Many members of the dog family are also capable of interbreeding and producing fertile offspring. This depends on how closely they are related. Molecular analysis divides the dog family into 4 divisions:
1. Wolf-like canids including the domestic dog, dingo, grey wolf, coyote, and jackal
2. South American canids
3. Old and New World red-fox-like canids, e.g. red fox and kit fox
4. Monotypic species, e.g. bat-eared fox, raccoon dog
The wolf, coyote, jackal, and domestic dog (including the dingo) have 78 chromosomes arranged in 39 pairs. They are interfertile (unless size or behavioural differences obstruct mating) and produce fertile offspring. The wolf, coyote, and golden jackal diverged around 3 to 4 million years ago. The African Wild Dog also has 78 chromosomes, but is considered distinct enough (with different physical features) to be placed in its own genus. Other members of the dog family diverged 7 to 10 million years ago and are less closely related. These cannot hybridise with the wolf-like canids and they have different numbers of chromosomes: the yellow Jackal has 74 chromosomes, the red fox has 38 chromosomes, the raccoon dog has 42 chromosomes, the Fennec fox has 64 chromosomes. Differences in chromosome number are not always a barrier to producing viable hybrids; it depends on the gene combinations in the hybrid and whether these allow an embryo to develop. However, large differences in chromosome number make female hybrids poorly fertile and male hybrids sterile. In the case of dogs and foxes, it appears that viable embryos are not formed.
According to “Dogs In Britain, A Description of All native Breeds and Most Foreign Breeds in Britain” by Clifford LB Hubbard, 1948 “So also do we find that in the Americas the aboriginal dog stock has been slightly influenced by the coyote (Canis latrans),whilst in the east the blood of the jackal (Canis aureus) has to an appreciable extent pulled most of the semi-wild Pariah Dogs into a uniformity of shape noticeably apparent even to the most casual observer. It will be seen from these references to the wolf, jackal and coyote that cross-breeding is not only possible between these animals and the true dog but is, in fact, fairly frequent; less so to-day, of course, when dog breeding is so strongly controlled by man and his fancies for this or that shape, but originally quite common. All members of the genus Canis interbreed freely and their progeny are as a rule quite fertile, whereas dogs do not mate with such animals as hyaenas, African ‘wild dogs’, foxes and the like. A few alleged fox-dog hybrids have been mentioned from time to time, but the details concerning the matings have not been authentic.”
People wanting to improve domestic dogs have sometimes bred them back to wolves; as well as compensating for genetic problems in domestic dogs, the wolf-dog hybrids tend to be dominant and less domesticable. Grey wolves have been crossed with wolf-like dogs e.g. German Shepherds, Malamutes. Many become a problem in adulthood, usually because the owner expects them to act like a domestic dog and is not prepared for their more wolf-like behaviour. The Saarloos breed is derived from such crosses. Parts of Europe, the wolf naturally hybridises with stray domestic dogs resulting in mongrel populations. The Australian Dingo (a feral, rather than wild, species) also hybridises freely with domestic dogs. Harmsworth Natural History (1910, main authors R Lydekker, Sir H Johnston & Prof JR Ainsworth-Davis) notes The dingo breeds freely with the various European dogs introduced by the colonists” and classes the dingo as a descendent of Asian domesticated dogs rather than a distinct species. Clifford LB Hubbard wrote in “Dogs In Britain, A Description of All native Breeds and Most Foreign Breeds in Britain” (1948): When early Scots settlers went to Australia some small working Sheepdogs (probably Welsh Sheepdogs or working Border Collies) were taken with them to help round up their then small flocks. The foundation stock appear to have been a British working Sheep-dog, of one or other of the above breeds, named “Caesar” and a cross-bred Dingo-Collie called “Kelpie”: of the resulting black-tan-and-red litter one bitch so resembled its dam that it was named “Kelpie II”
According to the “Illustrated Natural History” by the Rev JG Wood (1853, 1874): When Wolves and dogs are domesticated in the same residence, a mutual attachment will often spring up between them, although they naturally bear the bitterest hatred to each other. A mixed offspring is sometimes the result of this curious friendship, and it is said that these half-bred animals are more powerful and courageous than the ordinary dog. Mr. Palliser possessed a remarkably fine animal of this kind, the father of which was a white wolf and the mother an ordinary Indian dog. Its fur was white, like that of its wolf-parent. When ‘Ishmah’ as the dog was named, was first purchased from its Indian owners, he was so terrified at the white face of his new master, that he always ran away whenever he saw him, and could not be persuaded to come within two hundred yards. Ishmah was then tied up with a cord, but the moment that he was left to himself he held the cord to the ground with his paw, severed it in an instant with his sharp teeth, leaped out of the window, and dashed off to his former owners. After a while, however, he became reconciled to his white master, and proved to be a most faithful and useful ally; dragging a small sledge that contained the heavier necessaries of a hunter’s life, and partaking with his master all the pleasures and privations of a nomad existence. On account of wolfish ancestry, he was rather apt to run off and play with the young wolves instead of attending to his duty, but was never induced to throw off his allegiance. On one occasion the dog saved the life of his master by lying close to him on a bitterly freezing night, and with his long warm fur preserving him from the terrible death by frost.
December 13th, 1901 (various): BOSTOCK ANIMALS FIGHT. Bears, Lions and Hybrid Wolves in a Scrap—One Bear Killed. BOSTON, Dec. 12.—A fierce combat took place at Bostock’s animal arena today' between three black bears from India, two forest-bred lions and two hybrid wolves. As a result one bear is dead. Mr. Johnson, the menagerie superintendent, discovered the battle soon after it started, and called to his assistance Merman Weelon. Arthur Mullen and others, who entered the cage with hot irons and revolvers to quell the disturbance. Mullen was bitten in the calf of one leg, but his Injury is not dangerous. It was twelve minutes before the animals were separated. Then it was found that a 6-year-old bear, very valuable, had been bitten through the jugular vein. He died soon after. The hybrids are a cross between American bloodhounds and Siberian wolves and have been trained by Mme. Beaufort. They are very savage and probably they started the battle.There was great excitement during the combat, and Mr. Bostock regards it as fortunate that it occurred when no audience was present.
According to “Dogs In Britain, A Description of All native Breeds and Most Foreign Breeds in Britain” by Clifford LB Hubbard, 1948: The true Husky is not a large dog; the dogs of 27 inches height and even more are cross-bred with wolves or ‘white’ breeds (husky/hound cross-bred dogs) […]The Timber-Wolf Dog is not confined to one district but is spread throughout the areas from Alaska to north of Hudson Bay; the Yukon basin appears to be its adopted home to the largest extent. It is, as its name applies, a first-cross between the true Husky and the timber-wolf of the North American backwoods; gaunt, ferocious, the largest of its type and the most wolfish of all sled dogs. Owing to its size and strength it is capable of being a good leader or ‘king’ dog for a team, but otherwise it is not the best type for general use, as is the Husky itself which can be used in summer for pack carrying. Its heavy build handicaps it on very long journeys though for short heavy loads the breed is frequently in demand.
In general shape it is nearer the timber-wolf than the Husky; longer in back and coat (the coat does not stand stiffly out); longer couplings and greater angulation of the hindlegs; a more loping gait which is not the best for long journeys; the head usually having non-erect ears, scarcely any stop and a slightly longer muzzle. Colour is grey, or grey or sable with white throat, chest, belly, feet and tail-tip; rarely pure white. In height it stands about 27-28 inches, and in weight ranges from 85 to 100 pounds (outstanding specimens equal the height of an Irish Wolfhound and weigh as much as eight stone). The tail usually hangs low and is heavy in brush.
For many years, the German Shepherd Dog (called Alsatian in the UK due to two wars with the Germans) was believed to be a wolf-dog hybrid. Hubbard wrote in 1948: It must be understood that in the 1920s the breed was not only the best-loved but also the most-hates; its nom-de-mal of [Alsatian] Wolfdog, its lupine features, and the existence about that time of too many highly strung and nervous specimens (which whenever they bit anyone were referred in the Press as dangerous wolf-hybrids) together contributed to its decline. […] people have been told by those who should have known better, or who never will know better, that the German Shepherd Dog is half wolf […] should be authoritatively told that whatever wolf blood was in the breed about seventy-five years ago is bred entirely out now." (This is an interesting parallel with the types of dogs - based on appearance - demonised in the late 20th and early 21st century)
Dog genes also flow into wild wolf populations. Wolves acquired the genetic mutation for dark coat colour through mating with domestic dogs (Marco Musiani, University of Calgary, Canada and Greg Barsh, Stanford University, California in Science Magazine, 2009). Light colour coats predominate in wolves living on open tundra, but the dark colour is common in wolves in wooded or forested areas. The results came from a genetic analysis of wolf populations in Europe and North America. The studies suggest black wolves appeared after grey wolves bred with prehistoric domestic dogs that accompanied settlers into the continent 10,000-15,000 years ago. Those domestic dogs became extinct, however the genes from the extinct domestic dogs live on in the wolf population. Modern domestic dogs in North America are descended from European dogs over the last 500 years (some of which have again been crossed with wolves). A trait developed by humans in domestic dogs, which were developed from wolves, has found its way into the wolf population. Because wolves don't rely on camouflage for hunting success, there must be other reasons the mutation is beneficial to the woodland wolves. The protein involved in the coat colour mutation may be linked to the immune system. The change to the immune system may be the real advantage while the pigmentaiton is a side-effect.
In parts of Europe where wolves have been killed by shepherds, the remaining wolves have interbred with stray dogs. The strays have tended to be the blonde-furred sheepdogs (used to guard the flocks and merge in with the sheep) resulting in strains of cream-furred wolves. Unfortunately in this situation, low wolf numbers mean they risk being mongrelised out of existence and supplanted by feral dog populations.
DOG/FOX ALLEGED HYBRIDS
According to “Dogs In Britain, A Description of All native Breeds and Most Foreign Breeds in Britain” by Clifford LB Hubbard, 1948 “A few alleged fox-dog hybrids have been mentioned from time to time, but the details concerning the matings have not been authentic. Though perhaps strange, considering their external appearance, it perfectly true that the dog and the fox do not breed together and that because of the very structural and osteological differences which prohibit their free cohabitation they are scientifically classified apart, the dog being as we know of the genus Canis and the fox being in the genus Vulpes. At one time, until comparatively recently [recently to 1948] in fact, the fox was called Canis vulpes (and therefore accepted as a species of the true dog family), but with the advancement of cynological study and research it was finally given a genus of its own, being now generally believed to be quite apart from the true dog”.
An unconfirmed female fox-dog hybrid (terrier/fox) has been reported in the UK, unfortunately it was euthanized when the owner died because no suitable home could be found for it. Although gamekeeper folklore claims that Terrier bitches can produce offspring with dog Foxes, there are no authenticated hybrids between domestic dogs and Red Foxes. According to anecdote, "dox" (fox/bitch) hybrids are stronger and more vigourous than either parent and can be bred to other doxes. Thus far, all reported dox hybrids have turned out to be natural variation in the domestic dog. The great genetic mismatch between red foxes and domestic dogs makes the dox hybrid no more real than a cabbit (cat-rabbit) hybrid.
There has been a reported cross between a domestic dog and a South American fox, but the latter was most likely a fox-like wolf and not a true fox.
Harmsworth Natural History (1910, main authors R Lydekker, Sir H Johnston & Prof JR Ainsworth-Davis) notes the inability of the dog to form hybrids with the fox: ”Additional testimony that foxes have nothing to do with the origin of domesticated dogs is afforded by the fact that hybrids between the fox and the dog, notwithstanding numerous specimens of supposed hybrids of this sort which are from time to time brought to notice, are of extreme rarity, if not altogether unknown.
There was a report of a dox in Saskatchewan, Canada. This was believed to be the result of a miniature sheltie bitch with a wild fox (fox species not identified). One out of the three offspring survived, this being a sterile female that resembled a fox. However, the variability of dogs in appearance make it impossible to determine whether an animal is hybrid based on looks alone. I have not found genetic evidence to support the hybrid identity.
Coy-dogs (male coyote/female dog) have occurred e.g. a Coyote/Irish Setter mix (resembling a liver-coloured, spaniel-like dog) has been reported. Coydogs were once believed to be present in large numbers in Pennsylvania due to Coyotes being in decline and domestic dogs being available as mates. Most were probably naturally occurring red or blond Coyotes or feral dogs. The breeding cycles of dogs and coyotes are not synchronized and studies indicate that Coyote-Dog interbreeding is uncommon. If it such interbreeding was common, the Coyote population would acquire more and more dog-like traits.
The converse mating results in a Dogote and there is currently one known Dogote which arose from a male German Shepherd/female coyote mating in the wild. Hybrid pups were found after a female coyote was shot. The adult Dogote resembled a German Shepherd in colour.
DINGO/COYOTE & DINGO/WOLF HYBRIDS
Coyotes have also been crossed with Australian dingoes. This cross was made at a time the dingo was considered to be different species to the domestic dog.
The dingo has also been crossed with the wolf. According to Vol 1 of "Animal Life and the World of Nature" (1902-1903), Mr HC Brooke owned a female wolf which had become quite tame and was an excellent mother. This wolf was the mother of the wolf-dingo hybrid owned by Lord Walter Rothschild. The book also depicted a tame dingo owned and exhibited by Mrs HC Brooke, so it's likely that one of those dingoes was the sire of Lord Rothschild's hybrid.
Coy-wolves (Coyote/Wolf) have occurred in captivity or, rarely, in the wild where the choice of same-species mates has been limited. Coyote/Red Wolf hybrids have been found. Some consider the American Red Wolf is not to a true species because it can hybridize with both the Grey Wolf and the Coyote; however it is now known that hybridization between species (in general) happens more often than previously thought. Some consider it a Grey Wolf/Coyote hybrid and use this argument to prevent conservation of the Red Wolf. Some hybridization occurred when pure Red Wolves were in decline and interbred with more numerous Coyotes. More recent studies in many mammals shows that true species can and do hybridize and that the species boundary is preserved by geographic or behavioural separation (wolves will kill coyotes), not by genetic separation.
WOLF/JACKAL & DOG/JACKAL HYBRIDS
The Wolf and Jackal (an African wild dog) can interbreed and produce fertile hybrid offspring. In Russia, Dog/Jackal hybrids were bred as sniffer dogs because Jackals have a superior sense of smell and Huskies are good cold climate dogs. Male Jackal pups had to be fostered on a Husky bitch in order to imprint the Jackals on dogs. Female Jackals accepted male Huskies more easily. The half-bred Jackal-Dogs were hard to train and were bred back to Huskies to produce quarter-bred hybrids (quadroons). These hybrids were small, agile, trainable and had excellent noses. They are called Sulimov Dogs after their creator and may one day be registered as a working breed of dog.
EASTERN COYOTES – COYOTE X WOLF X DOG HYBRIDS
So-called Coywolves are living and evolving in forested areas of eastern North America, from Florida to Labrador. Scientists prefer the term Eastern Coyote. Because genetic tests in 2015 found them to be a mix of three species: predominantly coyote, but with a percentage of wolf and/or dog genes. The percentages vary depending on geographical location and on the sensitivity of the test used. Coyotes in the Northeast were 60%-84% coyote, 8%-25% wolf and 8%-11% dog. This mixture changes as you move south or east. In Virginia, these Coyotes are, on average, 85% coyote, 2% wolf and 13% dog. Those from the Deep South average 91% coyote, 4% wolf and 5% dog. None of the tested animals were purely coywolf, but a few were coydog with almost no distinguishable wolf genes. This means there is no “coywolf” species, but there are populations of coyotes with variable amounts of wolf and/or dog genes.
All Eastern Coyotes have evidence of past hybridization, but no evidence that they are still mating with dogs or wolves. Although genetically compatible, the three species only interbreed if they don’t have a same-species mate available. When isolated from their own species, the reproductive drive is so strong that they will overcome natural enmity to mate with a related species. Outside of this situation, wolves and large dogs will kill coyotes, while wolves may also kill domestic dogs.
There’s evidence of multiple hybridisation events in the three species. The gene for black coat colour in North American wolves and coyotes (and absent from European wolves) originated in dogs taken to the continent by prehistoric Native Americans. There was further hybridization in the 1800s when high demand for sled dogs meant that trappers and gold-miners used any suitably large dog for that purpose. Husky bitches on heat were sometimes tethered so that wolves would mate with them and produce offspring that were hardier than domestic dogs.
Around 100 years ago, wolf populations in the Great Lakes area were so low that wolves resorted to mating with coyotes. The hierarchical structure of wolf packs, where only the dominant pair breeds, means subordinate animals leave the pack in order to breed, and some evidently mated with coyotes. The estimated date of the most recent hybridization that created Eastern Ccoyotes was about 50 years ago with dogs. There are dark-coloured Eastern Coyotes in North Carolina resembling German Shepherd Dogs in colour and pattern. The colouration is believed to have entered the coyote gene pool around 50 years previously, although there have been deliberate German Shepherd/coyote hybrids in the last two decades, some of which were allegedly let loose into the wild due to their un-dog-like temperaments.
In the east, more recent coyote/dog hybridization has occurred at the leading edge of the coyote colonisation wave. Female coyotes reached upstate New York, where there were plenty of feral dogs, but no other coyotes. Modern Eastern Coyotes, and modern Great Lakes wolves, both have plenty of mates available these days and no longer interbreed. Coyotes also expanded north into Alaska, but there is currently no sign of hybridization in that range extension. In Central America, they have expanded from Mexico and into Central America. There are photographs of very dog-like coyotes at the southern edge of this coyote colonisation wave.
Hybridization across compatible species is a natural evolutionary phenomenon. It creates new genetic combinations, some of which have a survival advantage over the parent species. Wolf genes make the Eastern Coyotes a bit larger and more able to tackle the abundant deer in forested areas. It may make them more sociable and more likely to hunt in groups rather than pairs. Dog genes might make the coyote more bold and better able to survive in proximity to humans. It’s possible that several distinct geographical varieties of Eastern Coyotes will arise, and if they don’t interbreed they may evolve into separate species.
This extract from the "Book of The Dog" by William Youatt (1846) demonstrates nineteenth century beliefs about hybridisation between different species. It was known that a dog and wolf could be crossed and that the union would produce vicious mongrels, but there were claims of hybrids now known to be impossible: "There are some naturalists that even go so far as to state that the different varieties of dogs are sprung from, or compounded of, various animals, as the hyaena, jackal, wolf, and fox. The philosophic John Hunter commenced a series of experiments upon this interesting subject, and was forced to acknowledge that "the dog may be the wolf tamed, and the jackal may probably be the dog returned to his wild state. The ancient Cynegetical [dog] writers were not only acquainted with the cross between the wolf and dog, but also boasted the possession of breeds of animals, supposed to have been derived from a connection with the lion and tiger. The Hyrcanian dog, although savage and powerful beast, was rendered much more formidable in battle, or in conflict with other animals, by his fabled cross with the tiger. In corroboration of this singular, but not less fabulous belief, Pliny states that the inhabitants of India take pleasure in having dog bitches lined by the wild tigers, and to facilitate this union, they are in the habit of tieing them when in heat out in the woods, so that the male tigers may visit them."
In "Origin of Species" (1859) Charles Darwin wrote The German Spitz dog unites more easily than other dogs with foxes [...] certain South American indigenous domestic dogs do not readily cross with European dogs ..." and in "Darwinism An Exposition Of The Theory Of Natural Selection With Some Of Its Applications" (1889), Alfred Russel Wallace commented: "Dogs have been frequently crossed with wolves and with jackals, and their hybrid offspring have been found to be fertile _inter se_ to the third or fourth generation, and then usually to show some signs of sterility or of deterioration." (The deterioration was due to inbreeding among the hybrid offspring)
In "The Variation Of Animals And Plants Under Domestication" Charles Darwin wrote: "Buffon got four successive generations from the wolf and dog, and the mongrels were perfectly fertile together. [...] 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 [attributed to inbreeding]. ('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.) [...] Mr. Philip P. King, after ample opportunities of observation, informs me that the Dingo and European dogs often cross in Australia. [...] 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. [...] We have already seen how often savages cross their dogs with wild native species; and Pennant gives a curious account ('History of Quadrupeds' 1793 volume 1 page 238.) of the manner in which Fochabers, in Scotland, was stocked "with a multitude of curs of a most wolfish aspect" from a single hybrid-wolf brought into that district. [...] the jackal is prepotent over the dog, as is stated by Flourens, who made many crosses between these animals; and this was likewise the case with a hybrid which I once saw between a jackal and a terrier. "
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