Cattle Hybrids

Cattle, sheep, goats (Family Bovidae)

(Antelope are listed separately >>)

Mammalian Hybrids




Cattle hybrids:

A zubron
(European bison × cattle)

Abbreviation key >>

Internet citations >>

Glossary >>

Ammotragus lervia [Barbary Sheep | Aoudad]
× Capra hircus (♀) [Domestic Goat] CHR. HPF(♀♀). Breeding contact between feral domestic goats and Barbary sheep may occur both in the southwestern U.S., where barbary sheep are introduced, and in northern Africa, where they are endemic. Hybrids of both sexes have been produced, not only by artificial insemination, but also via ordinary coitus. Most are inviable, but it seems that in all cases where hybrids have been born alive the dam was a goat. Bunch et al. say a female hybrid was “quite adept at jumping heights in excess of 10 feet,” which exceeds the normal capabilities of either parental type. Hybrids have 2n = 57 with only three unpaired chromosomes. Females have been successfully backcrossed to Barbary rams. Haltenorth (1961) reports that at Halle Zoo in Germany a female hybrid crossed twice with an alpine ibex (Capra ibex), but that the first offspring died soon after, and the second, before, birth. Brentjes 1968; Bunch et al. 1977; Dain 1980; de Lavison 1863; Gray 1972; Gray and Simpson 1980 (pp. 3, 4); Haltenorth 1961; Milne-Edwards 1896; Moore et al. 1981; Petzsch 1957b, 1957a, 1957c, 1959.
× Ovis aries (♀) [Domestic sheep] Moore et al. (1981) obtained fertile embryos via artificial insemination, but none developed.

Anoa depressicornis [Lowland Anoa]
× Anoa quarlesi [Mountain Anoa] CHR?? NHR. CON: northern Sulawesi. Schreiber et al. suggest gene flow may explain a lack of distinct variation in the wild. They also say (p. 173) that “an anoa photographed by M. Pangau (Tropenzentrum Univerisät Göttingen) in the central region of the northern peninsula of Sulawesi appears to combine certain phenotypic traits of both the quarlesi and depressicornis morphotypes.” These anoas meet in an elongated contact zone (ACZ?) in the foothills of the northern peninsula. Burton et al. (p. 30) say there are at least two explanations for discrepancies between reported mtDNA and allozyme distance results: “First, mountain and lowland anoa are well-separated species but hybridization in the captive population caused the observed pattern of protein variation…Alternatively, the differences in the mtDNA sequences could be a chance effect caused by the sole female founder of the B. quarlesi species belonging to another mitochondrial lineage.” Schreiber et al. (p. 173) say that the low allozyme distances observed between so-called mountain and lowland anoas in zoos could result from hybridization, but they discount this explanation. Burton et al. 2005; Schreiber et al. 1999.

Note: McDonald (1978) suggested Bison bison emerged 4,000-5,000 years ago as the product of hybridization between two prehistoric bison (B. antiquus and B. occidentalis). Geist 1991b (pp. 295, 297).

Bison bison [American Bison]
See also: Bos frontalis × B. mutus; Bos frontalis × B. indicus.
× Bison bonasus (↔) [European Bison] CHR. DRS. HPF(♂&♀). These hybrids were commonly produced in the former Soviet Union. Lowek (1999, p. 1163) says B. bonasus populations now reestablished in the Caucasus “contain a small admixture of genes from B. bison.” Ansell also says that some hybridization of this type occurred in the efforts to save B. bonasus from extinction. Regarding this situation, the IUCN (Internet Citations: BIBON) says “A particular problem concerning the management of extant populations of European bison is the existence of hybrid herds, especially European × American bison hybrids living in the Caucasus. Two free-living hybrid herds have been established in the Caucasus Mountains, in close proximity to reintroduced free-living herds of the pure blood Lowland-Caucasian line. There are fears that all these animals will crossbreed, creating a mixture of various genotypes. According to Russian authors, the distances between herds are not so great, but the configuration of mountain ridges and valleys make it impossible for contact between them [to occur]. There are also two small semi-free herds of European and American bison hybrids in Toksove Forest Park (St. Petersburg) and the Mordovia Wildlife Reserve (Pucek et al. 2004).” These taxa are sometimes lumped. Iwanow and Philiptschenko describe three-way hybrids with Bos taurus. Ansell 1972 (p. 54); International Zoo Yearbook 1969 (p. 240), 1970 (p. 276), 1972 (p. 340), 1973 (p. 342), 1975 (p. 387), 1977 (p. 329), 1978 (p. 399), 1979 (p. 377), 1980 (p. 445), 1981 (p. 330); Iwanow and Philiptschenko 1916; Kliment 1989a, 1989b; Verkaar et al. 2002, 2003 (p. 565).
× Bos frontalis (♀) [Gayal | Mithan] CHR. DRS. Zoological Society of London 1878
× Bos grunniens (♂) [Domestic Yak] HPF(♀♀). This hybrid, known as a yakalo, was produced in the 1920s (Deakin et al. 1935). Pictures of takins (Budorcas taxicolor) have been misrepresented, either by mistake or hoax, as being this hybrid (e.g., Internet Citations: GENPR).
× Bos taurus (↔ usu. ♂) [European Domestic Cattle] CHR. HPF(♀♀). See the separate article “American Bison × Cattle.”
× Equus caballus [Horse] See the separate article “Jumarts.”

Note: The European Bison (Bison bonasus) is extremely similar in appearance to the American Bison (Bison bison), so it seems likely that any cross reported for the latter would also be feasible for the former, and vice-versa.

Bison bonasus [European Bison]
See also: Bison bison × Bos taurus.
× Bos taurus (↔ usu. ♀) [European Domestic Cattle] HPF(♀♀). CHR. This hybrid is known as a zubron. Ackermann 1898 (pp. 71-72); Baranov and Zakharov 1997; Dehnel 1960; Dziurdzik 1978; Fedyk and Krasinska 1980; Kania et al. 1982; Klevezal and Pucek 1987; Kliment 1989a, 1989b; Kobryńczuk and Krasińska 1987, 1991; Krasińska 1963, 1967, 1969, 1971a, 1971b, 1988; Krasińska and Suminski 1981; Ward et al 1999.

Note: Bos frontalis is thought to be the domesticated form of Bos gaurus.

Bos frontalis [Gayal | Mithan]
See also: Bison bison
× Bos gaurus [Gaur] CHR. HPF(♂&♀). Verkaar et al. 2003 (p. 565).
× Bos indicus (↔ usu. ♂) [Zebu] CHR. HPF(♀♀). Zebu females are often afraid of Gayal bulls and have to be blindfolded before mating will occur (see Simoons 1968). In Bhutan, where these hybrids are regularly produced, the traditional breeding program specifies that female hybrids be backcrossed to Zebu producing B₁, B₂, B₃, and B₄ generations. The hybrids of Mithan-Siri crosses in each generation are phenotypically distinct and the Bhutanese herders have a different name for each. Females of the B₄ generation are considered full-blooded zebu and are mated again to a mithan bull to start the cycle again. This practice has been carried on for at least a century, probably far longer. Female hybrids are superior milk producers; males make powerful draft animals, but are sterile (in the F₁ generation). The hump is less prominent in hybrids than in B. indicus. Winter et al. found no mature spermatozoa in F₁ testes, whereas spermatids occurred in B₁, and all stages of spermatogenesis, in B₂. Spermatogenesis appeared normal in B₃ males. At the Zoological Gardens of London, a female three-way hybrid, produced from a mating of a male American bison (Bison bison) with a female hybrid (B. frontalis ♀ × B. indicus ♂), later produced third-generation hybrids of both sexes with a B. bison male. Abraham Dee Bartlett, Superintendent of the London Zoo, gave the following account (Bartlett 1884, p. 400) of this series of crosses (the pedigree, to which the following description refers, is shown below): “In the first place, the bull Zebu (Bos indicus)was introduced to the cow Gayal (Bibos frontalis) and a female hybrid was born Oct 29, 1868 (A of pedigree). This animal (A) produced her first calf June 16, 1872, a second one Oct. 16, 1873, a third one Jan. 5, 1875, a fourth March 11, 1876, a fifth Nov. 2, 1878 these five calves were the produce of this female hybrid Gayal with the Zebu bull. She was now introduced to the male American Bison (Bison americanus), and on the 21st of May, 1881, she produced a female No. 2 (B of pedigree). It will be seen that this animal (B) is the produce not only by the intermixture of three well-marked species, but, according to our present definition, of three distinct genera. This remarkable animal the result of the triple alliance (Plate XXXIV) was last year introduced to the bull Bison and on the 12th of March, 1884 she produced a female (C of pedigree). This last individual, now eleven weeks old (Plate XXXV), is undistinguishable from a pure bred Bison of the same age.” A century later, Winter et al. (1984) conducted a modern study of the same kind of 3-way hybrids as those described by Bartlett. Ackermann 1898 (p. 70); Anonymous 1850 (p. 23); Bartlett and Bartlett 1900; Bartlett 1884; Faiz 1968; Flower 1929a (pp. 258-259); Haldane 1932; Hickman and Tenzing 1982; Kühn 1883; Namikawa et al. 1987; Phanchung et al. 2001; Sterndale 1884 (Appendix C); Winter et al. 1988a, 1988b; Zuckerman 1953 (p. 942). Internet Citations: PHAN. See also: Der zoologische Garten (1873, p. 36)

Bos gaurus [Gaur]
See also: Bos frontalis.
× Bos indicus (♀) [Zebu] CANHR. CON: southeast Asia. Bongso et al. report spontaneous hybridization of wild gaur and zebu. One theory of the origin of the Mithan (Bos frontalis) is that it is derived from this cross (Gupta et al. 1999). In Malaysia, a wild guar jumped a fence into a farm, gathered a harem of 200 cows, and impregnated many. Vietmeyer (1986) says the hybrid calves were golden-brown or black, “alert, vigourous, and feisty with handlers. At least for their first six months of life they grow 70 percent faster than cattle.” Southeast Asian Selembu cattle result from this cross. Jerdon (1874, p. 306) says Bos sylhetanus F. Cuvier, 1824 is this hybrid (also see: Anonymous 1850, p. 32; Cuvier 1819). Bongso et al. 1988; Felius 1995; Ward et al. 1999.
× Bos javanicus [Banteng | Bali Cattle] The rare Kouprey (Bos sauveli) occurs where B. gaurus and B. javanicus overlap (southeast Asia). It has characteristics of both gaurs and bantengs, so it has long been suggested that it may be a hybrid of this type. And on the basis of genetic data, Galbreath et al. (2006, 2007) argue that such is the case. But see contra: Hassanin and Ropiquet (2004, 2007a, 2007b), Hassanin et al. (2006), and Hedges et al. (2007). The IUCN rates the Kouprey as critically endangered. Internet Citations: DWIK.
× Bos taurus (♀) [European Domestic Cattle] CHR. HPF(♀♀). DRS. Riggs et al. 1997.

Video: Logging with dzo in China.

Video: Dzo (Bos grunniens × Bos indicus) in the Himalayas.

Bos grunniens [Domestic Yak]
See also: Bison bonasus.
× Bos indicus (↔) [Zebu] CHR. HPF(♀♀). A hybrid of this type is known as a dzo /(d)zō/ (pl dzo or dzos) or a zho /zhō/ (pl zho or zhos). According to Sharma (1954), there is much hybridization of yaks with zebu in Nepal, “the male progeny being infertile and the females fertile.” In these regions the offspring of yak and zebu cow are called urang, those of yak and zebu bull, dimjo. Sharma says that the urang is suited for altitudes of 2,000-3,500 m and yields more milk than either parent, and that the dimjo is a strong, swift work animal. Ackermann 1898 (p. 70); Anonymous 1850 (p. 23); International Zoo Yearbook 1965 (p. 346), 1973 (p. 341); Phillips et al. 1946; Zawadowski 1931; Zuckerman 1953 (p. 942).
× Bos mutus [Wild Yak] In a survey of the status of mammalian fauna on the Tibetan Plateau, Harris et al. (1999) say a mostly female herd of domestic yaks “elicited great interest among a number of solitary, male wild yaks in September 1997 (mating season for wild yaks), raising the specter of potential hybridization.”
× Bos taurus (♀ prob. ↔) [European Domestic Cattle] CHR. HPF(♂&♀). CON: central Asia. These hybrids are called yakows. Three yak at the Kansas City Zoo were found to have B. taurus mtDNA. Wishart et al. examined the testes of a single F₁ hybrid (yak × cow) and found spermatogonia were present, but no mature spermatozoa. However, Ivanova (1953) says, “1st generation bulls are mated either with Simmental or with hybrid cows” so at least some F₁ males are partially fertile.” In China these hybrids are kept at altitudes that are between those suitable for the two parents. Wild yaks are black or brown in color, whereas domesticated animals that are partly white have hybrid ancestry. Marco Polo during his twelfth-century travels learned that hybridization, apparently of this type, was carried out on a regular basis in eastern Asia. In his chapter describing the Kingdom of Erguiul, which lay in what is now in the northern part of China near Mongolia, he describes hybridization of cattle with what were probably domesticated yaks. His account (from Yule 1875, vol. 1, p. 266) runs as follows: “There are wild cattle in that country almost as big as elephants, splendid creatures, covered everywhere but on the back with shaggy hair a good four palms long. They are partly black, partly white, and really wonderfully fine creatures and the hair or wool is extremely fine and white, finer and whiter than silk. … There are also plenty of them tame, which have been caught young. They also cross these with the common cow, and the cattle from this cross are wonderful beasts, and better for work than other animals. These the people use commonly for burden and general work and in the plough, as well and at the latter, they will do full twice as much work as any other cattle being such very strong beasts.” Ackermann 1898 (p. 70); Aipu 1983; Felius 1995; Halnan 1989; Petzsch 1957a; Popescu 1969; Tumennasan et al. 1997; Ward et al 1999 (p. 54); Wishart et al. 1988; Zhang and Duan 1991. Internet Citations: YAKH.

Bos indicus [Zebu]
See also: Bos frontalis; B. gaurus; B. grunniens; B. mutus.
× Bos javanicus (↔) [Banteng | Bali Cattle] HPF(♀♀). Male F₁ hybrids are sterile or of very low fertility, and spermatogenesis is arrested at the late primary spermatocyte stage. In seven backcross hybrids, mean daily sperm production per gram of testis (DSPG) was lower than in purebred B. javanicus, but four (one 1/4 B. javanicus, one 3/4 B. javanicus, one from 5/8 parents B. javanicusmating inter se and one from 3/8 B. javanicus parents matinginter se) exhibited DSPG levels similar to the foundation stock. Semen from these latter seven, exhibiting complete spermatogenesis, was more variable in terms of sperm concentration, percentage of motile spermatozoa, and levels of spermatozoal abnormalities. In those hybrids where sperm production was reduced or absent, there was an elevated frequency of primary spermatocytes degenerating during late pachytene and diplotene, or of germinal cells degenerating at some later stage of development. In extreme cases, arrest of spermatogenesis is complete at the late primary spermatocyte stage. Nijman et al. analyzed hybridization of Banteng (Bos javanicus) and Zebu (Bos indicus) in southeast Asian cattle. The Indonesian Madura breed, used for the traditional bull racing competitions, have been shown by multiple molecular genetic criteria to be derived from this cross. Ward et al. say these animals were crossed on the island of Madura some 1500 years ago and that their descendants are currently distributed across Indonesia. They found high levels of zebu mtDNA (35%) in cattle from a Malaysian Bali cattle population. Similarly Kikkawa et al. say banteng mtDNA is prevalent in Indonesian zebus. Hybridization is considered a threat to the endangered Banteng. Ackermann 1898 (p. 71); Davis and Read 1985; Hoogerwerf 1970; National Research Council 1983; Kikkawa et al. 2003; McCool 1990; Nijman et al. 2003; Pathak and Kieffer 1979; Payne and Rollinson 1976; Ward et al. 1999.
× Bos mutus [Wild Yak] CAENHR(Asia). A website on the wild animals of Ladahk and Kashmir (KOUA) says wild yaks commonly cross with the Ladakhi cattle (Bos indicus). The hybrids are sure footed and used for riding and as beasts of burden in the difficult terrain, much as mules are used in other parts of the world. Hybridization between yaks has taken place throughout much of Asia. Schaller and Wulin 1996; Ward et al. 1999.
× Bos taurus [European Domestic Cattle] CHR. HPF(♂&♀). In the U.S., several new breeds are derived from this cross, important ones being the Beefmaster (Zebu × Shorthorn and Hereford), Brangus (Zebu × Angus), Charbray (Zebu × Charolais), and Santa Gertrudis (Zebu × Shorthorn). Partial resistance to rinderpest has apparently encouraged the introduction of zebu into Africa where extensive interbreeding of these types has long occurred, perhaps for millennia. Genetic analysis reveals higher concentrations of zebu genes in eastern Africa and, especially, Madagascar. The Sanga breed of southern and eastern Africa is derived from this cross. In the early twentieth century, zebu were brought to Brazil and were crossbred with European taurine (Charolais) cattle to produce a now popular breed known as the Chanchim. Craft (1938, p. 25) says the sex ratio in zebu-taurine hybrids is nearly the same as in pure B. taurus. Bongso et al. (1988) picture a three-way hybrid with Bos gaurus. Bradley et al. 1994, 1996; Canada Department of Agriculture 1961; Dollin et al. 1989, 1991a, 1991b; Epstein 1971; Frisch et al. 1997; Hanotte et al. 2000; Loftus et al. 1994; Manwell and Baker 1980; Nabours 1912; Nijman et al. 1999; Prichard 1836 (p. 141, citing von Zimmermann 1777); Sutherst et al. 1983; Verkaar et al. 2003 (p. 565).
× Bubalus bubalis (♂) [Domestic Water Buffalo] CHR?? CON: southeast Asia. Ackermann (1898, p. 69) says this cross usually fails because hybrid fetuses grow so large that they often die during delivery. However, Gray (1972, p. 125) says no progeny have been reported from this cross.

Bos javanicus [Banteng | Bali Cattle]
See also: Bos frontalis; B. gaurus; B. indicus.
× Bos taurus (♀) [European Domestic Cattle] CHR. HPF(♀♀). Male F₁ hybrids are sterile or of very low fertility and spermatogenesis is arrested at the late primary spermatocyte stage. International Zoo Yearbook 1971 (p. 286); McCool 1990; Pathak and Kieffer 1979; Shilova et al. 1990, 1991, 1992; Steklenev and Nechiporenko 1979.
× Canis familiaris [Dog] See the separate article "Dog-cow Hybrids."

Bos mutus [Wild Yak] See: Bos frontalis; B. grunniens; B. indicus.

Bos primigenius [Aurochs] (extinct)
× Bos taurus [European Domestic Cattle] According to Gotherstrom et al. (2005), genetic findings suggest that domestic cattle were first developed in the Near East, and that they were then hybridized with the aurochs to produce the modern Euopean breeds. And Beja-Pereira et al. et al. (2006) say that “We believe that contrasting emerging evidence calls now for a reevaluation of the single-origin hypothesis of the European cattle. In fact, (i) this hypothesis is based on limited DNA sampling of modern breeds from Southern Europe and Northern Africa and only six aurochsen from a single geographically restricted area in Northern Europe; (ii) the aurochs went extinct in Europe no more than 400 years ago and, hence, the cattle and local wild aurochsen coexisted for millennia during which interbreeding was possible; (iii) recently, a 4,000-year-old molar from Northern Spain, morphologically attributed to a cattle, was proved to have a mtDNA sequence of the British aurochs type, and the tooth could have belonged to a hunted aurochs morphologically misattributed to a cattle, but local domestication or cattle-aurochs hybridization cannot be excluded.”

Bos sauveli [Kouprey] See: Bos frontalis × B. javanicus.

Bos taurus [European Domestic Cattle] (2n=60)
See also: Bison bison; Bison bonasus; Bos frontalis; Bos gaurus; Bos grunniens; Bos indicus; Bos javanicus; Bos mutus; Bos primigenius.
× Alces alces(♂) [Moose | Eurasian Elk] CON: northern North America, northern Eurasia. See the separate article “Moose × Cow.”
× Bubalus bubalis (♂) [Domestic Water Buffalo] CHR?? Fu-Čžao (1959) says many hybrids have been produced in China, but this assertion seems not to have been substantiated elsewhere. Other studies report that this cross fails at an early stage of development. Kochhar et al. (2002) exposed cattle oocytes to buffalo sperm and buffalo oocytes to cattle sperm (in vitro). They found a higher cleavage rate in the former case than in the latter (86.3% vs. 40.8%). Also blastocysts hatched at a higher rate (25.9% vs. 8.8%). They did not evaluate the fate of hybrid embryos beyond the hatched blastocyst stage, they say (p. 162) that their findings “point to the viability of these pre-attachment hybrid embryos.” The directionality of the cross, on the basis of these early developmental studies, corresponds to that specified by Ackermann (1898) for a related cross (water buffalo male x zebu female). Patil and Totey (2003) also had poor results in vitro with the reciprocal cross. MacGregor (1941) says that when these animals are kept together, mixed matings are not unusual. International Zoo Yearbook 1967 (p. 321); Patil and Totey 2003; Tatham et al. 2003.
× Canis familiaris [Dog] See the separate article "Dog-cow Hybrids."
× Capra hircus [Domestic Goat] The following article appears on page 4, column 3, of the September 19, 1907, issue of the Hopkinsville Kentuckian, a newspaper published in Hopkinsville, Kentucky (source): “CURIOUS COWGOAT. Drops Dead From Heat Just as it Reached the City. A farmer from Trigg County arrived at the suburbs of Hopkinsville late Tuesday afternoon driving an animal monstrosity, half cow and half goat. When he reached the plant of the Hopkinsville Lumber Co., on 15th Street, the animal became overheated and died in the street. The owner was disconsolate, as he was bringing it to the city to sell it to the Hagenbeck show as a side show freak. [new paragraph] It was a male, black with white spots, had hair like a cow, a head like a goat, small short horns, weighed about 400 pounds and stood a little more than three feet height. It was four years old and was in good health and very fat when it was started from home. The owner said he lived 14 miles from the city, in the direction of Cadiz.” See also the separate article "Goat × Bighorn Sheep."
× Cervus elaphus (♂) [Red Deer | Elk | Wapiti] See the separate article “Red Deer × Cow.”
× Equus caballus [Horse] See the separate article “Jumarts.”
× Equus asinus [Ass] See the separate article “Jumarts.”
× Lepus europaeus [European Hare] See the separate article "Rabbit × Cow."
× Homo sapiens [Human] See the separate article “Domestic Cow × Human.”
× Macropus sp. [Kangaroo] The following notice appears on page 10, column 6, of the June 6, 1874, issue of The Argus, a newspaper published in Melbourne, Victoria, Australia (source). “The Evelyn Observer says:—‘One of those curious freaks of nature which every now and then crop up to astonish and mystify the untutored rustic and the learned savant alike occurred in the neighbourhood of Queenstown (Caledonia), a short time since. This lusus naturae was rendered the more remarkable, too, from the fact of there being several instances of a similar character occurring about the same time, viz., cows giving birth to calves minus the fore legs, and this happened, we are informed on reliable authority, in five different instances, each within a short time of the other. These monstrosities, our informant tells us, were as much like kangaroos as could be without being veritable marsupials and were possessed of two little fore paws instead of the usual and orthodox legs and feet, but they lacked that very necessary adjunct to kangaroo felicity, the tail, possessing only the comparatively useless member common to the bovine kind.’” A related report #1 >> A related report #2 >>
× Odocoileus virginianus [White-tailed Deer] An anecdotal report of such a hybrid exists (Access article).
× Oryctolagus cuniculus [Rabbit] See the separate article "Rabbit × Cow."
× Ovibos moschatus (♂) [Muskox] Ackermann (1898, p. 70) lists this cross, but the authors he cites (Bechstein 1789-1797; Bronn 1847; Hofacker and Notter 1828) seem not to mention it.
× Ovis aries [Domestic Sheep] See the separate article "Cow-sheep Hybrids."
× Ovis canadensis [Bighorn Sheep] See the separate article "Goat × Bighorn Sheep."
× Sus scrofa [Domestic Pig] There is a brief report of such a hybrid on page 6 of the February 5, 1901 issue of the El Paso Daily Herald, a newspaper published El Paso, Texas (source). It states that "A freak of nature was found by J. C. Yoakum, near Honey Grove, last week. It was half pig and half calf, the head and body being that of a calf, while the lower jaw, legs and feet were those of a hog. It was dead when found." There is also a brief mention in an advertiser dated Dec. 14, 1907 of such a hybrid, a conjoined triplet, as being for sale.
× Rangifer tarandus (♂) [Caribou | Reindeer] See the separate article “Reindeer × Cow.”
× Taurotragus oryx (♂) [Eland] CHR. CON: sub-Saharan Africa. Born out of a Friesland cow, a hybrid described by Mitchell had the dewlaps, ears, and hooves of an Eland. The gestation period was 280 days (compared to 254-277 days in T. oryx and 277-290 days in B. taurus). Boulineau 1932; Mitchell 1932; van Rooyen 1957. × Ursus arctos [Brown Bear] See the separate article about bear-cow hybrids.

Note: There are two chromosomal races of Bubalus bubalis, “river type” (2n = 50) and “swamp type” (2n = 48). In hybrids from the cross river-type male × swamp-type female, testicular biopsy revealed high levels of degenerating spermatocytes and abnormal spermatids (although all stages of spermatogenesis were in evidence). In many Asian countries hybrids of this type are commonly produced because they grow faster and have higher carcass weights. In Malaysia 50% of the beef supply is derived from this hybrid. Bongso and Hilmi 1983; Cockrill 1974; Dai et al. 1994; Winter et al. 1980 (p. 14).

Bubalus arnee [Wild Water Buffalo]
× Bubalus bubalis [Domestic Water Buffalo] ENHR(southern Asia). According to the IUCN (Internet Citations: BUBUB), which rates Bubalus arnee as endangered, “it is possible that no pure-bred Wild Water Buffaloes remain…any assessment of buffalo numbers is hampered by the difficulty of distinguishing between free-ranging domestic buffaloes, feral buffaloes, truly wild buffaloes, and hybrids between wild and other buffaloes. Individuals of Wild Water Buffalo and Domestic Water Buffalo are difficult to distinguish in some areas, and some domestic populations may be very closely related to (perhaps identical to) Wild Water Buffalo, as in Cambodia, where traditional forms of buffalo husbandry allow herds to range freely in forest areas (Timmins and Ou 2001; R. J. Timmins pers. comm. 2008).” Also see: Flamand et al. 2003.

Bubalus bubalis [Domestic Water Buffalo]
See also: Bos indicus; B. taurus; Bubalus arnee.
× Homo sapiens [Human] See the separate article “Water Buffalo × Human.”
× Syncerus caffer [African Buffalo] DRS. On the Internet, this hybrid, known as a buffalypso, is allegedly bred on game ranches. But the real buffalypso seems simply to be a new breed of water buffalo developed on the island of Trinidad without admixture from S. caffer.

Capra aegagrus [Wild Goat]
× Capra falconeri (♂) [Markhor] CAENHI(Chiltan Range, Pakistan). HPF(vh). Roberts (1977, 1997, pp. 287-288) implies that the Chiltan wild goat (C. a. chialtanensis) is derived from this cross because it is intermediate in morphology and range. In particular he notes that their horns “are intermediate in shape between the Markhor and Wild Goat.” Noting Roberts’ observation, Bouch and Groves (1990, p. 270) say that “an examination of [chialtanensis] specimens by one of us (CPG) leads him to concur with this [i.e., Roberts’] assessment.” Sterndale 1886b (p. 35).
× Capra hircus [Domestic Goat] CAENHR(Middle East, Crete). HPF(♂&♀). Shackleton (1997, p. 108) says F₁ hybrids are difficult to distinguish from pure wild goats, also known as agrimi, “in either appearance or behavior. (However, compared to agrimi, the hybrid’s coat appears to be slightly longer and interspersed with black hairs, the horns are curved outwards a little more and are more robust, and the resting tail is held more horizontal than downwards.)” Adamakopoulos et al. 1997; Roberts 1997 (p. 270).
× Ovis canadensis [Bighorn Sheep] CHR. See the separate article "Goat × Bighorn Sheep."
× Pseudois nayaur [Bharal] CHR. CON: Tajikistan, northern Pakistan. The Henry Doorly Zoological Gardens (Omaha, Nebraska, U.S.) reported hybrids of both sexes. International Zoo Yearbook 1979 (p. 382).

Capra caucasica [West Caucasian Tur]
× Capra cylindricornis [East Caucasian Tur] ENHR(central Caucasus). These turs exist only within a narrow region in the high Caucasus. Their horns differ markedly. The horns of hybrids are usually thinner than those of either parent. The hybrid zone is in Svanetiya, on the upper Ingur. Heptner et al. 1988 (pp. 833-837, Figs. 192, 193); Radde 1899; Vereshchagin 1938, 1967 (pp. 445-446).
× Capra falconeri [Markhor] CHR. DRS. International Zoo Yearbook 1974 (p. 395).
× Capra hircus (♀?) [Domestic Goat] CHR. CON: Caucasus. Gaebler 1885 (p. 242) reports a hybrid between “Capra pallasii” and domestic goat. International Zoo Yearbook 1969 (p. 243).
× Capra ibex [Alpine Ibex | Steinbock] CHR. DRS. Female hybrids have been reported. International Zoo Yearbook 1970 (p. 279), 1971 (p. 290).
× Capra sibirica [Siberian Ibex] CHR. DRS. Flower reports that forty-four hybrids were born in the Zoological Gardens of London between 1910 and 1927. However, these may refer to Capra cylindricornis × Capra sibirica, since Flower treats caucasica and cylindricornis as conspecific. Flower 1929a (p. 268); International Zoo Yearbook 1973 (p. 345).

Capra cylindricornis [East Caucasian Tur] See: Capra caucasica.

Capra falconeri [Markhor]
See also: Capra aegagrus; C. caucasica.
× Capra hircus (♀) [Domestic Goat] CHR. CON: mountains of southwest Asia. Breeders produce this cross in Pakistan. Between 1864 and 1915 twelve hybrids were born in the Zoological Gardens of London. Der Zoologische Garten 1863 (vol. 4, no. 4, p. 86); International Zoo Yearbook 1969 (p. 244). Internet Citations: TAXO.
× Capra nubiana [Nubian Goat] CHR. DRS. Ackermann (1898, p. 66) says a hybrid (“Capra megaceros × C. beden”) was born at London Zoo on April 24, 1864. C. beden is a synonym of C. nubiana, and megaceros (Kabul Markhor) is now treated as a subspecies of Capra falconeri. Flower 1929a.

Capra hircus [Domestic Goat] (2n=60)
See also: Ammotragus lervia; Capra aegagrus; C. caucasica; C. falconeri.
× Canis familaris [Domestic Dog] See the separate article "Domestic Dog × Domestic Goat"
× Capra ibex (↔ usu. ♂) [Alpine Ibex | Steinbock] CAENHR. HPF(♂&♀). CON: Alps. This cross was formerly common in zoos. The hybrids become increasingly similar to C. ibex with age. Hybridization between domestic goats and free-ranging ibexes occurs throughout much of the Alps (e.g., Aosta, Glarus, Grisons, Piedmont, Sondrio, Ticino, Valais). In the south-central Alps (Bregaglia Valley, Switzerland) feral domestic goats have bred with male ibexes. Resulting F₁ females bred again with male ibexes and a hybrid herd resulted. Recently state gamekeepers removed the domestic goats and hybrids in order to prevent interbreeding. Giacometti et al. (2004) recommend that domestic goat herds be carefully tended in regions where they might come into contact with ibexes and that such regions be surveyed to permit removal of hybrids and feral domestic goats. Hybrids are larger and heavier than ibex. Their horns are longer than those of ibex, and some males have horns lacking nodes. In some males, the first horn increment is longer than the second. In some cases the pelage color shows characteristics atypical for ibex such as obvious leg markings and dark brown color in young animals. This hybrid has been treated as a species (Capra lævicornis, Sundevall). Ackermann 1898 (pp. 67-68); Anonymous 1850 (pp. 151, 154); Ausserer 1947; Bächler 1918, 1935; Couturier 1962; Der Zoologische Garten (vol. 26, p. 346); Fitzinger 1864b; Giacometti et al. 2004; Stämpfli 1979; Stüwe and Grodinsky 1987; Wagner 1863 (p. 85).
× Capra nubiana (♂)[Nubian Goat] CHR. HPF(♀♀). CON: Middle East, northeastern Africa. These goats are crossed in Israel in efforts to produce better goat breeds. Abraham et al. 1989; Flower 1929a; International Zoo Yearbook 1969 (p. 243), 1970 (p. 279), 1971 (p. 290); Zuckerman 1953. Internet Citations: TAXO.
× Capra sibirica [Siberian Ibex] CHR. A hybrid was born in the Zoological Gardens of London on Aug. 5, 1926. This cross has been used by breeders in Pakistan. Flower 1929a. Internet Citations: TAXO.
× Capreolus capreolus (♂) [Western Roe Deer] Bechstein (1801, p. 491) lists this cross and says the offspring blend the characters of the parents. Ackermann (1898, p. 66) also lists this hybrid (citing Weidmanns Feierabende, ein neues Handbuch für Jaeger und Jagdfreunde, 1819, p. 33). See the separate article "Roe Deer × Domestic Sheep"
× Equus asinus [Domestic Donkey | Nubian Wild Ass] The early medical journal Miscellanea Curiosa (1677, vol. 8, supplement, Observation XLVI, p. 209) reports that a nanny goat, which was being kept in the same stable with donkeys, gave birth to a kid with donkey ears and a long, donkey-like tail. In other respects, "it resembled its mother." For an extensive discussion of a related cross, see also the separate article on cow-horse hybrids. Also see: Equus asinus × Ovis aries and Equus caballus × Capra hircus
× Equus caballus [Horse] 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 (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: Capra hircus × Equus asinus and Ovis aries × Equus asinus.
× Homo sapiens [Human] See the separate article “Caprinid-human hybrids.”
+ Lama glama [Llama] Although various later authors cite Pietro Rossi (1799, pp. 120-121) as having claimed that a male llama crossed with a goat doe to produce a hybrid, an examination of Rossi’s paper revealed that he merely quoted from Pietro Andrea Mattioli’s Epistolarum Medicinalium Libri Quinque (1561) to the effect that a male llama in captivity, deprived of females of its kind, mounted and inseminated a goat, but without a hybrid being produced.
× Odocoileus hemionus [Mule Deer] What seems to have been an alleged hybrid of this type is described in an old newspaper report (The Evening World, New York, December 3, 1892, p. 2, col. 7). The article says the animal in question is "part goat part deer," but does not specify the type of deer. However, due to the locale (San Francisco), the deer parent would have likely been O. hemionus. The report said the animal was male, in good health and three years of age. It looked generally like a deer but lacked antlers and had an indiscriminate diet like a goat.
× Odocoileus virginianus (♂) [White-tailed Deer] A female hybrid of this type was reported on page 17 of the May 16, 1916 issue of Farm and Fireside. The article, which includes a poor-quality photo of the alleged hybrid, reads as follows: “A New Dairy Hybrid. A Cuban subscriber of Farm and Fireside, E. H. Bogart, has a hybrid milch animal—a goat-deer cross—which is being used as a source of household milk supply. The mother of this hybrid is a pure white goat. The father is a wild deer that frequented the fields near the dwelling where the mother of the hybrid was kept. The deer and goat were seen to mate. The offspring of the deer and goat is colored almost identically like her father but in horns and conformation she closely resembles her mother. This hybrid produced about two quarts of milk daily for nearly a year, feeding mostly on native weeds and vines.” Given that the locale of the mating was Cuba, the deer in question would have been O. virginianus, given that no other deer is native to that island. Farm and Fireside was a semi-monthly national agricultural magazine based in Springfield, Ohio. It was published between 1878 and 1939.
× Ovis aries (↔ usu. ♂) [Domestic Sheep] CHR. HPF(♀♀). CON: worldwide in association with humans. See the separate article “Goat × Sheep.”
× Pseudois nayaur [Bharal] Bunch et al. (1978) say hybrid fetuses have been produced, but that none survived birth. However, see Capra aegagrus × Pseudois nayaur.
× Rupicapra rupicapra (♂) [Chamois] ONHR. See the separate article “Domestic Goat × Chamois >>.” × Sus scrofa [Domestic Pig] There is a brief report of such a hybrid on page 3 of the June 25, 1867 issue of The Western Democrat, a newspaper published Charlotte, North Carolina (source). It reads as follows: "HALF HOG AND HALF GOAT—A Mr. Farmer, of Henderson county, informs us that he had born on his place an animal that was evidently half pig and half goat. The head and shoulders were hog, but the other part of the body was goat."
× Testudines [Turtle] See the separate article “Turtle-sheep Hybrids.”
× Ursus arctos [Brown Bear] See the separate article "Bear-goat Hybrids.

Capra ibex [Alpine Ibex | Steinbock]
See also: Ammotragus lervia × Capra hircus; Capra caucasica; C. hircus.
× Capra sibirica [Siberian Ibex] CHR. DRS. International Zoo Yearbook 1969 (p. 243).
× Ovis aries [Domestic Sheep] CHR?? Blood tests suggested that the offspring of an ibex dam served by a ram was a hybrid. Théret and Makowiak 1958; Valot-Gostynsky 1955.

Capra nubiana [Nubian Goat] See: Capra falconeri; C. hircus.

Capra sibirica [Siberian Ibex] See: Capra caucasica; C. hircus; C. ibex.

Gazella subgutturosa [Goitered Gazelle] See: Antilope cervicapra; Gazella gazella, G. leptoceros, G. marica.

Hemitragus jemlahicus [Himalayan Tahr | Jharal]
See: Capra hircus.
× Axis axis (♀) [Axis Deer | Chital | Indian Spotted Deer] CHR. See the separate article "Axis Deer × Himalayan Tahr."

Naemorhedus baileyi [Red Goral]
× Naemorhedus griseus [Chinese Goral] CHR. BRO: northeastern India, northern Myanmar. The Beijing Zoo had a hybrid in 2014. Internet:

Naemorhedus griseus [Chinese Goral] See: Naemorhedus baileyi

Ovibos moschatus [Muskox] Moscow Zoo had hybrids between two types (moschatus, wardi) treated as races of O. moschatus. International Zoo Yearbook 1990 (p. 487). See also: Bos taurus.

Oreamnos americanus [Mountain Goat]
× Antilocapra americana [Pronghorn] A notice about tertium quid, speculated to be a hybrid of this kind, appeared on page 2 of the December 29, 1873, issue of the Indianapolis News, a newspaper published in Indianapolis, Indiana (source). It reads as follows: “An animal of the deer species was shot in the woods and its carcass sold to a man in East Saginaw, Michigan, the like of which is unknown to any of the people thereabouts. The animal seemed to he a cross between a mountain sheep and an antelope. It was pure white with the exception of a few dark colored spots down its back, and the fore legs were some six inches shorter than the hind.” This cross, which does not seem to be reported elsewhere, requires verification. Note also that Michigan, at the present day, is well outside the ranges of both the proposed parents.

Ovis ammon [Argali]
× Ovis aries (♀) [Domestic Sheep] CHR. HPF(♂&♀). CON: western and central Asia. Rumjancev et al. (Animal Breeding Abstracts 1936, p. 192) artificially inseminated 212 domestic ewes. After a gestation period of 121-156 days, 43 F₁ lambs were obtained. Isenžulov says that 400 domestic ewes were inseminated with sperm from F₁ rams. About 400 offspring were obtained in the following year (1937). In 1938, ~500 more B₂ and later-generation hybrids were obtained. According to Isenžulov, “the vast majority of the hybrids combine the desired characters of both parental forms. When transferred to a high altitude (2,500-3,000 m), they throve and showed good increase in body weight and in weight and length of fleece.” Kushner and Kitaeva (1938a) found that the hybrids’ blood profile was superior to that of the domestic parent with respect to suitability for breeding at high altitudes. A breed, the Arkhar-Merino, was eventually produced from this cross. Butarin 1935, 1938; Ewart 1914; Fedosenko and Blank 2005 (p. 11); Isenžulov 1938; Isenžulov et al.1965a, 1965b; Izrael 1936; Lus 1938; Rumjancev et al. 1935; Steinmetz 1940. For many additional citations relating to this cross see Gray (1972).
× Ovis canadensis [Bighorn Sheep] CHR. DRS. Bunch and Workman 1988.
× Ovis gmelinii [Mouflon] CANHR. HPF(♂&♀). CON: Parvar Protected Region, northern Iran. Bunch et al. (2006, p. 29) found that a disproportionately high number of offspring with 2n = 54 resulted from inter se matings of F₁ hybrids (2n = 55). Pliny mentions this hybrid (Historia Naturalis, Book VIII, ch. 75). Bunch et al. 1976, 2006; Fedosenko and Blank 2005 (p. 11); Nadler et al. 1971b; Valdez et al. 1978 (p. 65).
× Ovis vignei (♂) [Urial] CANHR. HPF(♂&♀). Hiendleder et al. say the affinities of its mtDNA suggest that a type of sheep from the Transcaspian Ust-Urt Plateau (arkal) is derived from this cross, “due to overlapping ranges of urial and argali, like the hybrid zones of mouflon and urial in Iran” (See: Ovis gmelinii × O. vignei). The mtDNA haplotype found in the Ust-Urt sheep was similar to O. ammon, which is consistent with the reported direction of the cross in captive matings. The horns of the F₁ hybrids are rounded in front (but hollowed out behind as in O. vignei). Sterndale (1886a) says, “Now as regards the colour of the skin, the Nyan or Ovis hodgsoni [i.e., O. ammon] has no black beard or throat-stripe, which O. vignei has. The half-bred animal shows no black, but the quarter-breddoes in a modified but decided degree. The half-bred turns also in summer to the colour of O. hodgsoni, having more of a blue-grey or lavender tint and less of the fawn colour of O. vignei; with the white throat of O. hodgsoni, it also gets the dark patch at the side of the neck. The skin of a quarter-bred specimen before me is of a bright fawn above, sides and rump white, and a black stripe down the middle of the throat.” Sterndale (1886b, p. 36) argues that a sheep formerly treated as a species (O. brookei) was this hybrid. The population severtzovi in northeastern Uzbekistan has been variously treated as either as a species (Ovis severtzovi Nasonov, 1914) or as a subspecies of O. ammon or O. vignei, and is a probable derivative of this cross (see Geist). Ewart 1914; Fedosenko and Blank 2005 (p. 11); Geist 1991a; Hiendleder et al. 2002 (p. 901) ; International Zoo Yearbook 1990 (p. 489); Sarkisov 1953; Sterndale 1886a, 1886b; Ward 1924.

Ovis aries [Domestic Sheep] (2n=54)
See also: Ammotragus lervia; Bos taurus; Capra hircus; C. ibex; Ovis ammon.
× Antilocapra americana (♂) [Pronghorn] Two hybrids of this kind were reported as having been produced at Egon Canyon, Nevada, “crosses between common ewe sheep and the antelope.” The notice appeared on page 2 of the August 23, 1870, issue of The Carson Daily Appeal, a newspaper published in Carson City, Nevada (source). This cross, which does not seem to be reported elsewhere, requires verification.
× Canis familiaris [Domestic Dog] See the separate article Dog-sheep hybrids.
× Capreolus capreolus [Western Roe Deer] See the separate article “Roe Deer × Domestic Sheep.” This would be an interfamilial cross.
× Cervus elaphus [Red Deer | Elk | Wapiti] Such a hybrid is described in a quoted passage toward the end of the separate article, “Sheep/Mule Deer Hybrids.” In addition Bronn (1847, p. 167) lists this cross, but he cites no sources.
× Cervus nippon (♀) [Sika Deer] In Kunming, China, at the Yunnan Wild Animal Park, a ram and a sika doe formed a mated pair and have been many times photographed mating with each other. Neither of these animals was interested in sexual partners of its own kind. No hybrids have as yet been reported from this particular pairing, but see also the separate article "Roe Deer × Domestic Sheep." Internet Citations: SMACK.
× Equus asinus [Domestic Donkey] 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: "Hoc tempore in aula regís Galliarum аnimal nutrírí aiunt ab anteriore parte asini posteríore ovis specíe. Sed praeter ínstitutum est bígenera hîc omnia persequi." 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. This cross would be inter-ordinal. 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 “Jumarts.”
× Homo sapiens [Human] See the separate article “Caprinid-human hybrids.”
× Lepus europaeus [European Hare] See the separate article "Domestic Sheep × European Hare."
× Lontra canadensis [Northern River Otter] See the separate article "Northern River Otter × Domestic Sheep."
× Macropus sp. [Kangaroo] On the front page, column 7, of the November 2, 1849, issue of Geelong Advertiser, an Australian newspaper published in Geelong, Victoria (source), the following notice appears: “A few days ago a lusus naturae, in the shape of a lamb with a kangaroo’s head, might he seen on the [sheep] station of Mr. T. Malcolm. The likeness was exact, ears, eyes, head, and even the incisors, which in sheep are [only] in the under jaw, were, in this lamb’s, in strict resemblance of its prototype, in the upper. It did not live more than a few hours after being yeaned, as it could not suck. In every other respect it was like an ordinary lamb.” A related report >>
× Odocoileus hemionus (♂) [Mule Deer] See the separate article “Sheep × Mule Deer.”
× Odocoileus virginianus [White-tailed Deer] There is an anecdotal report of such a hybrid on page 3 of the Aug. 9, 1898 Semi-weekly Interior Journal, a newspaper published Stanford, Kentucky (access report). Another such report appears on page 2 of the June 01, 1902 The Times, a newspaper published Richmond, Virginia (access report).
× Ovis canadensis (♂) [Bighorn Sheep] CANHR. HPF(♀♀). CON: western U.S. There have been repeated reports of bighorn rams coming down from the mountains to hybridize with domestic ewes. Hybrids have light coats, often tinged tan, and bald faces. Their legs are longer, and tails, shorter than in O. aries. Hybrid ewes have produced lambs in their first season (bighorn ewes normally first lamb in their second). The pelage is a mixture of hair and wool. Anonymous 1935; Pulling 1945; Young and Manville 1960.
× Ovis gmelinii [Mouflon] CAENHR. Common in captivity. CON: southern Europe, Middle East. Bunch et al. 1976; Ibrovic et al. 1989; International Zoo Yearbook 1962 (p. 239), 1965 (p. 349), 1966 (p. 407), 1967 (p. 326), 1971 (p. 291), 1972 (p. 345), 1973 (p. 346), 1974 (p. 395); Karoian and Antonian 1988; Masala et al. 1991; Mavrogenis and Herzog 1990; Sterndale 1886b (p. 35); Türcke and Tomiczek 1982; Uloth 1976. Internet Citations: TAXO. See also: Der zoologische Garten (1873, p. 36)
× Ovis nivicola [Siberian Bighorn Sheep] CHR. In Russia, at Yakut State Agricultural Academy, breeders have produced this cross in efforts to create a new, highly cold-resistant breed of sheep. The hybrids are called ovchubuks.
× Pseudois nayaur [Bharal] Oregon Department of Fish and Wildlife’s rules governing nonnative wildlife ban this hybrid from Oregon. However, Gray (1972) says such hybrids have not been produced. Internet Citations: OREGR.
× Sus scrofa [Domestic Pig] See the separate article "Domestic Pig × Domestic Sheep."
× Tetrao urogallus [Western Capercaillie] See the separate article "Western Capercaillie × Domestic Sheep."

Ovis canadensis [Bighorn Sheep] (2n=54)
See also: Capra hircus, Ovis ammon; O. aries.
× Ovis dalli (♂) [Dall’s Sheep] CHR. ENHI(southern Yukon, northern British Columbia). HPF(♀♀). CON: mountains of western Canada. A female hybrid grew exceptionally well and surpassed its mother in size at 18 mo. She resembled a Fannin sheep, which are hybrids between O. dalli dalli and O. d. stonei (see note immediately preceding O. dalli). A male backcross to bighorn resembled typical bighorn lambs. Since a population (stonei) is morphologically (Cowan 1940; Shackelton 1985, p. 1) and geographically intermediate, it is a PHP of this cross. In stonei there is variation from grizzled white (like O. dalli) to brown (like O. canadensis), but is generally grayer than O. canadensis. Contact between O. canadensis and stonei is in northeastern British Columbia. Hoefs and Nowlan 1997. Internet Citations: DIN.
× Ovis gmelinii [Mouflon] CHR. DRS. HPF(♂&♀). Hybrids of both sexes have been reported. They are indistinguishable in appearance from O. canadensis, but like O. gmeliniithey are resistant to lung worm. Bunch et al. 1976; Gray 1972 (p. 143); Gray and Simpson 1980 (p. 4); Nadler et al. 1974; Shackelton 1985 (p. 6); Williams et al. 1983.

Ovis dalli [Dall’s Sheep] (2n=54)
See also: Ovis canadensis.
× Ovis stonei [Stone’s Sheep] CAENHR(northwestern Canada). HPF. Natural hybridization occurs in the mountains of the Yukon and Northwest territories. These sheep have usually been treated as a single species since Sheldon reported the existence of a hybrid zone in 1919. However, Stone’s sheep is distinct in appearance from Dall’s sheep. A darker animal it is silver to blackish gray, with white muzzles, leg trim, and rump patches, whereas Dall’s sheep is entirely off-white, with the exception of some individuals having black tails. Natural hybrids are known as Fannin sheep and were once treated as a separate species (O. fannini, Hornaday 1901). Guthrie 1972; Hoefs and Barichello 1984; Hoefs and Nowlan 1997; Scotter 1980; Sheldon 1919. Internet Citations: ARCT.

Ovis gmelinii [Mouflon] (= musimon; orientalis=gmelinii+vignei)
See also: Ovis ammon; O. aries; O. canadensis.
× Ovis vignei [Urial] CAENHR. HPF(vh). A 150-kilometer-wide hybrid zone exists in northern Iran in the Alborz Mountains, in the region directly south of the Caspian Sea (Nadler et al. 1971b) and in southeastern Iran (Valdez et al. 1978). Sheep in western Iran (Mouflon) have a diploid count of 2n = 54, whereas those in northeastern Iran (Urial) have 2n = 58. Within the hybrid zone 2n varies from 54 to 58. Diploid counts increase across the zone from east to west. The hybrids are variable, but in general, morphologically intermediate. A website (see Internet Citations: TAXO) says, “The red sheep (listed as Ovis o. orientalis on page 13 of the IUCN Caprinae Action Plan) is a hybrid form found in Iran [in the Alborz Mountain hybrid zone], the result of interbreeding of different mouflon and urial subspecies…Some delegates felt that the scientific name for urial should be Ovis orientalis, others that it should be O. vignei. The Caprinae Action Plan uses orientalis, but that name may have first been used to designate the red sheep, which is a hybrid.” This northern hybrid zone extends south into the Kavir Desert (Siah Kuh range). The hybrids of the southeastern hybrid zone, which lies in Kerman Province, east of 55˚E in the Khabr and Baft mountains, are known as Kerman sheep and are derived from the cross Ovis gmelinii laristanica × O. vignei blanfordi. Bunch et al. 1976; Hiendleder et al. 2002 (pp. 892-893); International Zoo Yearbook 1970 (p. 280), 1971 (p. 291), 1973 (p. 346), 1977 (p. 334), 1978 (p. 404); Nadler et al. 1971b; Valdez et al. 1978.

Ovis nivicola [Siberian Bighorn Sheep] See:Ovis aries.

Ovis stonei [Stone’s Sheep] See:Ovis dalli.

Ovis vignei [Urial] See: Ovis ammon; O. gmelinii.

Pseudois nayaur [Bharal] See: Capra aegagrus, C. hircus; Ovis aries.

Note: The IUCN (Internet Citations: RUPIC) says that “many of the less numerous subspecies [of R. rupicapra] (e.g., R. r. balcanica, R. r. cartusiana, and R. r. tatrica) are threatened by the deliberate introduction of subspecies from other geographic areas (especially R. r. rupicapra), leading to hybridisation and genetic swamping (Shackleton 1997).”

Rupicapra pyrenaica [Pyrenean Chamois]
× Rupicapra rupicapra [Chamois] Genetic evidence indicates interbreeding of these two types of chamois occurred prehistorically. Rodriguez et al. 2007.

Syncerus range map Syncerus range map

Syncerus brachyceros [Lake Chad Buffalo]
× Syncerus nanus [Forest Buffalo, Dwarf Buffalo] See: Syncerus caffer × S. nanus.

Syncerus caffer [African Buffalo]
See also: Bubalus bubalis.
×Equus burchellii [Burchell’s Zebra] 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.
× Syncerus nanus (♀?) [Forest Buffalo, Dwarf Buffalo] CAENHR(e Africa). HPF(♀♀). These buffaloes have been treated both as separate species, and as subspecies of Syncerus caffer. The African buffalo, also known as the cape buffalo, is a much larger animal than the forest buffalo, typically twice as heavy. It is black with broad spreading horns, while the other is red with backswept horns and much smaller ears. A third population, brachyceros, occupying the belt of savanna along the southern boundary of the Sahara, is composed of animals very similar to those of caffer (see map). So it is probably reasonable to lump brachyceros and caffer together as the "savanna type" in contradistinction to nanus, the "rainforest type." Christy (p. 458) says the hybrids occur along the forest margin. In the uplands of Kigezi and Ruanda there are distinctive buffaloes, known under the name mathewsi (=cottoni). Since mathewsi is morphologically and geographically intermediate between caffer and nanus, it is a PHP of this cross. It has been treated as a separate species, known as the Virunga buffalo (Syncerus mathewsi). Groves and Grubb (p. 119) mention that some phenotypic caffer individuals have nanus mtDNA haplotypes, which suggests that the direction of this cross is female nanus × male caffer. Grubb (1972) applied the name Syncerus brachyceros to animals that were really hybrids between brachyceros and nanus (and termed the populations here described under the name brachyceros as S. aequinoctialis). Ansell 1972; Buckland and Evans 1978; Christy 1929; Cribiu et al. 1980; Groves and Grubb 2011; Grubb 1972; International Zoo Yearbook 1967 (p. 322), 1971 (p. 286), 1972 (p. 340), 1973 (p. 341), 1975 (p. 387), 1981 (p. 330); Kingdon 1982 (pp. 58, 63); Van Hooft et al. 2002. Internet Citations: SANDI.

Syncerus mathewsi [Virunga Buffalo] See: Syncerus caffer × S. nanus.

Syncerus nanus [Forest Buffalo, Dwarf Buffalo] See: Syncerus brachyceros, S. caffer.

Taurotragus derbianus [Giant Eland]
× Tragelaphus eurycerus [Bongo] Parapatric contact zone (Cameroon to southern Sudan, possibly also in western Africa). No hybrids as yet reported. If this cross occurs, it would presumably be between Bongo male and Eland female, the size difference would otherwise be too great. Bongo embryos have been successfully transferred to surrogate eland mothers (Dresser 1984). Kingdon 1982 (maps, pp. 44, 123).

Taurotragus oryx [Eland] (2n = 31♂/32♀)
See also: Bos taurus.
× Tragelaphus strepsiceros (♀) [Greater Kudu] CHR. CON: eastern and southern sub-Saharan Africa. Hybrids of both sexes have been reported. A male F₁ hybrid reported by Jorge et al. had primarily eland phenotypic characteristics. It had a strong libido but produced no sperm. Examination of the testis showed complete lack of germ cells. The parental karyotypes differ by two reciprocal translocations and one pericentric inversion (involving chromosomes 1 and 3, 5 and 11, and 9 respectively). All other chromosomes have identical banding patterns. Boulineau 1933; International Zoo Yearbook 1981 (p. 330); Jorge et al. 1976; Ruhe 1993; Van Gelder 1977a.
× Tragelaphus spekei (♀) [Sitatunga] CHR. CON: eastern and southern Africa. Two female hybrids have been reported. Benirschke et al. 1980; International Zoo Yearbook 1975 (p. 386); van Gelder 1977b (p. 16).

Tragelaphus angasii [Nyala] (2n = 55♂/56♀)
× Tragelaphus strepsiceros (♂) [Greater Kudu] CHR. CON: southeastern Africa. A hybrid produced serendipitously looked like a nyala and had nyala mitochondrial DNA. But genetic analysis based on nine microsatellite markers, chromosome number and chromosome morphology confirmed it was an F1 hybrid. Dalton et al. 2014.

Tragelaphus eurycerus [Bongo] (2n = 33♂/34♀)
See also: Taurotragus derbianus
× Tragelaphus spekei (♀) [Sitatunga] CHR. HPF(♀♀). CON: central Africa. The parents differ markedly in appearance. An F₁ hybrid was similar to Bongo in bearing horns having a heavily built body, large ears, a mane on her back, and vertical striping on the sides, but resembled Sitatunga with respect to hooves and tail, and in having horizontal rows of white spots on the thighs and flanks. Gestation of an F₁ hybrid was 309 days. A backcross to T. spekei has been reported. Koulischer et al. (p. 265) say that meiosis in a female F₁ hybrid was “unimpaired.” International Zoo Yearbook 1966 (p. 402), 1967 (p. 321), 1968 (Plates 47 and 48); 1971 (p. 285); Kingdon 1982 (p. 83); Koulischer et al. 1973; Tijskens 1968.

Tragelaphus imberbis [Lesser Kudu] (2n = 38)
× Tragelaphus scriptus (♀) [Bushbuck] CHR. CON: Tanzania, Kenya, Ethiopia, Somalia. Kingdon 1982 (pp. 83, 113).
× Tragelaphus spekei (♀) [Sitatunga] CHR(Giardino Zoologico di Roma). CON: central Africa. Benirschke et al. 1980; International Zoo Yearbook 1969 (p. 239), 1974 (p. 390); Kingdon 1982 (pp. 83, 113).

Note: Two populations (scriptus, sylvaticus), treated as races of Tragelaphus scriptus, have an extensive hybrid zone in eastern and southern sub-Saharan Africa, within which individuals vary widely in in striping pattern, coat color, and horn morphology. HPF(vh). Hybrids have been treated as races (e.g., bor, ornatus. Kingdon 1982, p. 98).

Tragelaphus scriptus [Bushbuck] (2n = 33♂/34♀)
See: Tragelaphus imberbis.
× Tragelaphus spekei (♀) [Sitatunga] CHR. CON: The Sitatunga overlaps broadly with the Bushbuck in central and western sub-Saharan Africa (i.e., in the range of T. s. scriptus). Benirschke et al. 1980; Kingdon 1982 (p. 83).

Tragelaphus spekei [Sitatunga] (2n = 30)
See also: Tragelaphus eurycerus; T. imberbis; T. scriptus.
× Tragelaphus strepsiceros [Greater Kudu] CHR. CON: eastern Africa. International Zoo Yearbook 1986 (p. 500).

Tragelaphus strepsiceros [Greater Kudu] (2n = 31♂/32♀)
See also: Taurotragus oryx; Tragelaphus angasii; T. spekei.
+ Giraffa camelopardalis [Giraffe] Although no hybrids of this type seem to have been reported, a male greater kudu was observed attempting to mate with a female giraffe, as documented in this video.

deer-cow hybrid Deer-cow hybrids?

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By the same author: Handbook of Avian Hybrids of the World, Oxford University Press (2006).

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