadruped would-be the zehorska."

A NEW ANIMAL.- ROMULUS THE ZEBRULE. Southland Times , Issue 19173, 10 October 1903, Page 4: Professor Cossar Ewart has been experimenting for some time with zebras to see if he could obtain an animal superior to the ordinary mule. During the last eight or nine years he has bred from his Burchell zebra, Matopo, and different classes of mares chiefly Shetland, Iceland and Clydesdale a number of hybrids, which have now reached a marketable value. Within the last twelve month; Hagenbeck, of Hamburg, has bought eight of the zebrules, , as the new animals are called, and in February last the Indian Government bought two, which have been sent to Quetta to be tried for mountain battery work. The illustrations show Matopo's eldest son, Romulus, when no was twenty-eight days old, and his head when full grown. His dam was Mulatto, one of Lord Arthur Cecil's Island of limn ponies. From the first Romulus was strong and hardy, and was easily broken to harness. He moves more like a zebra than a horse, and when young was somewhat restless when separated from his companions. Born on the 12th of August, 1890, he was presented to King Edward and carried off to Sandringham from the Royal Agricultural Society's Show at York in 1900. As a foal his color varied from orange to reddish brown, but since he shed his first coat he has become more of a dun. This is the first cross between a male zebra and an ordinary mare that has ever been bred ; but a few months later Baron de Parana, of Brazil, who has also been experimenting with zebras, obtained his first zebra -horse hybrid, and it is his belief that the zebrule will be the mule of the twentieth century, for strength, intelligence and alertness the hybrid is greatly indebted to its zebra sire, and it is more easily broken in than the ordinary mule.

ON HAIR IN THE EQUIDAE. By F. H. A. Marshall, B.A., F.R.S.E. Proceedings of Royal Society of Edinburgh (Read June 17, 1901.)

[Most of the paper looked at hair structure in horses and zebras as a way of distinguishing the species.]

ZEBRA-HORSE HYBRIDS.
[Vide Ewart, The Penycuik Experiments, London, 1899; and Guide to Zebra-Hybrids, Edinburgh, 1900.]
The hairs of several of Professor Ewart's zebra-horse hybrids have been examined and sections cut. Seeing that the dams of these animals belong to different breeds, it might at first be expected that we should find quite as much diversity in the character of the hybrid hairs as in those of the dams. Such, however, is not the case, for the hairs of the hybrids are for the most part constant in shape and in the relative development of the medulla and cortex. The measurements, taken as before, of the shoulder hairs of the hybrid "Norette," whose dam was a Shetland pony, are not widely different from those of the sire, the Burchell's zebra:: -

(1) Breadth of cortex on one side of medulla in three places
Breadth of hair in three places.0.0144 mm. 0.081 mm.
(2) Breadth of cortex on one side of medulla in three places
Breadth of hair in three places.0.0162 mm. 0.126 mm.
(3) Breadth of cortex on one side of medulla in three places
Breadth of hair in three places. 0.0198 mm. 0.054 mm.

They point to the conclusion that in the transmission of the character of the hair the Burchell's zebra is prepotent over the horse. In some cases, however, the hybrid hairs do not resemble those of the sire any more than those of the dam, but this is not because they depart from the hybrid type, but because the hairs of the dam happen to be not dissimilar to zebra hairs. It has been mentioned that horse hairs, owing to the large number of breeds of horses, are very variable, and so it is not to be wondered at that in some cases sections through horse hairs should resemble sections through zebra hairs. This is the case with sections taken through the hairs of the mane of Professor Ewart's Clydesdale mare, "Lady Douglas," the mane hairs of whose hybrid offspring "Brenda" are if anything more like those of the dam than those of the sire, " Matopo." A more typical case is that of the hybrid "Sir John" (Plate IV.). Here the dam was a skewbald Iceland pony, " Tundra," and the sire the Burchell's zebra. Sections through the mane hairs of "Tundra" are identical in appearance with sections through hairs of “Circus Girl," which are figured. Sections through the hairs of the hybrid offspring "Sir John," on the other hand, are in no way suggestive of those from the dam, but closely resemble those of the hybrids "Black Agnes" and "Brenda," one of which is figured (fig. 10). Professor Ewart has given reasons for the conclusion that of the existing species of zebras the Somali zebra approaches nearest to the ancestral type. He has also shown that the markings of the hybrids resemble the markings of the Somali zebra much more closely than those of the Burchell's zebra, and this resemblance he has ascribed to reversion. Now it cannot be said that the shoulder hairs of the hybrids, either in their shape, length, which is rather variable, or in the arrangement of the pigment, are at all suggestive of the same hairs in the Somali zebra. When, however, we compare the hairs of the mane the case is quite different. A section through a hair of the mane of a hybrid, such as the one figured, which is through such a hair in "Brenda," which in the mane hair characters is quite typical of the hybrids, shows a fairly even distribution of pigment and a circular line of demarcation between cortex and medulla, which are also what we find in a mane hair section from the Somali zebra. There is very little of that tendency of the pigment to become more thickly distributed towards the interior of the cortex, such as I have found in all sections through mane hairs of the Burchell's zebra. This is a curious result, and may, perhaps, like the peculiarities of the striping, be ascribed to reversion to the more ancestral type.

The Telegony Hypothesis.
Nathusius suggested that if the telegony hypothesis, or the hypothesis that subsequent offspring are infected by a previous sire be correct, we might expect to find evidence of it in the character of the hairs of the subsequent offspring. We have such a subsequent offspring in Professor Ewart's "Circus Girl." In 1897 the dam "Tundra" gave birth to a hybrid, "Hecla." In 1898 the subsequent foal "Circus Girl" was born, the sire being a bay Shetland pony. Just as "Circus Girl," both in make and colour, closely resembles her mother, so the hairs of the two animals are almost identical in character, and sections through the hairs of the manes are quite indistinguishable. There is nothing whatever suggestive of the Burchell's zebra " Matopo," which was the previous sire. The same remark is equally applicable, so far as I have seen, to the other subsequent foals in respect of their hair characters.

TELEGONY. The Pastoralists' review (Australia), Issue Vol. 16 No. 10. 15th December 1906
Sir, -Your issue of last month touches on the interesting subject of telegony and the conflict of views between scientists and practical stockbreeders. Among the former, Herbert Spencer and Darwin believed in telegony, while Weissman and Cossar Ewart would return a verdict of "not proven." The difficulty appears to be to distinguish between atavism and telegony, and between the former and discontinuous variation. Cossar Ewart made experiments by crossing the zebra site Matopo with mares of various breeds, one of which was a black Island of Rum pony Mulatto. She bred a hybrid, and was then put to a grey Arab, of pure lineage, from the Crabbet Park stud. The resulting foal had certain zebra-like markings on the croup and elsewhere, but Cossar Ewart discounts this by his theory, shared by Darwin, that the prehistoric horse was a striped animal. In 1897 40 per cent, of pure Arab foals dropped at Crabbet Park had a dorsal band, and one had, in addition, stripes on fore and hind legs.

To my mind the value of Ewart's experiments is somewhat discounted by the fact that hybrids are almost invariably sterile, hence characteristics might not he handed on by the sire of a hybrid, which might become in evidence with a female of his own race, but dissimilar variety. If telegony can be scientifically proved it is absolutely necessary that the female should be perfectly pure, and I think this could be guaranteed with sheep better than with any other animal. Artificial impregnation having become a proved success in horse-breeding, it would be interesting to make use of it in experiments with that animal, as by doing so the factor of "mental impression" on the dam could be eliminated, as it would be unnecessary for her to see the horse used in the first instance. Most of us who have been breeding stock have seen instances that appeared to support telegony, but in cases I have observed there was an element of doubt owing to lack of proof of purity of breeding in the dam. At any rate, if telegony is a fact, which I believe is the case, it is not invariable or even common; among a large number of cattle-dogs I have bred I have not seen a case of it.

With regard to stock of other kinds, a station manager has, with few exceptions, little chance to experiment exhaustively, but were some owners, like Sir S. McCaughey or others, to take the thing up, the vexed question could probably be set at rest, and stockbreeders all over the world would be interested and instructed. - H. W. GRIMANI-SMITH. Canal Creek Warwick.

A celebrated case to which Darwin attached importance was that of Lord Morton's mare, reported to the Royal Society in 1820. This mare, after bearing a hybrid by a quagga (a striped equine related to the zebra) produced, to a black Arabian horse, three-foals showing a number of stripes, and in one of them more stripes were present than in the quagga hybrid. This seems at first sight strong evidence in. favour of "infection" of the mare by the early quagga mate. But it appears that stripes are frequently seen in high-caste Arab horses, and colts cross-bred from such and other breeds of horse sometimes present far more distinct bars across the legs and other zebra-like markings than were seen in the late offspring of Lord Morton's Arabian mare. The fact appears to be that all the living species of the horse family (horses, asses, quaggas and zebras) are descended from an ancestry of "striped" equines, and are liable occasionally to "throw back" to their striped ancestry, more or less.

Professor Cossar Ewart determined some years ago to submit the matter to direct experiment, and has related his results in a book ("The Penycuik Experiments," 1899). The South African equine called the quagga, which was that used by Lord Morton, having become extinct, Professor Ewart made use of a richly striped Burchell's zebra. Thirty mares put to this animal produced seventeen hybrids, and subsequently these mares, put to horse stallions, produced twenty purebred foals. All the zebra hybrids were richly and very distinctly striped. Of the twenty later purebred horse-foals from the same mares three only presented stripe-like markings at birth, and these were few and indistinct. They disappeared when the foal's coat was shed. Their mothers were Highland mares. But the value of the faint striping in these three instances as evidence in support of telegony is at once destroyed by the fact that Professor Ewart obtained at the same time purebred foals from similar Highland mares which had never seen a zebra. Two of these purebred Highland foals showed stripes at birth, and one acquired stripes later; and further, whilst the stripes on the foals born after hybrids had been produced by their mothers disappeared with the foal's coat, the stripes on the three purebred colts whose mothers had never been near a zebra persisted for a longer period. Similar experiments confirmed these results, showing that traces of striping are no more likely to-occur on the offspring of a mare which has previously produced a mule with a zebra or an ass, than one on whose dam has neither seen nor been near to a zebra or an ass. Lord Morton's case thus falls to the ground.

Breeders of dogs are (or were) even more thoroughly convinced of the fact of telegony than breeders of horses. But- Sir Everett Millais, who devoted thirty years to the breeding of dogs and experiments on this question, states that he has never seen a case of telegony. And recent experiments of the most definite kind support his conclusion. Dalmatians, deerhounds, and retrievers have been used in these experiments. Many such experiments in telegony are accidentally or unwittingly made every year with dogs. An undesired crossing of two breeds takes place, but when subsequent pure breeding takes place no "telegonic" infection of the mother is observed. Cases believed in as due to telegony have on examination proved to be due to the carelessness of stablemen, denied or concealed by them, a dog having been allowed to escape temporarily from the kennels or to enter them uninvited. Another curious case was that of a rickety spaniel pappy, which was exhibited a few years ago at the Zoological Society and believed by the exhibitor to owe its bandy legs to “telegonic" infection of the mother by a dachshund, with which she was supposed to have mated a year or more before being put to the father of the spaniel. Its true nature was at once recognised by the experts present, the bandy legs being those caused by "rickets," and not like those of the well-known dachshund breed.

It is a remarkable proof of the belief in telegony - though not a proof of the reality of telegony - that amongst breeders of horses and dogs the selling value of a dam which has borne young to an inferior sire or to one of a distinct species, is largely diminished as compared with that of a dam which has been mated with a first-rate sire of her own breed. Darwin himself was led, by his enquiries into a similar occurrence in plants, to favour the notion that a sire could so "infect" a mare that her offspring by a later sire would in some instances show traces of the characters of the earlier sire.

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