EUGENE M. MCCARTHY, PHD GENETICS
Note: Horse hybrids, that is hybrid crosses involving the various members of Family Equidae, are more common than many people suppose. Some of these crosses occur in a state of nature on an ongoing basis. Others (not very well documented) are quite strange, such as cow-horse hybrids (see cross listing below).
|Zonkey - Zebra x Donkey|
Note: According to McClintock and Mochi (1976), the chromosome counts of zebras are 2n = 46 (E. grevyi), 2n = 44 (E. burchelli) and 2n = 32 (E. zebra). The domestic horse (E. caballus) has 2n = 64, and the ass (E. asinus), 2n = 62. The count for Przewalski’s Horse (E. ferus) is 2n = 66.
Note: Duff and Lawson (2004) list "wild ass" as the English common name of Equus africanus, stating that the range of the animal in question is northeastern Sudan, northeastern Ethiopia, and northern Somalia, the region where the animal often accorded the common name "Somali Wild Ass" resides. However, "wild ass" is a name traditionally associated with E. asinus, which is often equated with "domestic donkey." Actually, the wild animal most closely resembling the domestic donkey is often referred to as the "Nubian wild ass." It is often said to be extinct, but it's uncertain whether wild populations of the Nubian have been in existence since prehistoric times or whether currently existing wild populations are simply feral domestic asses (asses have been domesticated in Egypt and the Middle East for some 6,000 years). Its appearance differs from that of the Somali wild ass primarily in that it lacks the zebra-like leg striping of the latter. E. asinus was originally widespread in northern Africa and southwestern Asia. To avoid confusion, then, instead of following Duff and Lawson, I here will assign E. asinus the common names "Domestic Donkey | Nubian Wild Ass" and will call E. africanus "Somali Wild Ass."
Equus africanus [Somali Wild Ass]
× Equus asinus [Domestic Donkey | Nubian Wild Ass] CANHR. HPF(♀♀). Gray (1971, p. 94) notes two wild female hybrids "showed normal viability and fertility." Pocock (1911b, 993-994) says, "intermediate forms indeed, with very narrow spinal and shoulder stripes and a dusky patch on the ear, connect the Somali Ass with ordinary domestic varieties; and in all the many foals born in the [Zoological] Gardens [of London] between our Somaliland Ass and domestic asses of English and Spanish breeds, the ear-patch, shoulder and spinal stripes were present as in the dams." Natural hybridization is known to occur and is considered a threat to E. africanus. See: Equus asinus × E. grevyi. Rörig 1903 (p. 219); International Zoo Yearbook 1962 (p. 232), 1965 (p. 341). Internet Citations: ALWA.
× Equus burchelli (♀) [Zebra] In 1911, Pocock reported the cross Equus africanus × E. quagga as having occurred at the London Zoo. However, the alleged quagga mother he pictures (fig. 203, p. 992) is indistinguishable from an ordinary Burchell’s Zebra. See: Equus africanus × E. asinus; E. asinus × E. grevyi. Flower 1929a (p. 253); International Zoo Yearbook 1960 (p. 262), 1981 (p. 325); Pocock 1911b†.
× Equus quagga [Quagga] See: Equus africanus × E. burchelli.
× Equus zebra (♀) [Mountain Zebra] CHR. DRS. Gestation in one case took about 390 days. A hybrid foal from an E. africanus stallion bred to an E. zebra mare had two transverse shoulder stripes, leg bands and zebra-like ear stripes. Flower 1929a (p. 253); McClintock and Mochi (1976); Pocock 1911b; Rörig 1903 (p. 219).
Equus asinus [Domestic Donkey | Nubian Wild Ass] (2n = 62)
× Bos taurus [European Domestic Cattle] See the separate article "Jumarts."
× Equus burchelli (↔ usu. ♂) [Zebra] CHR. ENHI. HPF(♀♀). See the separate article "Zonkey."
× Equus caballus (↔ usu. ♀) [Domestic Horse] CHR. LFH. See the separate article "Mules (Donkey-horse hybrids)."
× Equus grevyi (↔ usu. ♂) [Grevy’s Zebra] CHR. Hybrids between male Grevy’s zebras and female asses are easily obtained and grow quickly. However, the hybrids are of low fertility. The reciprocal cross also occurs. Certain facts suggest the animal known as the Somali Wild Ass (E. africanus) is this hybrid: The first thing to consider is that the putative parents come into contact in the geographic region where E. asinus is known to occur. True wild asses disappeared from most of their African range during Roman times, from Asia, even earlier (Groves 1986), but the ass remains a fairly common beast of burden throughout its original range and feral stocks are widespread. Thus, feral herds of E. asinus exist in northeastern Africa (and, until recently, what may have been primevally wild populations in the form of the Nubian Wild Ass). Southward dispersal of these wild or feral asses would be expected to bring them into contact with E. grevyi in the Horn of Africa. The Somali Wild Ass occurs only in this expected region of contact. Moreover, wild E. africanus are known to interbreed with feral E. asinus (See Internet citation ALWA) and feral donkeys run with zebra herds in eastern Africa (Kingdon 1979, p. 143). E. asinus × E. grevyi hybrids are well known and have the same appearance as E. africanus (in particular the zebra-like leg stripes). E. africanus is therefore geographically intermediate and seemingly identical to captive hybrids of known parentage. So it is a PHP of this cross. This view is further supported by the facts that the Somali Wild Ass is rare (classified as critically endangered by the IUCN) and variable—typical characteristics of hybrids (also reports of Somali wild asses in captivity note that they are difficult to breed, which suggests they suffer from the reduced fertility often associated with hybridity). The hypothesis can be tested both by genetic testing and by observing whether E. africanus populations become more grevyi-like as the range of E. grevyi is approached, which would strongly suggest the existence a hybrid zone. Riley (1911) states that the average gestation time for production of Grevy’s zebra-ass hybrids in an ass dam is 387 days. Antonius 1944b, 1951a†; Churcher 1993 (p. 6); International Zoo Yearbook 1962 (p. 232); Flower 1929a (p. 253); Kaminski 1970; Mann 1938; Riley 1910†, 1911; Roberts 1929; Rommel 1913†; Rząśnicki 1930†, 1936†; Zuckermann 1953. Internet Citations: NATZ.
× Equus hemionus (↔) [Kulan | Mongolian Wild Ass | Dziggetai] CHR. Kulans probably come into contact with feral asses in Mongolia and northern China. Both male and female hybrids have been reported. Bannikov 1948; Brentjes 1969; de Lavison 1863; Flower 1929a (p. 254); International Zoo Yearbook 1990 (p. 468); Przibram 1910; Rörig 1903 (p. 218); Wagner 1863 (p. 85).
× Equus kiang [Kiang] A male hybrid was born in the London Zoological Gardens on Sept. 27, 1920. Flower 1929a (p. 254).
× Equus onager [Onager] CHR. Antonius 1944a†; International Zoo Yearbook 1970 (p. 267).
× Equus zebra (↔ usu. ♀) [Mountain Zebra] CANHR. CON: southern Africa. HPF(♂&♀). Short (1967) notes that the ejaculates of some hybrids contain spermatozoa. This partial fertility suggests (by Haldane’s Rule) that female hybrids, too, will sometimes be partially fertile. Penzhorn (1985) reported a natural hybrid produced by a mating feral jackass and an E. zebra mare. Ewart (in Riley) says that in about 1775, Lord Clive bred the first recorded zebra hybrid by crossing a female Mountain zebra (which he brought with him on returning from India) with a common ass, and that about a quarter of a century later, in 1801, a similar horse hybrid was bred in Italy. Antonius 1934b†, 1944a†, 1951a†; Benirschke 1964, 1967; Benirschke et al. 1964; Brown and Jenkins 1987; Chang et al. 1969; Geoffroy Saint-Hilaire 1806, 1808; Gray 1972; Flower 1929a (p. 253), 1931; International Zoo Yearbook 1966 (p. 398); MacClintock and Mochi 1976; Penzhorn 1985, 1988; Przibram 1910†; Riley 1911 (p. 229); Rząśnicki 1930†.
× Ovis aries [Domestic Sheep] Gesner (Historiae Animalium, Liber I, de Quadrupedibus viviparis, 1551, p. 19) writes, "At this time, in the palace of the king of France [probably Henri II (reigned 1547-1559)], an animal was raised, which in its anterior portion was like a donkey, but in its posterior parts, like a sheep. And, except in being of a dual nature, it followed each of these two distinct types of animals in all things" (translated by E.M. McCarthy, Original Latin). This animal, if it ever truly existed, would probably have been kept in the menagerie that Henri created at the Chateau de Madrid, his palace in the Bois de Boulogne. In addition, an old news report mentions such a hybrid, aged 16 years, as being present at the Ohio State Agricultural Fair held in Dayton, Ohio in September 1853. For an extensive discussion of a related cross, see also the separate article on cow-horse hybrids.
Note: In regions where they interface, hybridization occurs between all of the various populations treated as races of Equus burchelli (Grubb 1981, Fig. 5). Kingdon (1979) suggested that certain populations treated as races of E. burchelli are actually the products of hybridization between other such populations. Thus, he suggests hybridization between boehmi and crawshayi produced zambeziensis, and that crossing between burchelli and crawshayi produced both antiquorum and chapmanni.
Note: In crosses of Equus burchelli with other zebras, the striping patterns of the hybrids are generally intermediate between those of the parents (Grubb 1981). In crosses with the horse or ass, the striping in the hybrids is narrower than in the E. burchelli parents (Grubb 1981).
Equus burchelli [Burchell’s Zebra] (2n = 44)
See also: Equus asinus.
× Equus caballus (↔ usu. ♀) [Domestic Horse] See the separate article on zebra-horse hybrids.
× Equus ferus [Przewalski’s Horse] CHR. DRS. Treus et al. 1963.
× Equus grevyi [Grevy’s Zebra] CANHR. CON: Kenya. HPF(♀♀). A probable natural hybrid was reported by Keast (1965) Striping of hybrids is complete down to the hooves, though E. burchelli has white shanks and pasterns. Male hybrids produce abnormal spermatozoa. Keast (1965, p. 58) says a Kenyan taxidermist possessed the skin of a probable hybrid, which may have been of natural origin. Grubb (1981, p. 6) notes that these hybrids resemble E. zebra. These zebras differ with respect to karyotype. Benirschke 1977; Grubb 1981 (p. 6); International Zoo Yearbook 1990 (p. 469); Keast 1965; Ryder et al. 1978.
× Equus hemionus [Kulan | Mongolian Wild Ass | Dziggetai] CHR. DRS. Gray (1873, p. 38) said there was a stuffed specimen of this hybrid in the British Museum. Flower 1929a (p. 253); International Zoo Yearbook 1973 (p. 336), 1975 (p. 380); Kingdon 1979 (p. 138†); Wagner 1863 (p. 85).
× Equus quagga [Quagga] See: Equus burchelli × E. caballus.
× Equus zebra (♀) [Mountain Zebra] CHR. CON: Namibia, Angola. Most hybrids are female. LFH. Hybrid foals resemble E. burchelli, except for their larger ears and their hindquarters pattern. They have no dewlap. Wilson and Reeder (2005) state that "Equus wardi Ridgeway, 1910 is a hybrid between E. burchelli and E. zebra." Shortridge (1934), quoting Flower, says such a hybrid survived several years in the Paris Botanic Gardens. Flower 1929a (p. 253); McClintock and Mochi (1976); International Zoo Yearbook 1979 (p. 371), 1989 (p. 336); Pocock 1909. ×Syncerus caffer [African Buffalo] Hybrids of this type have not been reported. However, as a matter solely of speculation, the appearance of a wildebeest (Connochaetes sp.) seems to correspond with what might be expected from this cross.
Equus caballus [Domestic Horse] (2n = 64)
See also: Equus asinus; E. burchelli and the separate article about cow-horse hybrids.
× Alces alces (♂) [Moose | Elk] See the separate article about moose-horse hybrids.
× Camelus sp. [Camel] See the separate article "Camel × Horse."
× Capra hircus [Domestic Goat] The following brief announcement is from page 3 (column 3) of the April 15, 1899 issue of The Meridional, a newspaper published in Abbeville, Louisiana (Access Source): "What promises to prove an interesting freak of nature is now the property of F. P. Beauxis, of this town and will be on exhibition at his store after tomorrow. It is a goat-colt, if we may use the term, being an animal with the head and mane of a goat, and the body of a colt. It makes a noise very much like a goat. Its mane is soft as silk and the fore legs are somewhat peculiar shape having a foot very much like that of an ape." For an extensive discussion of a related cross, see also the separate article on cow-horse hybrids. Also see: Equus asinus × Ovis aries.
× Cervus elaphus [Red Deer | Elk | Wapiti] See the separate article on deer-horse hybrids.
× Equus ferus (♀) [Przewalski’s Horse] CANHR. CON: Mongolia. HPF(♀♀). E. caballus and E. ferus, which have often been treated as conspecific, are now usually treated separately. Wherever these horses come into contact in the wild, hybridization is likely. The IUCN (Internet Citations: FEREQ) says that for wild, reintroduced populations of Przewalski’s horse, which is endangered, hybridization is the primary threat to their survival. Moreover, it is suspected that captive stocks of so-called "E. ferus" are actually composed primarily of animals that are, at least in some degree, domestic horse hybrids. A backcross female (to an E. ferus male) had 2n = 65 and was very like E. ferus. A backcross of the same type was indistinguishable in appearance from E. caballus (with the exception of a prominent dorsal stripe). Ahrens and Stranzinger 2005; Allen and Short 1997†; Boyd and Houpt 1994; Hatami-Monazah and Pandit 1979; International Zoo Yearbook 1962 (p. 232), 1967 (p. 315), 1967 (p. 304) , 1969 (p. 232), 1970 (p. 266), 1971 (p. 278), 1973 (p. 336); Koulischer and Frechkop 1966; Ryder et al. 1978; Trommerhausen-Smith et al. 1979; Volf 2003; Wilson and Reeder 2005 (p. 1018).
× Equus grevyi (♂) [Grevy’s Zebra] CAONHR. CON: eastern and southern Africa. Grevy’s stallions do not readily mate with mares, and more services are required to achieve pregnancy than in the case, say of a jackass serving a mare. There is a high rate of spontaneous abortion, and surviving Grevy’s-horse hybrids of both sexes are of very low fertility (sterile?). However, Keast (1965, p. 58) notes that natural hybridization has been repeatedly reported. These horse hybrids are said always to be chocolate brown, whatever the color of the parent horse, with the Grevyi striping pattern. They also have the zebra’s tufted tail. Antonius 1944b, 1951a; Breen and Gill 1991; Crew and Smith 1930; Churcher 1993; Gray 1972; King 1967; King et al. 1966; Kingdon 1979 (pp. 138; 143)†; Lus 1938†; Mann 1938; Riley 1911; Roberts 1929; Rommel 1913†; Rzaśnicki 1936†; Short 1967; Webb 1952.
× Equus hemionus (♂) [Kulan | Mongolian Wild Ass | Dziggetai] CHR. Kulan-horse hybrids resemble kulans more than horses, but their tails and manes are long. Ackermann 1898 (p. 45); Antonius 1932a; Bannikov 1948; Przibram 1910; Rörig 1903 (p. 218).
× Equus onager (↔) [Onager] CHR. Brentjes 1969†; Rörig 1903.
× Equus quagga [Quagga] CHR. Bell 1837 (p. 392) states that "the Earl of Morton being desirous of obtaining a breed between the Horse and the Quagga, selected a young Mare of seven-eighths Arabian blood and a fine male of the latter species; and the produce was a female hybrid. The same Mare had afterwards first a Filly and afterwards a Colt by a fine black Arabian Horse. They both resembled the Quagga in the dark line along the back, the stripes across the forehead and the bars across the legs; in the Filly the mane was short stiff and upright, like that of the Quagga; in the Colt it was long, but so stiff as to arch upwards and hang clear of the sides of the neck; in other respects they were nearly pure Arabian as might have been expected from fifteen-sixteenths Arabian blood."
× Equus zebra (↔) [Mountain Zebra] CHR. CON: Africa. Reports of mountain-zebra/domestic horse hybrids are scanty indeed. Rörig 1903 (p. 219).
× Felis cattus [Domestic Cat] See the separate article “A Horse-cat Hybrid?.”
× Homo sapiens [Human] See the separate article “Human-horse Hybrids (Centaurs).”
× Macropus sp. On page 27 of the December 21, 1934, issue of The Advertiser, an Australian newspaper published in Adelaide, South Australia (source), an article entitled "FIFTY YEARS AGO, From "The Advertiser" December 22, 1884" quoted the following from a fifty-year-old report: “There is on view in Hindley street a living freak of nature, which is stated to be "half horse and half kangaroo." The animal, which is two years and ten months old, was bred on the Wild Horse Plains. The only portion of the creature that at all resembles a kangaroo is the near foreleg, which appears to be withered, and the hoof of which is parted.” However, this distant cross seems not to have been reported elsewhere, nor can even the report quoted be unambiguously interpreted as being consistent with such a hybrid. Mere deformity might just as well account for the phenotype in question.
× Odocoileus virginianus [White-tailed Deer] See the separate article on deer-horse hybrids.
× Sus scrofa [Domestic Pig] See the separate article "A Pig-horse Hybrid?."
Equus ferus [Przewalski’s Horse] (2n = 66) Przewalski’s horse hybrids are listed above under: Equus burchelli; E. caballus. E. ferus is often treated as a subspecies of Equus caballus.
Equus grevyi [Grevy’s Zebra] (2n = 46)
See also: Equus africanus; E. asinus; E. burchelli; E. caballus.
× Equus zebra (♀) [Mountain Zebra] CANHR. CON: eastern Africa. According to Groves (1974), attempts to breed Grevy’s stallions to mountain zebra mares are associated with high rates of abortion. However, entirely viable hybrids have been produced. For example, Gray (1972, p. 108) says a hybrid was born without difficulty at Schönnbrunn Zoo. It developed rapidly, and had "a lively temperament." King (1965, Fig. 10) describes a probable natural hybrid. International Zoo Yearbook 1973 (p. 336); Kingdon 1979 (p. 138†); McClintock and Mochi (1976); Rząśnicki 1930, 1951†.
× Ursus arctos [Brown Bear] See the separate article "A Bear-horse Hybrid?"
Equus hemionus [Kulan | Mongolian Wild Ass | Dziggetai] (2n = 54)
See also: Equus asinus; E. burchelli; E. caballus.
× Equus kiang (♀) [Kiang] CHR. CON: northern China? HPF. International Zoo Yearbook 1978 (p. 391), 1984/1984 (p. 545).
× Equus onager (↔) [Onager] CHR. DRS. According to the IUCN (Internet Citation: ONAG1), supposed onagers reintroduced in Israel are actually hybrids of this type. Flower 1929a (p. 254); International Zoo Yearbook 1973 (p. 336), 1975 (p. 380), 1978 (p. 391), 1980 (p. 437), 1981 (p. 324), 1982 (p. 427), 1984/1984 (p. 545), 1988 (p. 474).
× Equus zebra (♀) [Mountain Zebra] CHR. Hybrids were bred at London Zoo in the 1830s. Rörig 1903 (p. 218); Flower 1929a (p. 253).
Equus kiang [Kiang] See: Equus asinus; E. hemionus.
Equus onager [Onager] See: Equus asinus; E. hemionus.
Equus quagga [Quagga] See: Equus burchelli× E. caballus.Equus zebra [Mountain Zebra] (2n = 32) See: Equus africanus; E. asinus; E. burchelli; E. burchelli × E. grevyi, E. grevyi; E. hemionus.
By the same author: Handbook of Avian Hybrids of the World, Oxford University Press (2006).
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