The Variation of Animals and Plants under Domestication — Volume 1

Turning to the Old World, some European dogs closely resemble the wolf; thus the shepherd dog of the plains of Hungary is white or reddish-brown, has a sharp nose, short, erect ears, shaggy coat, and bushy tail, and so much resembles a wolf that Mr. Paget, who gives this description, says he has known a Hungarian mistake a wolf for one of his own dogs. Jeitteles, also, remarks on the close similarity of the Hungarian dog and wolf. Shepherd dogs in Italy must anciently have closely resembled wolves, for Columella (vii. 12) advises that white dogs be kept, adding, "pastor album probat, ne pro lupo canem feriat." Several accounts have been given of dogs and wolves crossing naturally; and Pliny asserts that the Gauls tied their female dogs in the woods that they might cross with wolves. (1/19. Paget 'Travels in Hungary and Transylvania' volume 1 page 501. Jeitteles 'Fauna Hungariae Superioris' 1862 s. 13. See Pliny 'History of the World' (English translation) 8th book ch. 40 about the Gauls crossing their dogs. See also Aristotle 'Hist. Animal.' Lib. 8 c. 28. For good evidence about wolves and dogs naturally crossing near the Pyrenees, see M. Mauduyt 'Du Loup et de ses Races' Poitiers, 1851; also Pallas in 'Acta Acad. St. Petersburgh' 1780 part 2 page 94.) The European wolf differs slightly from that of North America, and has been ranked by many naturalists as a distinct species. The common wolf of India is also by some esteemed as a third species, and here again we find a marked resemblance between the pariah dogs of certain districts of India and the Indian wolf. (1/20. I give this on excellent authority, namely Mr. Blyth (under the signature of Zoophilus) in the 'Indian Sporting Review' October 1856 page 134. Mr. Blyth states that he was struck with the resemblance between a brush-tailed race of pariah-dogs, north-west of Cawnpore, and the Indian wolf. He gives corroborative evidence with respect to the dogs of the valley of the Nerbudda.)

With respect to Jackals, Isidore Geoffroy Saint-Hilaire (1/21. For numerous and interesting details on the resemblance of dogs and jackals see Isid. Geoffroy St. — Hilaire 'Hist. Nat. Gen.' 1860 tome 3 page 101. See also 'Hist. Nat. des Mammiferes' par Prof. Gervais, 1855 tome 2 page 60.) says that not one constant difference can be pointed out between their structure and that of the smaller races of dogs. They agree closely in habits: jackals, when tamed and called by their master, wag their tails, lick his hands, crouch, and throw themselves on their backs; they smell at the tails of other dogs, and void their urine sideways; they roll on carrion or on animals which they have killed; and, lastly, when in high spirits, they run round in circles or in a figure of eight, with their tails between their legs. (1/22. Also Guldenstadt 'Nov. Comment. Acad. Petrop.' tome 20 pro anno 1775 page 449. Also Salvin in 'Land and Water' October 1869.) A number of excellent naturalists, from the time of Guldenstadt to that of Ehrenberg, Hemprich, and Cretzschmar, have expressed themselves in the strongest terms with respect to the resemblance of the half-domestic dogs of Asia and Egypt to jackals. M. Nordmann, for instance, says, "Les chiens d'Awhasie ressemblent etonnamment a des chacals." Ehrenberg (1/23. Quoted by De Blainville in his 'Osteographie, Canidae' pages 79, 98.) asserts that the domestic dogs of Lower Egypt, and certain mummied dogs, have for their wild type a species of wolf (C. lupaster) of the country; whereas the domestic dogs of Nubia and certain other mummied dogs have the closest relation to a wild species of the same country, viz. C. sabbar, which is only a form of the common jackal. Pallas asserts that jackals and dogs sometimes naturally cross in the East; and a case is on record in Algeria. (1/24. See Pallas in 'Act. Acad. St. Petersburgh' 1780 part 2 page 91. For Algeria, see Isid. Geoffroy St. — Hilaire 'Hist. Nat. Gen.' tome 3 page 177. In both countries it is the male jackal which pairs with female domestic dogs.) The greater number of naturalists divide the jackals of Asia and Africa into several species, but some few rank them all as one.

I may add that the domestic dogs on the coast of Guinea are fox-like animals, and are dumb. (1/25. John Barbut 'Description of the Coast of Guinea in 1746.') On the east coast of Africa, between latitude 4 deg and 6 deg south, and about ten days' journey in the interior, a semi-domestic dog, as the Rev. S. Erhardt informs me, is kept, which the natives assert is derived from a similar wild animal. Lichtenstein (1/26. 'Travels in South Africa' volume 2 page 272.) says that the dogs of the Bosjemans present a striking resemblance even in colour (excepting the black stripe down the back) with the C. mesomelas of South Africa. Mr. E. Layard informs me that he has seen a Caffre dog which closely resembled an Esquimaux dog. In Australia the Dingo is both domesticated and wild; though this animal may have been introduced aboriginally by man, yet it must be considered as almost an endemic form, for its remains have been found in a similar state of preservation and associated with extinct mammals, so that its introduction must have been ancient. (1/27. Selwyn, Geology of Victoria; 'Journal of Geolog. Soc.' volume 14 1858 page 536 and volume 16 1860 page 148; and Prof. M'Coy in 'Annals and Mag. of Nat. Hist.' (3rd series) volume 9 1862 page 147. The Dingo differs from the dogs of the central Polynesian islands. Dieffenbach remarks ('Travels' volume 2 page 45) that the native New Zealand dog also differs from the Dingo.)

From this resemblance of the half-domesticated dogs in several countries to the wild species still living there, — from the facility with which they can often be crossed together, — from even half-tamed animals being so much valued by savages, — and from the other circumstances previously remarked on which favour their domestication, it is highly probable that the domestic dogs of the world are descended from two well-defined species of wolf (viz. C. lupus and C. latrans), and from two or three other doubtful species (namely, the European, Indian, and North African wolves); from at least one or two South American canine species; from several races or species of jackal; and perhaps from one or more extinct species. Although it is possible or even probable that domesticated dogs, introduced into any country and bred there for many generations, might acquire some of the characters proper to the aboriginal Canidae of the country, we can hardly thus account for introduced dogs having given rise to two breeds in the same country, resembling two of its aboriginal species, as in the above- given cases of Guiana and of North America. (1/28. These latter remarks afford, I think, a sufficient answer to some criticisms by Mr. Wallace, on the multiple origin of dogs, given in Lyell's 'Principles of Geology' 1872 volume 2 page 295.)

It cannot be objected to the view of several canine species having been anciently domesticated, that these animals are tamed with difficulty: facts have been already given on this head, but I may add that the young of the Canis primaevus of India were tamed by Mr. Hodgson (1/29. 'Proceedings Zoological Soc.' 1833 page 112. See also on the taming of the common wolf, L. Lloyd 'Scandinavian Adventures' 1854 volume 1 page 460. With respect to the jackal, see Prof. Gervais 'Hist. Nat. Mamm.' tome 2 page 61. With respect to the aguara of Paraguay see Rengger's work.), and became as sensible of caresses, and manifested as much intelligence, as any sporting dog of the same age. There is not much difference, as we have already shown and shall further see, in habits between the domestic dogs of the North American Indians and the wolves of that country, or between the Eastern pariah dogs and jackals, or between the dogs which have run wild in various countries and the several natural species of the family. The habit of barking, however, which is almost universal with domesticated dogs, forms an exception, as it does not characterise a single natural species of the family, though I am assured that the Canis latrans of North America utters a noise which closely approaches a bark. But this habit is soon lost by dogs when they become feral and is soon reacquired when they are again domesticated. The case of the wild dogs on the island of Juan Fernandez having become dumb has often been quoted, and there is reason to believe (130. Roulin, in 'Mem. present. par divers Savans' tome 6 page 341.) that the dumbness ensued in the course of thirty-three years; on the other hand, dogs taken from this island by Ulloa slowly reacquired the habit of barking. The Mackenzie-river dogs, of the Canis latrans type, when brought to England, never learned to bark properly; but one born in the Zoological Gardens (1/31. Martin 'History of the Dog' page 14.) "made his voice sound as loudly as any other dog of the same age and size." According to Professor Nillson (1/32. Quoted by L. Lloyd in 'Field Sports of North of Europe' volume 1 page 387.), a wolf-whelp reared by a bitch barks. I. Geoffroy Saint-Hilaire exhibited a jackal which barked with the same tone as any common dog. (1/33. Quatrefages 'Soc. d'Acclimat.' May 11, 1863 page 7.) An interesting account has been given by Mr. G. Clarke (1/34. 'Annals and Mag of Nat. Hist.' volume 15 1845 page 140.) of some dogs run wild on Juan de Nova, in the Indian Ocean; "they had entirely lost the faculty of barking; they had no inclination for the company of other dogs, nor did they acquire their voice" during a captivity of several months. On the island they "congregate in vast packs, and catch sea-birds with as much address as foxes could display." The feral dogs of La Plata have not become dumb; they are of large size, hunt singly or in packs, and burrow holes for their young. (1/35. Azara 'Voyages dans l'Amer. Merid.' tome 1 page 381; his account is fully confirmed by Rengger. Quatrefages gives an account of a bitch brought from Jerusalem to France which burrowed a hole and littered in it. See 'Discours, Exposition des Races Canines' 1865 page 3.) In these habits the feral dogs of La Plata resemble wolves and jackals; both of which hunt either singly or in packs, and burrow holes. (1/36. With respect to wolves burrowing holes see Richardson 'Fauna Boreali-Americana' page 64; and Bechstein 'Naturgeschichte Deutschlands' b. 1 s. 617.) These feral dogs have not become uniform in colour on Juan Fernandez, Juan de Nova, or La Plata. (1/37. See Poeppig 'Reise in Chile' b. 1 s. 290; Mr. G. Clarke, as above; and Rengger, s. 155.) In Cuba the feral dogs are described by Poeppig as nearly all mouse-coloured, with short ears and light-blue eyes. In St. Domingo, Col. Ham. Smith says (1/38. Dogs, 'Nat. Library' volume 10 page 121; an endemic South American dog seems also to have become feral in this island. See Gosse 'Jamaica' page 340.) that the feral dogs are very large, like greyhounds, of a uniform pale blue-ash, with small ears, and large light-brown eyes. Even the wild Dingo, though so anciently naturalised in Australia, "varies considerably in colour," as I am informed by Mr. P.P. King: a half-bred Dingo reared in England (1/39. Low 'Domesticated Animals' page 650.) showed signs of wishing to burrow.

[From the several foregoing facts we see that reversion in the feral state gives no indication of the colour or size of the aboriginal parent-species. One fact, however, with respect to the colouring of domestic dogs, I at one time hoped might have thrown some light on their origin; and it is worth giving, as showing how colouring follows laws, even in so anciently and thoroughly domesticated an animal as the dog. Black dogs with tan-coloured feet, whatever breed they may belong to, almost invariably have a tan- coloured spot on the upper and inner corners of each eye, and their lips are generally thus coloured. I have seen only two exceptions to this rule, namely, in a spaniel and terrier. Dogs of a light-brown colour often have a lighter, yellowish-brown spot over the eyes; sometimes the spot is white, and in a mongrel terrier the spot was black. Mr. Waring kindly examined for me a stud of fifteen greyhounds in Suffolk: eleven of them were black, or black and white, or brindled, and these had no eye-spots; but three were red and one slaty-blue, and these four had dark-coloured spots over their eyes. Although the spots thus sometimes differ in colour, they strongly tend to be tan-coloured; this is proved by my having seen four spaniels, a setter, two Yorkshire shepherd dogs, a large mongrel, and some fox-hounds, coloured black and white, with not a trace of tan-colour, excepting the spots over the eyes, and sometimes a little on the feet. These latter cases, and many others, show plainly that the colour of the feet and the eye-spots are in some way correlated. I have noticed, in various breeds, every gradation, from the whole face being tan-coloured, to a complete ring round the eyes, to a minute spot over the inner and upper corners. The spots occur in various sub-breeds of terriers and spaniels; in setters; in hounds of various kinds, including the turnspit-like German badger-hound; in shepherd dogs; in a mongrel, of which neither parent had the spots; in one pure bulldog, though the spots were in this case almost white; and in greyhounds, — but true black-and-tan greyhounds are excessively rare; nevertheless I have been assured by Mr. Warwick, that one ran at the Caledonian Champion meeting of April 1860, and was "marked precisely like a black-and-tan terrier." This dog, or another exactly the same colour, ran at the Scottish National Club on the 21st of March, 1865; and I hear from Mr. C.M. Browne, that "there was no reason either on the sire or dam side for the appearance of this unusual colour." Mr. Swinhoe at my request looked at the dogs in China, at Amoy, and he soon noticed a brown dog with yellow spots over the eyes. Colonel H. Smith (1/40. 'The Naturalist Library' Dogs, volume 10 pages 4, 19.) figures the magnificent black mastiff of Thibet with a tan-coloured stripe over the eyes, feet, and chaps; and what is more singular, he figures the Alco, or native domestic dog of Mexico, as black and white, with narrow tan-coloured rings round the eyes; at the Exhibition of dogs in London, May 1863, a so-called forest dog from North-West Mexico was shown, which had pale tan-coloured spots over the eyes. The occurrence of these tan-coloured spots in dogs of such extremely different breeds, living in various parts of the world, makes the fact highly remarkable.

We shall hereafter see, especially in the chapter on Pigeons, that coloured marks are strongly inherited, and that they often aid us in discovering the primitive forms of our domestic races. Hence, if any wild canine species had distinctly exhibited the tan-coloured spots over the eyes, it might have been argued that this was the parent-form of nearly all our domestic races. But after looking at many coloured plates, and through the whole collection of skins in the British Museum, I can find no species thus marked. It is no doubt possible that some extinct species was thus coloured. On the other hand, in looking at the various species, there seems to be a tolerably plain correlation between tan-coloured legs and face; and less frequently between black legs and a black face; and this general rule of colouring explains to a certain extent the above-given cases of correlation between the eye-spots and the colour of the feet. Moreover, some jackals and foxes have a trace of a white ring round their eyes, as in C. mesomelas, C. aureus, and (judging from Colonel H. Smith's drawing) in C. alopex, and C. thaleb. Other species have a trace of a black line over the corners of the eyes, as in C. variegatus, cinereo-variegatus, and fulvus, and the wild Dingo. Hence I am inclined to conclude that a tendency for tan-coloured spots to appear over the eyes in the various breeds of dogs, is analogous to the case observed by Desmarest, namely, that when any white appears on a dog the tip of the tail is always white, "de maniere a rappeler la tache terminale de meme couleur, qui caracterise la plupart des Canides sauvages." (1/41. Quoted by Prof. Gervais 'Hist. Nat. Mamm.' tome 2 page 66.) This rule, however, as I am assured by Mr. Jesse, does not invariably hold good.]

It has been objected that our domestic dogs cannot be descended from wolves or jackals, because their periods of gestation are different. The supposed difference rests on statements made by Buffon, Gilibert, Bechstein, and others; but these are now known to be erroneous; and the period is found to agree in the wolf, jackal, and dog, as closely as could be expected, for it is often in some degree variable. (1/42. J. Hunter shows that the long period of seventy-three days given by Buffon is easily explained by the bitch having received the dog many times during a period of sixteen days ('Phil. Transact.' 1787 page 353). Hunter found that the gestation of a mongrel from wolf and dog ('Phil. Transact.' 1789 page 160) apparently was sixty-three days, for she received the dog more than once. The period of a mongrel dog and jackal was fifty-nine days. Fred. Cuvier found the period of gestation of the wolf to be ('Dict. Class. d'Hist. Nat.' tome 4 page 8) two months and a few days, which agrees with the dog. Isid G. St. — Hilaire, who has discussed the whole subject, and from whom I quote Bellingeri, states ('Hist. Nat. Gen.' tome 3 page 112) that in the Jardin des Plantes the period of the jackal has been found to be from sixty to sixty-three days, exactly as with the dog.) Tessier, who has closely attended to this subject, allows a difference of four days in the gestation of the dog. The Rev. W.D. Fox has given me three carefully recorded cases of retrievers, in which the bitch was put only once to the dog; and not counting this day, but counting that of parturition, the periods were fifty-nine, sixty-two, and sixty-seven days. The average period is sixty-three days; but Bellingeri states that this applies only to large dogs; and that for small races it is from sixty to sixty-three days; Mr. Eyton of Eyton, who has had much experience with dogs, also informs me that the time is apt to be longer with large than with small dogs.

F. Cuvier has objected that the jackal would not have been domesticated on account of its offensive smell; but savages are not sensitive in this respect. The degree of odour, also, differs in the different kinds of jackal (1/43. See Isid. Geoffroy St. — Hilaire 'Hist. Nat. Gen.' tome 3 page 112, on the odour of jackals. Col. Ham. Smith in 'Nat. Lib.' volume 10 page 289.); and Colonel H. Smith makes a sectional division of the group with one character dependent on not being offensive. On the other hand, dogs — for instance, rough and smooth terriers — differ much in this respect; and M. Godron states that the hairless so-called Turkish dog is more odoriferous than other dogs. Isidore Geoffroy (1/44. Quoted by Quatrefages in 'Bull. Soc. d'Acclimat.' May 11, 1863.) gave to a dog the same odour as that from a jackal by feeding it on raw flesh.

The belief that our dogs are descended from wolves, jackals, South American Canidae, and other species, suggests a far more important difficulty. These animals in their undomesticated state, judging from a widely-spread analogy, would have been in some degree sterile if intercrossed; and such sterility will be admitted as almost certain by all those who believe that the lessened fertility of crossed forms is an infallible criterion of specific distinctness. Anyhow these animals keep distinct in the countries which they inhabit in common. On the other hand, all domestic dogs, which are here supposed to be descended from several distinct species, are, as far as is known, mutually fertile together. But, as Broca has well remarked (1/45. 'Journal de la Physiologie' tome 2 page 385.), the fertility of successive generations of mongrel dogs has never been scrutinised with that care which is thought indispensable when species are crossed. The few facts leading to the conclusion that the sexual feelings and reproductive powers differ in the several races of the dog when crossed are (passing over mere size as rendering propagation difficult) as follows: the Mexican Alco (1/46. See Mr. R. Hill's excellent account of this breed in Gosse's 'Jamaica' page 338; Rengger 'Saugethiere von Paraguay' s. 153. With respect to Spitz dogs, see Bechstein's 'Naturgesch. Deutschlands' 1801 b. 1 s. 638. With respect to Dr. Hodgkin's statement made before Brit. Assoc. see 'The Zoologist' volume 4 for 1845-46 page 1097.) apparently dislikes dogs of other kinds, but this perhaps is not strictly a sexual feeling; the hairless endemic dog of Paraguay, according to Rengger, mixes less with the European races than these do with each other; the Spitz dog in Germany is said to receive the fox more readily than do other breeds; and Dr. Hodgkin states that a female Dingo in England attracted the male wild foxes. If these latter statements can be trusted, they prove some degree of sexual difference in the breeds of the dog. But the fact remains that our domestic dogs, differing so widely as they do in external structure, are far more fertile together than we have reason to believe their supposed wild parents would have been. Pallas assumes (1/47. 'Acta Acad. St. Petersburgh' 1780 part 2 pages 84, 100.) that a long course of domestication eliminates that sterility which the parent-species would have exhibited if only lately captured; no distinct facts are recorded in support of this hypothesis; but the evidence seems to me so strong (independently of the evidence derived from other domesticated animals) in favour of our domestic dogs having descended from several wild stocks, that I am inclined to admit the truth of this hypothesis.

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, 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.