Old World Monkey Hybrids

Family Cercopithecidae

cercopithecidae

EUGENE M. MCCARTHY, PHD GENETICS

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macaca nemestrina
Pig-tailed Macaque
(Macaca nemestrina)
Macaca mullata
Rhesus Macaque
(Macaca mullata)

Allenopithecus nigroviridis [Allen’s Swamp Monkey]
× Chlorocebus aethiops (♀) [Grivet Monkey] CHR(Naples, Italy). DRS. A viable male hybrid was reported. Chiarelli 1961.

Cercocebus sp.
× Macaca sylvanus (♂) [Barbary Macaque] CHR(Madison, Wisconsin, U.S.). DRS. A male hybrid was reported. Craft 1938 (Table 4).

Cercocebus agilis [Agile Mangabey]
× Lophocebus albigena (♀) [Grey-cheeked Mangabey] CHR. CON: northern Dem. Rep. Congo. Hill 1974 (p. 138).
× Cercocebus atys (♂) [Sooty Mangabey] CHR. DRS. Hill 1974 (p. 138); International Zoo Yearbook 1962 (p. 224).
× Cercocebus torquatus [Red-capped Mangabey] Parapatric contact zone in Cameroon (East Dja Reserve, southern Cameroon 3˚00´N, 13˚37´E). No hybrids as yet reported. Gartlan and Struhsaker 1972; Groves 1978 (p. 14).

Cercocebus atys [Sooty Mangabey]
See also: Cercocebus agilis.
× Cercocebus lunulatus (♀) [White-crowned Mangabey] CAONHR. Hill describes skins of probable natural hybrids obtained from Liberia. They were likely taken in nearby western part of Ivory Coast, where Booth (1958) says contact occurs in a probable hybrid zone (between the Nzo and Sassandra rivers). These taxa are not treated separately by Duff and Lawson (2004), but F₁ males are usually stillborn (as are some females). Monstrosities and stillbirths occur at high rates, too, in later-generation hybrids. See also Cercocebus atys × C. torquatus. Groves (1978, pp. 11-12) mentions a probable hybrid "in Paris" from Guiglo, Ivory Coast. Hill 1974 (pp. 137, 138, 146-149), Zuckerman 1953 (p. 942).
× Cercocebus torquatus (usu. ♀) [Red-capped Mangabey] CHR. ENHI. DRS. HPF(♂&♀). Some hybrids are inviable. However, at least one lived 20 years. Another, born at the London Zoo in 1932, had blue eyes that led Zuckerman (1933, p. 97) to comment that "all other mangabeys I have seen, in fact almost all monkeys and apes, have unmistakable brown eyes." The father was albino. A hybrid from the reciprocal cross lived two days. Groves (1978, p. 11) says lunulatus is morphologically "intermediate in many respects between atys and torquatus." It is also geographically intermediate and, thus, a PHP of this cross. Ashley Montagu 1950; Hill 1974 (p. 138); International Zoo Yearbook 1971 (p. 265), 1972 (p. 318), 1973 (p. 322); Zuckerman 1933, 1953 (pp. 830, 942).
× Macaca fascicularis (♀) [Long-tailed Macaque] CHR. DRS. A hybrid listed by Ashley Montagu (1950) lived three months. Sclater reported one from the reciprocal cross that died in utero when the mother died near term in a fall. The macaque sire of the latter hybrid also mated with a Mandrill (Mandrillus sphinx) which later birthed a viable hybrid. Sclater says, "there seems no reason to doubt that the Macaque was the father of both the young ones, there being no male Mandrill nor Mangabey which could have had access to them, and the Mandrill’s young one having a short tail." The Proceedings of Philadelphia Academy of Natural Sciences (1890, p. 446), too, list a specimen in the collection of the academy's museum, of a hybrid between "Macacus cynomolgus and Cercocebus fuliginosus," which would also be a hybrid of this sort, or something very like it. Hill 1974 (p. 139); Sclater 1878 (p. 791); Zuckerman 1933, 1953 (p. 942).
× Macaca nemestrina (♀) [Pig-tailed Macaque] CHR(Philadelphia). DRS. A hybrid lived almost two years. Ashley Montagu 1950.
× Mandrillus sphinx (♀) [Mandrill] CHR. DRS. The Brookfield Zoo, Chicago, Illinois (USA) has a hybrid. Internet Citations: AMIEE, VGFER.

Golden-bellied Mangabey
Golden-bellied Mangabey (Cercocebus chrysogaster) Image: Wikimedia
Olive Baboon
Olive Baboon
(Papio anubis)

Cercocebus chrysogaster [Golden-bellied Mangabey]
× Papio anubis (♀) [Olive Baboon] CHR. CON: northwestern Dem. Rep. Congo? A hybrid was born at the Mount Kenya Animal Orphanage in 2003. Ngugi 2003.

Cercocebus galeritus [Tana River Mangabey]
× Lophocebus albigena (♀?) [Grey-cheeked Mangabey] CHR. DRS. International Zoo Yearbook 1962 (p. 224).

Cercocebus lunulatus [White-crowned Mangabey]
See also: Cercocebus atys; Cercocebus atys × C. torquatus.
× Cercopithecus mitis (♂) [Blue Monkey] CHR(Philadelphia). DRS. Two hybrids died soon after birth. Concerning taxonomic treatment of C. lunulatus, see Cercocebus atys × C. lunulatus. Ashley Montagu 1950; Hill 1974 (p. 139).
× Cercocebus torquatus [Red-capped Mangabey] CHR(Wassenaar, Netherlands). International Zoo Yearbook 1970 (p. 254), 1971 (p. 265).

Cercocebus torquatus [Red-capped Mangabey]
See also: Cercocebus lunulatus.
× Cercopithecus mitis (♂) [Blue Monkey] CHR. DRS. A hybrid died soon after birth. Ashley Montagu 1950.
× Macaca fascicularis (↔) [Long-tailed Macaque] CHR. DRS. Chiarelli 1961.
× Macaca mulatta [Rhesus Macaque] CHR. DRS. Baker 1988 (p. 53). Internet Citations: ZOOH.
× Macaca nemestrina [Pig-tailed Macaque] CHR. DRS. Chiarelli 1961.
× Mandrillus leucophaeus (♀) [Drill] CHR(Edinburgh). Chiarelli 1961, 1967; Hill 1970 (p. 483).
× Mandrillus sphinx (♀) [Mandrill] CHR(Cincinnati). CON: Rio Muni, Gabon. A hybrid was colored like a Mandrill, but had a black head, a black stripe down its spine, and a long black tail. It died of an illness at the age of four months. Martin et al. 1974; Zipperlen 1896.

Note: Two populations (kolbi, monoïdes), treated as races of Cercopithecus albogularis, are connected by a geographically and morphologically intermediate population (kibonotensis) in southern Kenya and northern Tanzania. Meester and Setzer (1971, Part 3, p. 20) note "Altogether, kibonotensis appears to be transitional between kolbi, with a broad white collar, and monoïdes with a white throat."

Cercopithecus albogularis [Sykes’ Monkey]
× Cercopithecus mitis (↔) [Blue Monkey] CAENHR(northeastern Tanzania). HPF(vh). There is a hybrid zone near Karatu. Hybrids are easily produced in captivity. Booth 1968; International Zoo Yearbook 1962 (p. 225), 1974 (p. 372), 1983 (p. 316).
× Cercopithecus mona (♂) [Mona Monkey] CHR. DRS. Gijzen 1963; Hill 1966 (p. 381, citing Gijzen).
× Chlorocebus pygerythrus (♀) [Vervet Monkey] CHR. In the abstract for a presentation given at a conference, Bramblett (1992) says that a shortage of space at the Department of Anthropology, University of Texas, Austin resulted in the introduction of an adult male Sykes’ monkey into a vervet group composed of adult vervets of both genders. "Sexual facilitation," Bramlett goes on to say, "seemed to make him a popular consort among vervet females. Several hybrid conceptions occurred and two live hybrid births followed in the spring of 1985. Shortly afterwards a mixed species group (half vervet and half Sykes’ monkeys) was established. The hybrids and their mothers were incorporated into this mixed group so they had social companions (adults and playmates) of both species to serve as role models. One of these hybrids (DU, a male, born June 9, 1985) is described as he reaches adulthood. Behaviorally and socially he is a vervet and is imbedded into his mother’s social group. Physically he becomes more Sykes’-like as he matures. He is much larger than the other vervets and there would be little reason to consider a vervet contribution to his phenotype if his history were not known."

Cercopithecus ascanius [Red-tailed Monkey]
× Cercopithecus doggetti [Silver Monkey] NHR(northwestern Tanzania). De Jong and Butynski 2010; Detwiler 2002, 2005.
× Cercopithecus mitis (prob. ♀) [Blue Monkey] ONHR(southwestern Uganda, southwestern Kenya). HPF(♀♀). These monkeys differ markedly in appearance. Hybridization occurs in the Budongo, Itwara, and Kibale forests. Aldrich-Blake (1968) says the two use very similar postures to solicit mating. He describes a wild female hybrid. It differed from C. mitis in having a white nose spot and in lacking black on her arms, and from a red-tail in the color and pattern of her cheek fur, the absence of brown or red on the back and tail, and in the underside being pale gray instead of creamy white. She carried a hybrid baby that could have passed for a blue if not for its white nose spot. Its appearance together with the fact that the mother lived with a troop of blues suggest it was a backcross to C. mitis. That the mother hybrid lived with blues suggests her own mother was a blue (since as a general rule, hybrid mammals remain with the mother after birth and become part of her social group). Coimbra-Filho et al. 1993 (p. 117) say hybrids have been reported from at least four localities and that hybrid may be at an advantage in the Butinsky 1982 (p. 7); De Jong and Butynski 2010; Oates 1985 (p. 41); Struhsaker et al. 1988.
× Cercopithecus pogonias (♂) [Crowned Monkey] NHR. CON. central Africa (n of the Congo). Hybrids of both sexes are reported. Gautier and Gautier say calls of a one-month old female hybrid contained attributes of both parental calls. Dugoujon et al. 1982b; Gautier and Gautier 1977; Gautier and Gautier-Hion 1982; International Zoo Yearbook 1980 (p. 424); Martin et al. 1974; Muleris et al. 1985. Internet Citations: DPZ.

× Erythrocebus cephus [Patas Monkey] ENHI(southeastern Central African Republic, between the Sangha and Ubangi). Grubb et al. (2003, p. 1336) say a population (ngottoensis) is "geographically and morphologically intermediate between C. cephus and C. ascanius." It is therefore a PHP of this cross. Colyn 1999.

Cercopithecus petaurista [Lesser Spot-nosed Monkey] See: Cercopithecus cephus.

Cercopithecus cephus [Moustached Monkey]
See also: Cercopithecus ascanius.
× Cercopithecus erythrogaster [Red-bellied Monkey] ENHI(se Nigeria, w Cameroon). Meester and Setzer (1971, Part 3, p. 23) say that Cercopithecus erythrotis, which is geographically intermediate between these monkeys "combines the characters found in C. erythrogaster, on the one hand (black nape and under parts ash grey), and in C. cephus, on the other (pale fringe on the ears, russet tail)." In addition C. erythrotis, is known to hybridize extensively with C. cephus in southwestern Cameroon. Taken together, these facts suggest C. erythrotis as a PHP of this cross. See Cercopithecus cephus × C. erythrotis.
× Cercopithecus erythrotis [Cameroon Red-eared Monkey] ENHR(southwestern Cameroon). HPF(vh). These hybrids are common south of the Sanaga River below Edéa. Facial coloration is variably intermediate. Hill (1966) placed these monkeys in different superspecies. See Cercopithecus cephus × C. erythrogaster. Struhsaker 1970 (pp. 374-375).
× Cercopithecus mitis [Blue Monkey] Oates (1985, p. 41) suggests that Cercopithecus signatus (Greater White-nosed Monkey), based on three skins of uncertain provenance, may be this hybrid.
× Cercopithecus mona (♂) [Mona Monkey] CHR. CON: southwestern Cameroon, Gabon. Hill 1966 (p. 381); Gray 1972; Martin et al. 1974.
× Cercopithecus nictitans [Greater Spot-nosed Monkey] NHR(Lopé Reserve, Gabon). A single natural hybrid has been reported. De Jong and Butynski 2010; Tutin 1999.
× Cercopithecus petaurista [Lesser Spot-nosed Monkey] CHR. DRS. A male hybrid has been reported. International Zoo Yearbook 1978 (p. 376).
× Erythrocebus patas (♂) [Patas Monkey] CHR(St. Paul Zoo, Minnesota, USA). DRS. International Zoo Yearbook 1960 (p. 256). Internet Citations: SD167.

Cercopithecus denti [Dent’s Monkey]
× Cercopithecus doggetti [Silver Monkey] NHR(Nyungwe National Park, southwestern Rwanda). B. A. Kaplin pers. comm. in Detwiler et al. 2005.

Cercopithecus diana [Diana Monkey]
× Canis familiaris [Domestic Dog] See the separate article "Domestic Dog × Primate"
× Cercopithecus roloway [Roloway Monkey] CHR(Duisburg Zoo, Germany). CON: Ghana, Ivory Coast. These two monkeys have been treated, at times, as conspecific. International Zoo Yearbook 1970 (p. 256).

Cercopithecus doggetti [Silver Monkey]
See also: Cercopithecus ascanius; Cercopithecus denti.
× Cercopithecus kandti [Golden Monkey] ENHR(southwestern Uganda). HPF(vh). Kingdon (p. 240) says the taxon schoutedeni "would seem to be described from animals that are intermediate in almost every particular, and which show every shade of gradation between C. m. kandti and S. m. doggetti. "These taxa are sometimes lumped. Kingdon 1971 (pp. 235, 240).
× Cercopithecus lhoesti [L’Hoest’s Monkey] Kingdon (1971, pp. 244-245) says these monkeys occur in mixed troops in the Kayonza Forest, but he does not mention hybrids.
× Cercopithecus mona (♂) [Mona Monkey] CHR. DRS. Gijzen (1963) describes in detail a male hybrid born at Antwerp Zoo. International Zoo Yearbook 1965 (p. 334).

Cercopithecus erythrogaster [Red-bellied Monkey]
See also: Cercopithecus erythrotis.
× Cercopithecus mona (♀) [Mona Monkey] CHR. CON: southwestern Nigeria. A female hybrid has been reported. International Zoo Yearbook 1971 (p. 267).
× Cercopithecus sclateri [Sclater’s Monkey] NHR(Niger Delta, Nigeria). Two natural hybrids were reported. De Jong and Butynski 2010 (p. 2).

Cercopithecus erythrotis [Cameroon Red-eared Monkey]
See: Cercopithecus cephus.
× Cercopithecus nictitans [Greater Spot-nosed Monkey] NHR. CON: Bioko Island, Equatorial Guinea. A hybrid is pictured in The Proceedings of the Zoological Society of London (1907, Issue 3, Plate XL).

Cercopithecus hamlyni [Owl-faced Monkey]
× Cercopithecus lhoesti (♀) [L’Hoest’s Monkey] CHR. CON: eastern Africa (between Lake Victoria and the Lualaba). Viable female hybrids are known. Chiarelli 1961; Martin et al. 1974.
× Cercopithecus mona (♀) [Mona Monkey] CHR. DRS. Chiarelli 1961.
× Cercopithecus mitis [Blue Monkey] CHR. CON: northeastern Dem. Republic of Congo. A male hybrid was reported. International Zoo Yearbook 1987 (p. 406).

Cercopithecus kandti [Golden Monkey] See: Cercopithecus doggetti.

Cercopithecus lhoesti [L’Hoest’s Monkey] See: Cercopithecus doggetti; C. hamlyni.

Note: Two populations (kolbi, stuhlmanni), treated as races of Cercopithecus mitis, hybridize along the Eastern Rift Valley (southwestern Kenya). Kingdon says a type, boutourlinii, native to Ethiopia (between Lake Tana and Lake Turkana) is similar to kolbi × stuhlmanni hybrids, and suggests that boutourlinii may be descended from interbreeding kolbi and stuhlmanni colonists. Hill (1966, p. 401) also says boutourlinii is "perfectly intermediate" and says albotorquatus (southern Somalia, around Baldoa) is too. All of these taxa, with the exception of stuhlmanni, were once treated as separate species. Hill (1966) treated kolbi as a race of C. albogularis. Kingdon 1971 (pp. 235, 239-240).

Cercopithecus mitis [Blue Monkey]
See also: Cercopithecus albogularis; C. ascanius; C. hamlyni; C. lhoesti; C. lunulatus; C. torquatus.
× Chlorocebus pygerythrus (♂) [Vervet Monkey] NHR. A probable male hybrid was found at Diani on the south coast of Kenya (about 30 miles south of Mombasa) in December 2008, and two additional probable hybrids have since been located, one at Diani, and one at Ngong Forest Sanctuary, Nairobi. De Jong and Butynski note that "young juvenile C. pygerythrus sometimes become well-integrated into C. mitis groups and probably grow up in them." The same authors provide tables comparing many of the hybrids’ traits to those of their parents. They also describe (p. 2) the first hybrid as "intermediate to the parent species in most respects, although the color of the face, neck collar, dorsum, back of the legs, sides, and tail appears to be slightly more like C. mitis, while body shape, and color of the iris, eyelids, shoulders, and ventrum appear to be slightly more like C. pygerythrus. The scrotum of the Diani hybrid is intermediate in size and color between C. mitis and C. pygerythrus. The muzzle of the Diani hybrid seems to be longer and more pointed than the muzzle of either C. mitis or C. pygerythrus. Interestingly, unlike either parent species, the Diani hybrid has a faint nose spot (recalling C. nictitans and members of the ‘Cercopithecus cephus species-group’)." They further note (p. 11) that the "adult male hybrid at Diani exhibits some phenotypic characters (for example, pale grey nose spot) and behaviors (for example, active scent-marking), which are absent or rare in both parent species." De Jong and Butynski 2010. Internet Citations: BUTYN
× Chlorocebus tantalus [Tantalus Monkey] CHR? CON: central Africa. De Jong and Butynski (2010, p. 9) state that "Erhart et al. (2005) report on two captive-bred C. mitis × C. pygerythrus hybrids, but these are probably C. mitis × C. tantalus hybrids (T. Rowell pers. comm.)."

Cercopithecus mona [Mona Monkey]
See also: Cercopithecus albogularis; C. doggetti; C. cephus; C. hamlyni.
× Cercopithecus neglectus (♀) [De Brazza’s Monkey] CHR. CON: eastern Dem. Rep. Congo (east of the Lualaba). Chiarelli 1961.
× Cercopithecus nictitans [Greater Spot-nosed Monkey] CHR. CON: Cameroon. Gyekye 1966; International Zoo Yearbook 1967 (p. 306); Martin et al. 1974.
× Cercopithecus pogonias (♂) [Crowned Monkey] ONHR(western Cameroon). Struhsaker (1970, pp. 372-373) describes three hybrids observed near Idenau (4°14´N, 8°59´E) and Tinaso (3°37´N, 9°57´S). Dugoujon et al. 1982a; Martin et al. 1974. Internet Citations: DPZ.
× Chlorocebus aethiops [Grivet Monkey] CHR. DRS. In Germany, both the Braunschweig and the Straubing zoos have had hybrids. Benzien 1959; International Zoo Yearbook 1968 (p. 294).

Cercopithecus neglectus [De Brazza’s Monkey] See: C. hamlyni.

Cercopithecus nictitans [Greater Spot-nosed Monkey]
See also: Cercopithecus cephus; C. erythrotis; C. mona.
× Cercopithecus petaurista [Lesser Spot-nosed Monkey] CHR(Duisberg Zoo, Germany). CON: northern Liberia. International Zoo Yearbook 1970 (p. 255).

Cercopithecus petaurista [Lesser Spot-nosed Monkey] See: C. nictitans.

Cercopithecus pogonias [Crowned Monkey] See: C. ascanius; C. mona.

Cercopithecus roloway [Roloway Monkey] See: C. diana.

Cercopithecus sclateri [Sclater’s Monkey] See: C. erythrogaster.

Cercopithecus signatus [Greater White-nosed Monkey]See: Cercopithecus cephus × C. mitis.

Chlorocebus aethiops [Grivet Monkey]
See also: Allenopithecus nigroviridis; Cercopithecus mitis; C. mona.
× Chlorocebus pygerythrus [Vervet Monkey] CAENHR(e Africa). Parapatric contact zone (southwestern Ethiopia). There is a hybrid population in Gama-Gofa Province, Ethiopia (w of Lake Abaya). Dandelot 1959; Dandelot and Prevost (1972, p. 615); International Zoo Yearbook 1978 (p. 375), 1987 (p. 406); 31st Report of the Zoological Society of Philadelphia 1903 (p. 23).
× Chlorocebus sabaeus (♂) [Green Monkey] CHR(Erfurt, Germany). DRS. International Zoo Yearbook 1969 (p. 222).
× Erythrocebus patas [Patas Monkey] CHR. CON: eastern Sudan, western Ethiopia. International Zoo Yearbook 1974 (p. 372), 1978 (p. 354), 1980 (p. 424).
× Macaca mulatta (♀) [Rhesus Macaque] CHR. DRS. This hybrid occurred at the Zoological Gardens of London in 1873. Zuckerman 1953 (p. 942).
× Macaca radiata (♂) [Bonnet Macaque] CHR. DRS. Gunning 1910; Zuckerman 1953.
× Macaca sinica [Toque Macaque] CHR. DRS. Gunning 1910; Przibram 1910.
× Papio anubis [Olive Baboon] Aldrich-Blake (1968, p. 19) mentions a case of a male Grivet living with a troop of Olive Baboons and mating with the females of the troop, but no hybrids were reported.

Note: Grubb et al. (2003, p. 1331) say "Napier (1981) reported intergradation in a broad tract extending through Dandelot’s [i.e., Dandelot 1959] hybrid zone [i.e., Chlorocebus pygerythrus × C. tantalus] northeastward to Harar in Ethiopia. Apparently the zone involves Cercopithecus djamdjamensis, C. aethiops hilgerti and C. pygerythrus nesiotes in addition to C. tantalus budgetti and C. pygerythrus centralis."

Chlorocebus cynosuros [Malbrouck Monkey]
× Chlorocebus pygerythrus [Vervet Monkey] Parapatric contact zone (s. Africa). No hybrids as yet reported. These taxa are often lumped. Dandelot 1959 (Fig. 6).
× Chlorocebus tantalus [Tantalus Monkey] Parapatric contact zone (Dem. Rep. Congo). No hybrids as yet reported. These taxa are often lumped. Dandelot 1959 (Fig. 6).

Chlorocebus pygerythrus [Vervet Monkey]
See also: Cercopithecus albogularis; C. mitis; Chlorocebusaethiops; C. cynosuros.
× Chlorocebus sabaeus (♀) [Green Monkey] CHR. DRS. International Zoo Yearbook 1981 (p. 312), 1982 (p. 414), 1983 (p. 316); 34th Report of the Zoological Society of Philadelphia 1906 (p. 10).
× Chlorocebus tantalus [Tantalus Monkey] ENHR(e Africa). HPF(vh). Hybrid zone is in southwestern Uganda and northeastern Dem. Rep. Congo (upper White Nile Valley, near Lake Edward). Dandelot 1959; Kingdon 1971.
× Erythrocebus patas (♂) [Patas Monkey] CHR. CON: northwestern Kenya, northern Uganda and southern Sudan. A female hybrid was born in captivity in 1975. Matsubayashi et al. 1978.
× Lophocebus albigena [Grey-cheeked Mangabey] Hill (1974, p. 190) says an L. albigena male had a C. pygerythrus female as mate, but no hybrids are reported.
× Macaca fascicularis [Long-tailed Macaque] Citing Gunning (1910) and Przibram (1910), Hill (1966, p. 379) lists this cross. However, Gunning reports (and Przibram lists) only C. lalandei (i.e., C. pygerythrus) × Macacus sinica.
× Macaca radiata [Bonnet Macaque] Citing Gunning (1910), Hill (1974, Table 33) lists this cross. However, Gunning reports only C. lalandei (i.e., C. pygerythrus) × Macacus sinica.
× Macaca sinica (♂) [Toque Macaque] CHR(Pretoria, S. Africa). DRS. Gunning (1910) reports a viable male hybrid. He says it was "gesund und außerordentlich munter" (i.e., healthy and extraordinarily lively). Like its sire, it had an unpigmented face.

Chlorocebus sabaeus [Green Monkey]
See also: Chlorocebus aethiops; C. pygerythrus.
× Chlorocebus tantalus [Tantalus Monkey] CHR. Parapatric contact zone (upper Volta River Valley). International Zoo Yearbook 1977 (p. 305).
× Erythrocebus patas [Patas Monkey] NHR(Saloum Delta National Park, Senegal). A single natural hybrid has been reported. De Jong and Butynski 2010; Galat-Luong 1996; Galat et al. 1993.
× Macaca mulatta (♀) [Rhesus Macaque] CHR? DRS. The Zoological Society of London’s occurrence sheet for March 22, 1873 notes the birth of a hybrid at the London Zoo "between Cercopithecus callitrichus M[ale]. and Macacus erythraeus F[emale]." Zuckerman 1931 (p. 338), 1933 (p. 96), 1953 (pp. 833, 942).

Chlorocebus tantalus [Tantalus Monkey] See: Cercopithecus mitis, Chlorocebuscynosuros; C. pygerythrus; C. sabaeus.

Colobus angolensis [Angola Pied Colobus] (= palliatus)
× Colobus polykomos [Western Pied Colobus] CHR. DRS. Dutrillaux et al. 1981.
× Colobus ruwenzorii [Rwenzori Black-and-white Colobus] ENHR. Hybrid zones exist in the Uele-Oubangui river system and near Lake Kivu (Internet Citations: ANGRU). Although angolensis and ruwenzorii were long treated as separate species, they have been lumped in recent taxonomy.

Colobus guereza [Guereza Colobus]
× Colobus polykomos [Western Pied Colobus] CHR. DRS. HPF. International Zoo Yearbook 1988 (p. 457); Lang 1973.

Colobus polykomos [Western Pied Colobus]
See also: Colobus angolensis; C. guereza.
× Colobus vellerosus [Geoffroy’s Pied Colobus] ENHR(Ivory Coast). These taxa are sometimes lumped. A population, dollmani, which has been treated as a subspecies of both of C. polykomos and of C. vellerosus, is actually composed of hybrids of this type (Groves et al. 1993). Studies show that dolmani is morphologically and geographically intermediate between polykomos and vellerosus, as well as far more variable than either parent. However, the construction of a dam has now resulted in the inundation of most of the region of contact between the Sassandra and Bandama rivers, where hybridization formerly occurred. Groves et al. 1993; Grubb et al. 2003 (p. 1343); Oates and Trocco 1983; Nowak 1999 (vol. 1, p. 604).

Erythrocebus patas [Patas Monkey] See: Cercopithecus cephus; Chlorocebus aethiops; C. pygerythrus; C. sabaeus.

Lophocebus sp. [Mangabey]
× Papio cynocephalus (♂) [Yellow Baboon] ENHR. In 2005, a new taxon Lophocebus kipunji was described, based on a single specimen and observations of living animals in relict forests in Tanzania. Its initial assignment to Lophocebus was based on its overall morphology, but subsequent genetic studies indicating it was sister taxon to common baboons (Papio) led to its generic separation, as Rungwecebus. However, Burrell et al. (2009) present a strong case that it actually originated by hybridization between a mangabey of some time (Lophocebus sp.) and a yellow baboon (Papio cynocephalus) females and state that "We believe this to be the first case among mammals in which a natural occurrence of inter-generic hybridization can be shown to have resulted in a new, distinct, long-surviving taxon." Burrell et al. 2009.

Note: Two populations (albigena, osmani), treated as subspecies of Lophocebus albigena, have a narrow hybrid zone in central Cameroon (Groves 1978). A hybrid between albigena and a third type (johnstoni), also treated as a race of L. albigena, has also been reported (International Zoo Yearbook 1989, p. 317).

Lophocebus albigena [Grey-cheeked Mangabey]
See also: Cercocebus agilis; C. galerita; Chlorocebus pygerythrus.
× Lophocebus aterrimus (♂) [Black Mangabey] CHR. DRS (ranges are separated by Congo River). Hybrids have the bushy tail of L. albigena, but otherwise resemble L. aterrimus. These taxa were recently split. Hill 1974 (p. 139); International Zoo Yearbook 1967 (p. 305), 1968.

Lophocebus aterrimus [Black Mangabey]
See also: Lophocebusalbigena.
× Lophocebus obdenboschi (♀)[Opdenbosch’s Mangabey] CHR. CON: southeastern Dem. Rep. Congo. Hill (1974, p. 190) mentions three hybrids. They resembled L. aterrimus, but their whiskers were not wholly black. These taxa are often lumped.

Lophocebus kipunji [Kipunji] See: Lophocebus sp. × Papio cynocephalus

Lophocebus obdenboschi [Opdenbosch’s Mangabey] See: Lophocebus aterrimus.

Note: In a report on hybridization in macaques, Bernstein and Gordon (1980, p. 145) say that "hybrids reported in this study, and in the literature, demonstrate that many species with the genus Macaca are capable of fertile cross-matings and that the offspring of such mating are no less fertile than the parental generation."

Macaca arctoides [Stump-tailed Macaque]
See also: Macaca assamensis × M. fascicularis.
× Macaca assamensis (♀) [Assamese Macaque] CHR(Berlin). CON: northern Myanmar. An Assamese female produced hybrids in three successive gestations. All survived. International Zoo Yearbook 1969 (p. 220).
× Macaca fuscata [Japanese Macaque] CHR. DRS. Ruhr-Zoo Gelsenkirchen had two hybrids. Internet Citaions: DPZ.
× Macaca mulatta (♀) [Rhesus Macaque] CHR(Vienna). ENHI? Several captive hybrids are reported. Some markers suggest these macaques are more alike genetically where they are in contact (Thailand), which suggests extensive gene flow. Antonius 1951b; Cronin et al. 1980 (p. 42); Eudey 1980 (p. 81).

Macaca assamensis [Assam Macaque]
See also: Macaca arctoides.
× Macaca fascicularis (♀) [Long-tailed Macaque] ENHR(southeast Asia). HPF. Brandon-Jones et al. 2004 (p. 112) say "Macaca arctoides was first published in abstract by Isidore Geoffroy Saint-Hilaire (1830). Genetic evidence from mtDNA and Y-chromosome DNA suggest that M. arctoides originated from hybridization of M. assamensis/thibetana-like males with M. fascicularis-like females (Tosi et al. 2000, 2003)." However, according to distribution maps in Fooden (1980), M. thibetana does not come into direct contact with M. fascicularis, so hybridization must have been with M. assamensis (unless distributions were much different at the time of hybridization).
× Macaca nemestrina (♀) [Pig-tailed Macaque] CHR. CON: Burma, northeastern Bangladesh, Assam. Bernstein and Gordon (1980, Table 6-1) report a stillborn hybrid.
× Macaca radiata (♂) [Bonnet Macaque] CHR. DRS (possible former contact). Viable hybrids of both sexes have been reported. The body pelage of a hybrid resembles M. assamensis, but ear shape, tail length and shape, and crown hair resemble M. radiata. Acharjyo and Mishra 1976, 1982.
× Macaca thibetana [Père David’s Macaque] Parapatric contact zone (southern China). No hybrids as yet reported.

Macaca cyclopis [Taiwan Macaque]
× Macaca fuscata [Japanese Macaque] CAONHR. HPF. DRS. Extensive crossbreeding of the introduced M. cyclopis with the native M. fuscata in Wakayama Prefecture was genetically confirmed. The proportion of M. cyclopis genes was 57.4% (Kawamoto 2005). Also, numerous 3-way hybrids (listed as "M. fuscata × M. cyclopis × M. fascicularis") were bred in Oshima, Japan (International Zoo Yearbook 1963; 1972, p. 317; 1973, p. 322).

Macaca fascicularis [Long-tailed Macaque] (= irus)
See also: Cercocebus atys; Macaca assamensis; Macaca cyclopis × M. fuscata.
× Macaca maura (♀) [Moor Macaque] CHR. DRS. Bernstein and Gordon (1980, Table 6-1).
× Macaca mulatta (↔ usu. ♀) [Rhesus Macaque] CAENHR. CON: southeast Asia. HPF(♂&♀). The hybrid zone is about 2,000 km long. It extends from southeastern Bangladesh, southward through Myanmar to about ~17˚N, whence it turns due west through Thailand; at the Laos border it turns southeast, running through eastern Cambodia and southern Vietnam (see map in Fooden 1997). Fooden (1997, p. 228) evaluated macaques in the contact zone using relative tail length (RTL) as a measure (i.e., the ratio of tail length to head and body length. He comments that "in most parts of the extensive ranges of M. fascicularis and M. mulatta, RTL is strongly differentiated. North of ca. 5˚N, RTL in M. fascicularis tends to decrease latitudinally and therefore tends to approach that in M. mulatta." He also notes that the transition from long-tailed M. fascicularis to short-tailed M. mulatta is geographically abrupt (a graph of RTL across the contact zone shifts discontinuously). This finding probably results from the relative inviability of hybrids. Indeed, captive crosses indicate that some hybrids are inviable (although others reach maturity). The hybrids are greyish in color. A tuft is present (as in M. fascicularis) in young hybrids, but later disappears. An F₁ female had an unpigmented face, as in M. mulatta, but a long tail (though not so long as in M. fascicularis). However, Fooden (1997) says the pelage of natural hybrids is variably intermediate (which suggests the occurrence of extensive backcrossing). Genetic studies based on Y-chromosome DNA carried out by Tosi et al. confirm the occurrence of natural hybridization. They say (p. 172) that monkeys from as far into the territory of M. fascicularis as Tum Chompol, Thailand (~200 km from the contact zone) "share exactly the same SRY and TSPY sequences with M. mulatta from China and Burma." They suggest that the subpopulation of M. fascicularis where M. mulatta Y-chromosome DNA (north of the Isthmus of Kra) "may require a subspecific designation distinct from southern conspecifics." They also suggest that mtDNA has not moved across the contact zone in either direction because of the extreme female philopatry characteristic of these macaques. These macaques have hybridized extensively, too, in Hong Kong (on the Kowloon Peninsula) since the introduction of long-tails in the 1950s. Hybrids now make up ~30% of the population there (long-tails comprise only 2%). Anderson 1881 (p. 65); Ashley Montagu 1950; Avise and Duvall 1977; Bernstein and Gordon (1980, Table 6-1); Burton and Chan 1996; Eudey 1980; Flower 1929a (p. 23); Fooden 1964, 1971, 1997; Groves 2001; Hamilton 1914; Hill 1972, 1974 (Table 33 and pp. 474, 504-506, 542); Honacki et al 1982 (p. 235); International Zoo Yearbook 1959, 1960, 1961, 1965, 1966, 1969; Melnick and Kidd 1985 (p. 150); Niemeyer 1868; Southwick and Southwick 1983; Tosi et al. 2000, 2002; von Knottnerus-Meyer 1904; Wong and Ni 2000; Zuckermann 1953 (p. 942).
× Macaca nemestrina (usu. ♀) [Pig-tailed Macaque] CAONHR(Malay Peninsula). HPF(♂&♀). Tail of hybrid is intermediate in length and carried in an arch. Fur resembles M. fascicularis, but posture and gait recall M. nemestrina. Data are limited for natural hybrids, but the observation of two hybrids in a single troop suggests they are fairly common. Bernstein and Gordon picture a three-way hybrid: M. fascicularis ♂ × (M. mulatta × M. nemestrina)♀. Ashley Montagu 1950; Avise and Duvall 1977; Bernstein 1966, 1968, 1974; Bernstein and Gordon 1980; Bongso and Hilmi 1981a, 1981b; Flower 1929a (p. 23); Gentry 1872; Groves 2001; Hill 1972, 1974 (Table 33); International Zoo Yearbook 1966 (p. 388), 1974 (p. 370), 1975 (p. 363); Ledbetter et al. 1979; Niemeyer 1868; Ridley 1906; Zuckermann 1931 (p. 338), 1953 (p. 942).
× Macaca nigra (♀) [Sulawesi Crested Macaque] CHR. DRS. A hybrid lived for seven hours after death. Ashley Montagu 1950; Chiarelli 1961; Hill 1974 (p. 835); Martin et al. 1974.
× Macaca philippinensis [Philippine Macaque] CHR. CON: Palawan, Sulus? The National Zoological Park (USA) had hybrids. These taxa are now usually lumped. International Zoo Yearbook 1959, 1960, 1961, 1962.
× Macaca radiata [Bonnet Macaque] Hill (1974 Table 33) lists this cross, but the works he cites (Przibram 1910, p. 111; Rörig 1903, p. 290) cite Fitzinger (1864a) who lists only M. fascicularis × M. sinica.
× Macaca sinica (♂) [Toque Macaque] CHR. ENHI. HPF. DRS. Tosi et al. (2003) argue genetic evidence indicates the Stump-tailed Macaque (M. arctoides) is derived from ancient hybridization of this type. Reports of captive hybridization all seem to trace back to a single female hybrid reported by Fitzinger (1864).
× Macaca silenus (♀) [Lion-tailed Macaque] CHR(Rostock, Germany). DRS. A female hybrid lacked the mane of M. silenus. Her tail was almost as long as her head and body together. Hill 1974 (Table 33); International Zoo Yearbook 1971 (p. 265); Linke 1971; Linke and Ritscher 1980.
× Macaca thibetana (♀) [Père David’s Macaque] NHR(Hong Kong). The natural ranges of these macaques are disjunct, but both occur on the Kowloon Peninsula, where they are introduced. Care of M. fascicularis infants by M. thibetana adults may result in imprinting and encourage later hybridization. Burton and Chan 1987, 1996 (pp. 393, 398).
× Mandrillus leucophaeus (♀) [Drill] CHR. DRS. A male hybrid born at Hannover Zoo lived two hours. The same female also produced a hybrid with a Macaca maura male and with a Macaca nemestrina male. Knottnerus-Meyer 1904.
× Mandrillus sphinx (♀) [Mandrill] CHR. DRS. A hybrid, which lived, was born in the London Zoo on Oct. 13, 1878. It had a short tail. See: Cercocebus atys × Macaca fascicularis. Flower 1929a (p. 23); Sclater 1878.
× Papio cynocephalus (♂) [Yellow Baboon] CHR. Przibram (1910, p. 111) lists this cross as Cynocephalus hamadryas × Macaca cynomolgus. Schöpff 1871.

Macaca fuscata [Japanese Macaque]
See also: Macaca arctoides; M. cyclopis.
× Macaca mulatta [Rhesus Macaque] CAENHR(Hong Kong). HPF. These macaques have disjunct natural ranges, but both occur on the Kowloon Peninsula, where M. fuscata is introduced. Burton and Chan 1996; Fujita 1990; International Zoo Yearbook 1962 (p. 224), 1972 (p. 317), 1973 (p. 322); Wolfe 1981.
× Macaca silenus (♂) [Lion-tailed Macaque] CHR(Manila). DRS. International Zoo Yearbook 1971 (p. 265).

ammospermophilus harrisii
Distributions of six hybridizing Sulawesi macaques: Macaca hecki (Heck’s Macaque), Macaca maura (Moor Macaque), Macaca nigra (Sulawesi Crested Macaque), Macaca nigresens (Black Macaque), and Macaca tonkeana (Tonkean Macaque).

Macaca hecki [Heck’s Macaque]
× Macaca nemestrina (♀) [Pig-tailed Macaque] CHR. DRS Bernstein and Gordon (1980, Table 6-1) report an aborted hybrid.
× Macaca nigra (♀) [Sulawesi Crested Macaque] CHR(Berlin). DRS. Viable hybrids were reported. Fooden 1969b; Hill 1974 (p. 799); International Zoo Yearbook 1971 (p. 265), 1972 (p. 318).
× Macaca nigrescens [Black Ape] ENHR(Indonesia). The hybrid zone is on Sulawesi’s northern peninsula near Gorontalo (see map above). Ciani et al. 1989; Groves 1980, 2001; Watanabe and Matsumura 1991.
× Macaca ochreata [Booted Ape] CHR(Berlin). DRS. International Zoo Yearbook 1971 (p. 265), 1972 (p. 318), 1978 (p. 373).
× Macaca tonkeana [Tonkean Macaque] ENHR(Indonesia). At the base of Sulawesi’s northern peninsula, north of Palu, there is a narrow (less than 20km-wide) hybrid zone (see map above). Although M. hecki and M. tonkeana differ with respect to multiple morphological traits, they are sometimes lumped due to hybridization. In a personal communication to the IUCN, however, M. Richardson says that this hybrid zone appears to be disappearing "due to human encroachment." Bynum et al. 1997a, 1997b; Ciani et al. 1989; Fooden 1969b; Gotoh et al. 2001; Groves 1980, 2001; Watanabe and Matsumura 1991; Watanabe et al. 1991a.

Macaca maura [Moor Macaque]
See also: Macaca fascicularis.
× Macaca nemestrina [Pig-tailed Macaque] CHR. Hill (1974, Table 33) mentions a putative hybrid of this type captured in Sumatra. M. maura is not native to Sumatra, so any such hybrid either occurred in captivity or resulted from human introduction of M. maura. Viable hybrids of both sexes have been produced. International Zoo Yearbook 1962 (p. 223), 1967 (p. 304), 1971 (p. 264), 1972 (p. 317); Schwarz 1934b.
× Macaca nigra (↔ usu. ♂) [Sulawesi Crested Macaque] CHR. DRS. Chiarelli 1973 (citing Bernstein); Fooden 1969b; Hill 1974 (pp. 799, 835); International Zoo Yearbook 1968 (p. 293), 1969 (p. 220), 1970 (p. 254), 1977 (p. 304), 1986 (p. 476), 1987 (p. 406).
× Macaca tonkeana (↔ usu. ♂) [Tonkean Macaque] HPF(♂&♀). CAENHR(Indonesia). There is a narrow (~30km-wide) hybrid zone at the north end of Sulawesi’s southwestern peninsula (see map above). These macaques are very different in appearance and have remained distinct outside the hybrid zone. However, in a genetic evaluation of macaques within the zone, Evans et al. 2001 found that a "minimum [of], 27-28 of 36 individuals (>75%) sampled in the contact zone are first- or second-generation hybrids and that hybridization has been occurring for multiple generations." In the hybrid zone, Y-chromosome DNA is mainly from M. tonkeana, which indicates a biased directionality to the cross. In addition, Brandon-Jones et al. (2004, p. 121) say a population on Malenge Island may be derived this cross. This population has been treated as a species, the Togean, Balantak macaque (Macaca togeana, Cynopithecus togeanus). Bernstein and Gordon (1980, Table 6-1); Ciani et al. 1989 Evans et al. 2001; Froehlich and Supriatna 1996; Groves 1980, 2001; Supriatna 1991.
× Mandrillus leucophaeus (♀) [Drill] CHR? DRS. A hybrid died soon after birth. Its mother also produced a hybrid with a Macaca nemestrina male, and with a Macaca fascicularis male. Knottnerus-Meyer 1904.

Macaca mulatta [Rhesus Macaque]
See also: Cercocebus torquatus; Chlorocebus aethiops; C. sabaeus; Macaca arctoides; M. cyclopis × M. fuscata; M. fascicularis; M. fuscata.
× Macaca nemestrina (↔ usu. ♂) [Pig-tailed Macaque] CHR. HPF(♂&♀). Bernstein and Gordon (Table 6-1) report three fertile males, but females are here presumed fertile by Haldane’s Rule. Parapatric contact zone in southeast Asia, but no natural hybrids have as yet been reported. Contact is between M. mulatta and leonina, a population often lumped with M. nemestrina. Duff and Lawson (2004) treat leonina separately, as Macaca leonina (Northern Pig-tailed Macaque). Nesturkh (1967, p. 47) says a male hybrid at Sukhumi Medico-Biological Station, in what was then the U.S.S.R., produced two female offspring in crosses with a Papio hamadryas female. Antonius 1951b; Avise and Duvall 1977; Bernstein 1974; Bernstein and Gordon 1980; Chiarelli 1973; Hill 1974 (Table 33); International Zoo Yearbook 1968 (p. 292), 1971 (p. 264), 1972 (p. 317).
× Macaca radiata (♂) [Bonnet Macaque] CHR(Leipzig, Germany). A male hybrid lived at least four years. Parapatric contact zone in peninsular India. Koyama and Shekar (1981) and Fooden et al. (1981) found that the boundary between these macaques lies further south than had previously been supposed. Approximately, it lies along a line extending from the northern end of the Western Ghats (20˚47´N), through the Manjra Plateau, to the Velikonda Range of the Eastern Ghats (15˚46´N), instead of along the Tapti and Godavari rivers, as had been thought. These same workers observed mixed mulatta-radiata troops in the boundary region, but reported no hybrids. Only a few such troops were observed, but the limited data suggest that in a natural setting the association is usually between M. radiata males and M. macaca females, which corresponds to the reported direction of the cross in captivity. Ashley Montagu 1950; Flower 1929a; Landois 1896b; Zuckerman 1931 (p. 338), 1953 (p. 942).
+ Ovis aries [Domestic Sheep] A video (MACSHP) on YouTube showing a captive rhesus macaque male attempting to have sex with a sheep demonstrates that very distinct animals will sometimes try to mate.
× Papio cynocephalus [Yellow Baboon] CHR. DRS. Ledbetter et al. 1979.
× Papio hamadryas (♀) [Hamadryas Baboon] CHR. DRS. Testicular biopsy of a single male indicated sterility. Moore et al. describe him as the sole long-term survivor of 26 hybrid pregnancies resulting from matings of this type. See Macaca mulatta × M. nemestrina. Ashley Montagu 1950; Moore et al. 1999.

Note: Macaca nemestrina here includes both Macaca leonina (Northern Pig-tailed Macaque) and M. nemestrina (Sunda Pig-tailed Macaque), which are listed separately by Duff and Lawson (2004). Thus, some crosses reported as involving Macaca nemestrina and listed here under that name may have actually involved M. leonina. Kloss (1919) notes the occurrence of natural hybrids between these two macaques in southernmost part of Thailand (Trang) where they have a parapatric contact zone. Hill 1974 (p. 680).

Macaca nemestrina [Pig-tailed Macaque]
See also: Cercocebus atys; C. torquata; Macaca assamensis; M. fascicularis; M. hecki; M. maura; M. mulatta.
× Macaca nigra (↔) [Sulawesi Crested Macaque] CHR. DRS. HPF(♀♀). Most hybrids are stillborn or aborted. Some reach maturity. Avise and Duvall 1977; Bernstein 1974; Bernstein and Gordon 1980; Hill 1974 (p. 835) International Zoo Yearbook 1986 (p. 476).
× Macaca silenus (♂) [Lion-tailed Macaque] CHR. DRS HPF(♀♀). Zukowsky describes several hybrids in detail. Ashley Montagu 1950; Avise and Duvall 1977; Bernstein and Gordon 1980; Chiarelli 1961, 1973; Steinbacher 1941 (p. 366); Zukowsky 1952.
× Macaca tonkeana (↔) [Tonkean Macaque] CHR. DRS. Avise and Duvall 1977; Bernstein and Gordon 1980 (Table 6-1).
× Mandrillus leucophaeus (♀) [Drill] CHR. DRS. It is unclear from the report whether this hybrid survived. The mother also produced a hybrid with a Macaca maura male and with a Macaca fascicularis male. Knottnerus-Meyer 1904.
× Papio ursinus (♀) [Chacma Baboon] CHR. DRS. Blyth (1863) briefly describes a dead infant male hybrid. It was preserved in alcohol at the Indian Museum in Calcutta (Anderson 1881, p. 82). Anderson 1881; Mollika 1863.

Macaca nigra [Sulawesi Crested Macaque]
See also: Macaca fascicularis; M. maura; M. nemestrina.
× Macaca nigrescens [Gorontalo Macaque] ENHR(Indonesia). The hybrid zone is near western end of Sulawesi’s northern peninsula between Inobonto and Pinolosian (see map above). These macaques are sometimes lumped due to hybridization. Ciani et al. 1989; Groves 1980, 2001; Watanabe and Matsumura 1991.
× Macaca silenus (♂) [Lion-tailed Macaque] CHR. DRS. Hybrids of both sexes have been reported. Avise and Duvall 1977; Bernstein and Gordon (1980, Table 6-1); International Zoo Yearbook 1973 (p. 322).
× Macaca tonkeana (♀) [Tonkean Macaque] CHR. DRS. Avise and Duvall 1977; Bernstein and Gordon (1980, Table 6-1).

Macaca nigrescens [Gorontalo Macaque] See: Macaca nigra.

Macaca ochreata [Booted Macaque]
See also: Macaca hecki.
× Macaca tonkeana [Tonkean Macaque] ENHR(Indonesia). The hybrid zone is at the head of Sulawesi’s southeastern peninsula near Walu (see map above). Groves 2001; Watanabe et al. 1991b.

Macaca philippinensis [Philippine Macaque] See: Macaca fascicularis.

Macaca radiata [Bonnet Macaque]
See also: Chlorocebus aethiops; Macaca assamensis; M. fascicularis.
× Macaca sinica (↔) [Toque Macaque] CHR. DRS. Flower 1929b (p. 23); Hill 1937, 1974 (Table 33 and p. 699); International Zoo Yearbook 1969 (p. 219).

Macaca silenus [Lion-tailed Macaque] See: Macaca fascicularis; M. nemestrina; M. nigra.

Macaca sinica [Toque Macaque] See: Chlorocebus aethiops; Macaca fascicularis; M. radiata.

Macaca sylvanus [Barbary Macaque] See: Cercocebus sp.

Macaca thibetana [Père David’s Macaque] See: Macaca assamensis; M. fascicularis.

Macaca togeana [Togean Macaque | Balantak Macaque] See: Macaca maura × M. tonkeana.

Macaca tonkeana [Tonkean Macaque ] The tonkean macaque hybridizes with M. ochreata, M. maura and M. hecki in areas where their ranges overlap (see map above). See: Macaca hecki; M. maura; M. nigra; M. ochreata.

Mandrillus leucophaeus [Drill]
See also: Cercocebus torquatus; Macaca fascicularis; M. maura; M. nemestrina.
× Mandrillus sphinx (♂) [Mandrill] CHR. Parapatric contact in southwestern Cameroon, Gabon. Although most zoos now frown on hybridization and take stringent measures to prevent it, this hybrid was formerly very common in captivity. On the basis of the face color of the females, the purity of the Melbourne Zoo mandrill population was questioned. Genetic analysis showed all members of the population had drill mtDNA, which indicates the original female founder was hybrid. Chiarelli 1973; International Zoo Yearbook 1960 (p. 257), 1961 (p. 258), 1962 (p. 225), 1965 (p. 333), 1967 (p. 305), 1968 (p. 294), 1969 (p. 221), 1970 (p. 255), 1972 (p. 319), 1974 (p. 371), 1978 (p. 374), 1979 (p. 353), 1981 (p. 311); Painter et al. 1993. Internet Citations: DPZ.
× Papio hamadryas (♂) [Hamadryas Baboon] CHR. DRS. Chiarelli 1973; Gray 1972; International Zoo Yearbook 1961 (p. 258).
× Papio papio (♂) [Guinea Baboon] CHR(London). A hybrid lived one day. Hill 1970 (p. 218).

Mandrillus sphinx [Mandrill]
See also: Cercocebus atys; C. torquatus; Mandrillus leucophaeus.
× Papio anubis (♂) [Olive Baboon] CHR. CON: southern Cameroon. A viable male hybrid was reported. Chiarelli 1973.
× Papio cynocephalus (♂) [Yellow Baboon] CHR(Philadelphia, U.S.). DRS. A female hybrid (listed as Papio langheldi × Papio sphinx) lived one day. Ashley Montagu 1950.
× Papio hamadryas (♀) [Hamadryas Baboon] CHR. DRS. Viable hybrids of both sexes have been reported. Chiarelli 1973; International Zoo Yearbook 1973 (p. 323), 1975 (p. 363), 1978 (p. 374), 1979 (p. 353), 1980 (p. 423), 1984/1985 (p. 520).

Nasalis larvatus [Proboscis Monkey]
× Pygathrix nemaeus (♂) [Red-Shanked Douc Langur] CHR. DRS. A hybrid born at Zoo Erfurt (Germany) on August 27, 1981 lived nearly five years. It died of a bezoar. International Zoo Yearbook 1983 (p. 317). Internet Citations: DOUC.

Note: Baboons of the genus Papio comprise a set of parapatric populations spread over most of sub-Saharan Africa. According to Newman et al. (2004, p. 25), "All baboon allotaxa appear to be capable of producing viable and fertile offspring when crossed. Active, natural, and genetically complex hybrid zones have been found wherever the critical interfaces have been investigated in the field."

Papio anubis [Olive Baboon] (= doguera)
See also: Cercocebus chrysogaster; Chlorocebus aethiops.
× Papio cynocephalus (♀ prob. ↔) [Yellow Baboon] HPF(♀♀). CAENHR(eastern Africa). Hybridization occurs across Kenya and Tanzania, from Lake Turkana to the northern end of Lake Tanganyika. In Kenya these types are largely separated by a waterless region with very scattered bush, but contact occurs in the southwestern Amboseli Basin, where mixed groups, including hybrids have been observed. Probable hybrids have also been observed on the opposite side of the basin, between Ithumba and Simba. The IUCN (Internet Citations: ABCYN) says that "there is a broad clinal hybrid zone between [the] Laikipia District, just to the north-east and east of Mt. Kenya, and the Lower Tana River, Kenya coast. Baboons in this more than 200-km wide region are intermediate and cannot be readily allocated to either P. anubis or P. cynocephalus (baboons become increasingly "yellow-like" in their phenotypes towards the Kenya Coast; T. Butynski and Y. de Jong pers. comm.)." Newman et al. found nearly identical mtDNA haplotypes in olive and yellow baboons, which suggests backcrossing has occurred and that at least female hybrids are fertile. Due to hybridization, these very different looking baboons are sometimes lumped. Samuels and Altmann report that a P. anubis male had no trouble joining a P. cynocephalus group. It assimilated rapidly and thereafter mated with several P. cynocephalus females. Grubb et al. 2003; International Zoo Yearbook 1962, 1965, 1966, 1971, 1972, 1973, 1975; 1977; Jolly 1993; Kingdon 1971 (p. 177); Lucotte and Dubouch 1980; Maples and McKern 1967; Newman et al. 2004; Samuels and Altmann 1986. Internet Citations: ZOOH.
× Papio hamadryas (♀) [Hamadryas Baboon] CAENHR. CON: Ethiopia. Probable ACZ (P. anubis occurs above P. hamadryas). In the Awash Valley, morphological and genetic traits vary clinally across the zone, which is about 20-km wide. Within the hybrid zone, some troops are composed entirely of hybrids. Zinner et al. (2001) say contact is probable also in Eritrea. Chiarelli (1973) says captive crosses are more successful (multiple viable hybrids of both sexes), when the female parent is P. hamadryas. In the opposite direction he lists a single hybrid that lived an hour. Inviability of hybrids with the cross reversed may account, then, for Shotake’s (1981, p. 302) observation that natural matings between P. hamadryas males and P. anubis femalesdid not result in genetically identifiable progeny the following year. Jolly (1993, p. 79) notes brown pelage in western P. hamadryas is probably the result of gene flow from adjacent P. anubis (Kummer was probably the first to make this suggestion). Beehner et al. 2005; Brett et al. 1977; Chiarelli 1961, 1973; Hill 1970 (pp. 17, 29, 151, 217), 1977 (p. 305); International Zoo Yearbook 1984/1985 (p. 520), 1987 (p. 406); Iwamoto 1980; Kawai and Sugawara 1976a, 1976b; Kummer 1968 (pp. 23, 121); Kummer et al. 1970; Nagel 1971, 1973; Newman 1997; Phillips-Conroy 1978; Phillips-Conroy and Jolly 1981, 1986; Shotake 1981; Shotake et al. 1977; Sugawara 1979, 1982; Woolley-Barker 1999. Internet Citations: DPZ.
× Papio papio (♀?) [Guinea Baboon] CAENHR(southern Mali, Guinea, Sierra Leone). HPF. Jolly 1993 (p. 79) says, "the brown eastern Guinea [Baboon] populations could well be the result of the introgression of Anubis genes, pointing to a Guinea-Anubis hybrid zone." Chiarelli 1973; Hill 1970 (pp. 29, 150); International Zoo Yearbook 1969 (p. 221), 1971 (p. 266), 1972 (p. 318), 1974 (p. 371); Pollock (cited by Grubb et al. 2003, p. 1326).
× Theropithecus gelada [Gelada Baboon] ONHR(Bole Valley, Ethiopia). Three probable hybrids were reported. De Jong and Butynski 2010; Dunbar and Dunbar 1974.

Note: Analysis of morphometric characters indicates that two populations (cynocephalus, kindae), treated as races of Papio cynocephalus, hybridize between Lake Tanganyika and Lake Malawi (southeastern Africa). Formerly, kindae, a very distinct form, was treated as a separate species, Papio kindae (Kinda Baboon). Ansell 1978; Freedman 1963; Hill and Carter 1941; Jolly 1965, 1993; Lönnberg 1919.

Papio cynocephalus [Yellow Baboon]
See also: Lophocebus sp., Macaca fascicularis; M. mulatta; Mandrillus sphinx; Papio anubis.
× Papio hamadryas (♂?) [Hamadryas Baboon] CHR. Common in captivity. CON: Juba River valley (at Ethiopian-Somalian border). Of five male, and four female, hybrids born at Madrid Zoo in 1976, three and two, respectively, reached maturity. The Berlin Zoo is said to have had a hybrid in 1882. International Zoo Yearbook 1968 (p. 293), 1969 (p. 220), 1971 (p. 266), 1978 (p. 374), 1980 (p. 423), 1981 (p. 311), 1982 (p. 413), 1984/1985 (p. 520), 1986 (p. 477), 1987 (p. 406), 1988 (p. 456), 1989 (p. 317), 1990 (p. 438).
× Papio papio (♀?) [Guinea Baboon] CHR. DRS. Hybrids of both sexes are reported. Ashley Montagu 1950; International Zoo Yearbook 1969 (p. 221), 1970 (p. 255), 1971 (p. 266), 1972 (p. 318), 1973 (p. 323); Peláez 1982.
× Papio ursinus [Chacma Baboon] CHR. ENHI. CON: eastern Zambia. Hill (1974, p. 340) describes a population of eastern Zambia, and of adjacent Malawi and Mozambique, jubilaeus, as "somewhat intermediate between P. cynocephalus and P. ursinus, being long-limbed with brown hands, but with the cranial characters of P. ursinus, of which it is now regarded as a dwarf race." P. cynocephalus is much smaller than P. ursinus. Since jubilaeus is also geographically intermediate, it can be regarded as a PHP of this cross. In addition, Newman et al. (2004, p. 25) say "gray-footed chacma baboons" are geographically and morphologically intermediate between typical chacmas and yellows. Hill (1974, pp. 340-342, also Plates IX and X); International Zoo Yearbook 1960 (p. 257), 1974 (p. 371), 1977 (p. 305).

Papio hamadryas [Hamadryas Baboon]
See also: Macaca mulatta; Macaca mulatta × M. nemestrina; Mandrillus leucophaeus; M. sphinx; Papio anubis; P. cynocephalus.
× Pan troglodytes (♀) [Chimpanzee] DRS. These animals have been observed mating in captivity, but no hybrids were reported. Matsubayashi et al. 1978.
× Papio papio (♀) [Guinea Baboon] CHR. DRS. These hybrids were formerly common in captivity. Jolly (1993, p. 79) notes that the ranges of these very similar baboons lie more than 5,000 km apart and that they are separated by the range of P. anubis (which hybridizes with both). This configuration, then, may represent a split range situation similar to that seen in the Hooded and Carrion crows (Corvus cornix, C. corone), or the Black Crested and Northern White Cheeked gibbons (Hylobates concolor, H. leucogenys). Chiarelli 1973; Hill 1970 (pp. 205, 217); International Zoo Yearbook 1968 (p. 293),1970 (p. 255), 1971 (p. 266), 1973 (p. 323) 1975 (p. 363); Petzsch 1951; Steinmetz 1940.
× Theropithecus gelada [Gelada Baboon] CHR. CON: Ethiopia. International Zoo Yearbook 1982 (p. 413); Martin et al. 1974; van Gelder 1977b.

Papio kindae [Kinda Baboon]. See note immediately preceding Papio cynocephalus.

Papio papio [Guinea Baboon]
See also: Papio anubis; P. cynocephalus; P. hamadryas.
× Papio ursinus (♂) [Chacma Baboon] CHR(Lisbon). DRS. Six male hybrids were reported (International Zoo Yearbook 1970, p. 255). In an old report, Landois (1896a, p. 98) says the sire was three times the size of the dam and that he was an attentive father ("besorgtester Vater"). International Zoo Yearbook 1968 (p. 293), 1969 (p. 221), 1970 (p. 255), 1971 (p. 266), 1973 (p. 323), 1989 (p. 317).
× Theropithecus gelada (♀) [Gelada Baboon] CHR. DRS. Chiarelli (1973, p. 302) says a hybrid was viable and fertile, but does not specify the hybrid’s sex. International Zoo Yearbook 1975 (p. 364).

Papio ursinus [Chacma Baboon] See: Macaca nemestrina; Papio cynocephalus; P. papio.

Presbytis chrysomelas [Sarawak Langur]
× Presbytis femoralis [Banded Monkey] NHR(Malaysia). Probable hybrids (from Miri, Sarawak) were described as a species (Presbytis arwasca Miller 1934). Brandon-Jones 1999; Brandon-Jones et al. 2004 (p. 124).
× Presbytis rubicunda [Red Langur] ENHR??. Banks (1929) offered evidence that a population in western Sarawak, cruciger (Red-banded Langur), is derived from this cross, but Hill (1960, p. 32) believed cruciger was "an erythristic mutant of chrysomelas."

Note: Hill (1960, p. 32) notes that in Selangor Province, in western Malaya, hybridization occurs between two populations (nubigena, siamensis) treated as races of Presbytis melalophos. In addition, Hill describes another population, australis, as an "intermediate race between femoralis and siamensis," two other populations treated as races of P. melalophos.

Presbytis comata [Grizzled Langur]
× Presbytis fredericae [Banded Monkey] ONHR(w Java). Probable hybrids are known from Linggo (7˚06´S,109˚35´E), Mt. Sawal (7˚12´S,108˚16´E) and Ceringin (7˚21´S 108˚30´E). Duff and Lawson (2004) lump these taxa. Brandon-Jones et al. 2004 (p. 124); Nijman 1997, 2001.

Presbytis femoralis [Banded Monkey] See: Presbytis chrysomelas.

Presbytis hosei [Grey Langur]
× Presbytis rubicunda [Red Langur] NHR(Malaysia)?? White leaf-monkeys of an unknown type have been observed in the Danum Valley and in the Ulu Segama Reserve (Sabah). They associated both with P. hosei, and with P. rubicunda. Stünkel (2003, p. 61) says they may hybrids.

Presbytis melalophos [Mitred Langur]
× Presbytis thomasi [North Sumatran Langur] Parapatric contact zone in the northern part of Sumatra. No hybrids as yet reported. Oates 1994.

Presbytis thomasi [North Sumatran Langur] See: Presbytis melalophos.

Presbytis rubicunda [Red Langur] See: Presbytis chrysomelas; P. hosei. Two populations (ignitus, rubidus), treated as races of P. rubicunda, hybridize in the northern part of Sarawak (Baram). Hill 1960 (p. 32).

Pygathrix cinerea [Grey-Shanked Douc Langur]
× Pygathrix nemaeus [Red-Shanked Douc Langur] CHR. ENHR. CON: Vietnam. According to the IUCN, there is strong evidence that P. cinerea "hybridizes with Red-shanked Doucs (P. nemaeus) in a small sympatric distribution area of Quang Nam Province (Nadler 1997, Nadler and Roos in press)." See: Pygathrix nemaeus × Pygathrix nigripes. Internet Citations: CINER.

Pygathrix nemaeus [Red-Shanked Douc Langur]
See also: Nasalis larvatus.
× Pygathrix nigripes [Black-Shanked Douc Langur] CHR. ENHR. CON: Vietnam. According to the IUCN, Pygathrix nigripes "hybridizes with the Red-shanked Douc Langur, Pygathrix nemaeus, where their distributions meet in Viet Nam." They also say the critically endangered Grey-Shanked Douc Langur (Pygathrix cinerea) "has been considered either a subspecies of P. nemaeus, or a hybrid of P. nemaeus and P. nigripes (Nadler 1995)," and that nemaeus "is strongly suspected to hybridize in the wild with both P. cinerea and P. nigripes. However, knowledge of the zones of transition between the three taxa is limited…(Timmins and Duckworth 1999; Nadler et al. 2004). Both these species have at times been considered conspecific with P. nemaeus." Internet Citations: NEMAE, NIGRI.

Rungwecebus kipunji [Kipunji] See: Lophocebus sp. × Papio cynocephalus

ammospermophilus harrisii
Spatial relationships of grey langurs in peninsular India (per IUCN, 2012).

Semnopithecus dussumieri [Southern Plains Grey Langur]
× Semnopithecus entellus [Northern Plains Grey Langur] ENHR(eastern India). A lengthy hybrid zone exists between these two langurs in eastern India (see map above). Brandon-Jones 2004 (p. 1572)
× Semnopithecus hector [Tarai Grey Langur] NHR(northern India)?? Brandon-Jones et al. (2004, p. 92) says a specimen from Gorakhnath Temple (c. 26˚45ʹ N 83˚24ʹE), about four kilometers northwest of Gorakhpur, Uttar Pradesh was a probable hybrid of this type (NB: S. dussumieri as it is treated here, and as it is recognized by the IUCN, is a composite of Brandon-Jones’s S. entellus achates and S. e. achises). Another such specimen was from the Kusmi Forest (c. 26˚45ʹ N 83˚30ʹE). A third was from Mohaddipur North (c. 26˚45ʹ N 83˚24ʹE), near Gorakhpur. However, IUCN distribution maps show the ranges of dussumieri and hector as disjunct, and Brandon-Jones (p. 1566) notes that the fact that hybridization of this type actually occurs needs confirmation. Also see Roonwal (1979).
× Semnopithecus hypoleucos [Black-footed Grey Langur] ENHR(India) CON: see map above. A series of specimens from the vicinity of Karwar (Karnataka, India) were variably intermediate. A hybrid zone exists in the Western Ghats, and probably extends from about 12˚ to 20˚ N). Brandon-Jones 2004 (pp. 1554, 1557).
× Semnopithecus priam [Tufted Grey Langur] ENHR (southern India). A hybrid zone exists where these two langur populations interface (see map above). Pithecus entellus priamellus (Pocock 1928) is known only from the holotype (from Sharnelli Estate, Nelliampathy Plateau, Kerala, India). Geissmann (1995, p. 134) says it is probably a hybrid derived from this cross. He also mentions a troop of probable hybrids of this type. NB: S. dussumieri as it is treated here, and as it is recognized by the IUCN, is a composite of the two populations Brandon-Jones calls S. entellus achates and S. e. achises. Brandon-Jones 2004 (p. 1557).
× Trachypithecus johnii (♀) [Hooded Langur | Nilgiri Langur] ENHR(southern India). There is a hybrid zone in southern part of the Western Ghats. So-called Brown Langurs seen in T. johnii troops in the vicinity of Agastyamalais and Anaimalais are probably hybrids of this type. Their pelage and calls are intermediate. Brandon-Jones (p. 1563) says the holotype specimen for Semnopithecus hypoleucos is a hybrid of this type. Some experts in the field believe S. hypoleucos itself is a hybrid population derived from this cross (Molur et al. 2003). NB: Note that S. dussumieri as it is treated here, and as it is recognized by the IUCN, is a composite of Brandon-Jones’s S. entellus achates and S. e. achises. Brandon-Jones 2004 (pp. 1553, 1555); Hohmann 1988; Hohmann and Hertzog 1985; Oates 1982.
× Trachypithecus vetulus [Purple-faced Langur] CHR. DRS. A hybrid lived an hour. Hill 1936.

Semnopithecus entellus [Northern Plains Grey Langur]
See also: Semnopithecus dussumieri.
× Trachypithecus pileatus [Capped Langur] CHR. CON: possibly in western Bangladesh. London Zoo had a hybrid in 1912. There is a parapatric contact zone in Bengladesh and eastern India. Edwards 1996 (p. 33). Internet Citations: ZOOH.

Semnopithecus hector [Tarai Grey Langur] See: Semnopithecus dussumieri.

Note: Some experts in the field believe Semnopithecus hypoleucos is a hybrid population derived from interbreeding of Semnopithecus dussumieri and Trachypithecus johnii. Molur et al. 2003.

Semnopithecus hypoleucos [Black-footed Grey Langur]
See also: Semnopithecus dussumieri; Semnopithecus dussumieri × Trachypithecus johnii.
× Semnopithecus priam [Tufted Grey Langur] NHR(southern India). A hybrid zone exists in the southern part of the Western Ghats (between 10˚ and 12˚ N). The name elissa Pocock, 1928 was based on probable hybrids (from Nagarhole, Karnataka). Brandon-Jones et al. 2004 (pp. 1557, 1562).
× Trachypithecus johnii [Hooded Langur | Nilgiri Langur] ENHR(southern India). CON: Nilgiri Hills. An intermediate population exists in the Karnataka-Kerala border area. The name aeneas Pocock, 1928 was based on probable hybrids (from Nagarhole, Karnataka). Brandon-Jones 2004 (p. 1562); Hohmann and Hertzog 1985.

Semnopithecus priam [Tufted Grey Langur]
See also: Semnopithecus dussumieri; S. hypoleucos.
× Trachypithecus johnii (♀) [Hooded Langur | Nilgiri Langur] ENHR(southern India). CON: Nilgiri and Annamalai hills. Brandon-Jones (p. 1583) says T. johnii females leave their natal troops, join S. priam troops, and then hybridize. Brandon-Jones 2004 (pp. 1553, 1556, 1559-1562, 1583); Hohmann and Hertzog 1985.
× Trachypithecus obscurus (♂) [Dusky Langur] CHR. DRS. Hill 1939.
× Trachypithecus pileatus (♂) [Capped Langur] CHR. DRS. LVH (hybrids are typically stillborn or aborted). Hill 1957 (p. 70).
× Trachypithecus vetulus [Purple-faced Langur] Brandon-Hill (2004, p. 1560) says that where these two langurs meet in Sri Lanka hybridization occurred in the past, probably, or still does.

Theropithecus gelada [Gelada Baboon] See: Papio anubis; P. hamadryas; P. papio.

Trachypithecus auratus [Javan Langur | Ebony Leaf Monkey]
× Trachypithecus cristatus [Silvered Langur | Silvered Leaf Monkey] CHR. DRS. Waters et al. ( 2001) say "the Javan langur group [at Bristol Zoo, UK] began with one male and three females. These animals were believed to be of the subspecies Trachypithecus auratus auratus. The original male died in 1985 without siring offspring. Females were then sent to Twycross Zoo to mate with their breeding male and returned to Bristol where they gave birth. Another male arrived at Bristol in 1989. He was later suspected to have belonged to a Sumatran subspecies of T. cristatus, so his offspring were thought to be hybrids, as were the offspring from the breeding male at Twycross (Waters, 1994). Recent taxonomic investigations of the Bristol group have suggested that they may be ebony/silver leaf monkey hybrids or of unknown origin (J.B. Carroll, pers. comm.)." Bristol euthanized two males suspected to be hybrids of this type (Walker et al. 2001).

Trachypithecus barbei [Tenasserim Langur] See: Presbytis obscurus × P. phayrei

Trachypithecus cristatus [Silvered Langur | Silvered Leaf Monkey]
See also: Trachypithecus auratus.
× Trachypithecus obscurus [Dusky Langur] CHR. CON: southeast Asia. International Zoo Yearbook 1979 (p. 355).
× Trachypithecus phayrei (♀) [Phayre’s Langur] CHR. Parapatric contact zone in southeast Asia. A notice in the Proceedings of the Zoological Society of London about a hybrid of this type read as follows: "The Secretary read the following extract of a letter addressed to him by Babu Ram Bramha Sányál, C.M.Z.S., dated Zoological Garden, Calcutta, July 27, 1893:—’I am not aware whether closely allied species of Semnopitheci have ever interbred anywhere. They are rather exclusive in their ideas in respect to matrimonial relationship. Anyhow, such an event has just happened in this Garden. The Phayre’s Langur (Semnopitheeus phayrei, Blyth) has given birth to a young one—a lovely little babe of a delicate light orange colour. As there has been no other male in the same cage except the S. cristatus, there is no doubt of the young one being a hybrid between these two species. These Monkeys have been living together since 1880, and although they agreed very well, they were never observed to be over friendly. Even now the male does not appear to take any interest in the offspring.’" Sányál (1858-1908) was the first superintendent of the Alipore Zoological Gardens in Kolkata. Sányál 1893.

Trachypithecus johnii [Hooded Langur | Nilgiri Langur] See: Semnopithecus dussumieri; S. hypoleucos; S. priam.

Trachypithecus obscurus [Dusky Langur]
See also: Semnopithecus priam; Trachypithecus cristatus.
× Trachypithecus phayrei [Phayre’s Langur] CHR. NHI. Parapatric contact zone (head of Malay Peninsula). A taxon, Presbytis barbei (Tenasserim Langur) is based on two specimens originally described (Blyth 1847) as being from Tenasserim, a region along the Burma-Thailand border between the ranges of P. obscurus and P. phayrei. Geissmann et al. discovered a specimen of unknown provenance in the Bangkok Zoo that they believe to be a living representative of this taxon. Its mtDNA is closely similar to, but not identical to that of P. obscurus and P. phayrei, and so may fall within the range of variation for one or both. These facts suggest P. barbei as a PHP of this cross. Geissmann et al. 2004; International Zoo Yearbook 1968 (p. 295), 1969 (p. 222), 1970 (p. 256).

Trachypithecus phayrei [Phayre’s Langur] See: Trachypithecus cristatus; T. obscurus.

Trachypithecus pileatus [Capped Langur] See: Semnopithecus priam.

Trachypithecus vetulus [Purple-faced Langur] See: Semnopithecus dussumieri; S. priam.

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


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