Golden Snub-nosed Langur (Pygathrix roxellana)

MORPHOLOGY: This species has a sacculated stomach and symbiotic bacteria in the stomach to assist in the breakdown of cellulose (Gao and Liu, 1995). The teeth, masseter muscle and temporalis muscle of the golden snub-nosed langur are stronger as compared to other Chinese monkeys, which assist in the processing of food that contains a large amount of roughage (Gao and Liu, 1995). This species has high levels of sexual dimorphism as compared to other Old World monkeys (Jablonski and Ruliang, 1995). Males have a pair of fleshy flaps on their upper lip (Qi, 1986). This species has an upturned snub nose, hence the common name (Qi, 1986). The average body mass for an adult male golden snub-nosed langur is around 15.0 kilograms, and for the female it is between 6.5 and 10.0 kilograms (Fleagle, 1988). The golden snub-nosed langur has a dental formula of 2:1:2:3 on both the lower and upper jaws (Ankel-Simons, 2000). This species moults its fur between August and October (Davison, 1982). The male's penis is black and the scrotum is bluish white in coloration (Davison, 1982). The perineum of the male is bright blue in a narrow line, with sky blue skin surrounding the ischial callosities (Davison, 1982). The coloration of the male's perineum mimics the facial coloration, which is convergent with the mandrill, Mandrillus sphinx (Davison, 1982). Adult males also have secondary sexual characteristics including: large canines, upper-lip warts, and long hair on the back (Liang et al., 2000). Large canines start to appear at 4.9-5.4 years, upper-lip warts begin to appear from 5.4-6.2 years, and back hair begins to appear from 5.6-6.3 years of age (Liang et al., 2000).

The infant has a natal coat that is dark gray on the dorsal side and whitish gray on the ventral side (Rapaport and Mellen, 1990, 1987). Qi (1986) reports that the newborn has long black hair on the head and back. The hair on the ears of the infant is white (Rapaport and Mellen, 1990). The skin of the infant is white except for the area around the mouth which is pale pink and the palms and fingers which are dark pink (Rapaport and Mellen, 1990). Qi (1986) reports that the eyes and the nostril bridge are bluish-green in color of the newborn. Gold colored hairs begin to appear on the infant at about 9 days of age (Rapaport and Mellen, 1990). At around day 12, the area between the eyes become bluish in color (Rapaport and Mellen, 1990).

This species has three subspecies each having their pelage coloration:

• Pygathrix roxellana roxellana: The dorsal pelage of this subspecies is dull golden-red (Wang et al., 1998). The shoulders, forelimbs, and hindlimbs are a dusky blackish-brown color (Wang et al., 1998). This subspecies has a relatively short tail and relatively long hindlimbs as compared to the other subspecies (Wang et al., 1998).
• Pygathrix roxellana qinlingensis: The overall coloration of this subspecies is described as being "brilliant golden" (Wang et al., 1998).
• Pygathrix roxellana hubeiensis: The dorsal pelage of this subspecies is pale golden-reddish (Wang et al., 1998). This subspecies is the most pallid of the subspecies of Pygathrix roxellana (Wang et al., 1998). This subspecies also has a relatively long tail, longer than the other two subspecies (Wang et al., 1998).

RANGE:
The golden snub-nosed langur is found in the country of China. This species lives in montane, conifer forests where snow cover can last for 6 months or more (Allen, 1938; cited in Davison, 1982). In Qinling Mountains, Shaanxi province, this species lives in mixed deciduous-broadleaf evergreen forests from 1200 to 3000 meters (Gao and Liu, 1995). The forest types this species lives in are conifer, mixed-conifer broadleaf, and deciduous broadleaf (Happel and Cheek, 1986).

This species has three subspecies each having differing ranges:

• Pygathrix roxellana roxellana: This subspecies is found in western Sichuan and southern Gansu, from the Dadu River in the east to the Sichuan Basin in the west (Wang et al., 1998).
• Pygathrix roxellana qinlingensis: This subspecies is found from in the south of the Qinling Mountains, which include: Taibei, Zhouzhi, Fuping, Ninshaan, Shiquan, and Zhashui of southern Shaanxi (Wang et al., 1998).
• Pygathrix roxellana hubeiensis: This subspecies is found in the Shennongjia forest region and areas adjacent, Fangxian, Xingshan, and Badong in western Hubei and Wushan in southeastern Sichuan (Wang et al., 1998).

ECOLOGY:
The golden snub-nosed langur is a folivorous species, but will also eat seeds, fruit, bark, insects, and small vertebrates. On the ground they forage for wild onions. During the summer at Baihe Nature Reserve the primary food source for this species was mature leaves of Prunus and Sorbus (Kirkpatrick et al., 1999). In winter the number of tree species foraged upon greatly decreases during this time (Li et al., 2002). The diet changes in winter where the golden snub-nosed langur primarily forages for lichens found on trees, fruit of Pinus armandii, bark, and insects under bark (Li et al., 2002; Li, 2001). In winter preferred feeding trees tend to be larger than other trees of a given species (Li et al., 2002). Larger trees may be selected because larger trees potentially offer more food, provide better protection from predators, and increase foraging efficiency because they could see other sources of food around them better (Li et al., 2002). Pinus armandii and Abies fargesii are evergreen coniferous trees species that are preferred by the golden snub-nosed monkeys in winter (Li et al., 2002). In winter Litsea ichangensis and Salix walliciana are also preferred tree species (Li et al., 2002). Kirkpatrick et al. (1999) found that in winter the golden snub-nosed langur mostly lichens from the family Usneaceae and the buds and nonlignified bark of pencil-thin twigs. Large troops may split in the winter months to spread more widely in search of lichens, buds, and bark (Kirkpatrick et al., 1999, 1998).

At Shennongjia Nature Reserve the golden snub-nosed langur forages on 51 species of trees comprising 24 families (Li, 2001). The diet in the spring consists of buds and young leaves of 21 tree species, lichen and bark from the trees, insects from under the bark of 6 tree species, and the flowers of Cerasus discadenia (Li, 2001). The diet in the summer consists of young leaves of 20 tree species, lichens, bark, insects from under the bark of 4 tree species, and the fruit of Litsea ichangensis (Li, 2001). The diet in autumn consists of fruits from 17 tree species, young leaves of 5 tree species, lichen, bark, and insects from under the bark of 7 tree species (Li, 2001). The diet in winter consists of Pinus armandii fruits, lichens, bark, and insects from under the bark of 8 tree species (Li, 2001). Also in winter individuals come down to the ground to consume herbs and insects from under stones (Li, 2001). In winter insects are a significant food source and provide protein for the golden snub-nosed langurs (Li, 2001). Snow is eaten in winter by this species (Su et al., 1998).

Group sizes range according to the season, in winter the groups are smaller (20 to 30 individuals), and in the summer time they can be quite large (up to 600 individuals). The home range of this species is largest during the spring and smallest in the summer (Li et al., 2000). Li et al. (2000) found that autumn and winter home ranges are similar in size. The spring home range in the Qinling Mountains consist mostly of deciduous broadleaf forest (82.4%) (Li et al., 2000). The summer home range in the Qinling Mountains consists of 58.6% deciduous broadleaf forest and 40.5% coniferous and deciduous broadleaf mixed forest (Li et al., 2000). The home range in autumn in the Qinling Mountains consists of 76.5% deciduous broadleaf forest and 23.5% coniferous and deciduous broadleaf forest (Li et al., 2000). Groups of golden snub-nosed langurs tend to stay away from areas of human activity, especially commercial logging (Li et al., 1999; Su et al., 1998). One study found that the noise of chain saws upsets members of this species (Li et al., 1999). This is a semi-terrestrial and diurnal species. Golden snub-nosed langurs will spend 15.3% of the time on the ground, 24.3% in the low stratum of the forest, 28.2% in the middle stratum of the forest, and 32.2% in the canopy of the forest (Ren et al., 2001). Adult males are more frequently seen on the ground foraging for food and/or guarding the group against terrestrial predators (Ren et al., 2001). Adult females and immatures rarely descend to the ground level (Ren et al., 2001).

Predators of the golden snub-nosed monkey include: the red dog, Cuon alpinus, wolf, Canis lupus, Asiatic golden cat, Catopuma temmincki, leopard, Panthera pardus, golden eagle, Aquila chrysaetos, and goshawk, Accipiter gentilis (Hu, 1998; cited in Li et al., 2002; Zhang et al., 1999b). After a goshawk took a juvenile member of a group, for the rest of the afternoon the adult males kept to the periphery of the group with the young in the center and the spread of the group when foraging was reduced to a third of its former size (Zhang et al., 1999b).

LOCOMOTION:
The golden snub-nosed langur moves through the forest and on the ground quadrupedally (Fleagle, 1988). When the adult male walks on the ground quadrupedally, the tail is held in a typical macaque or baboon-like hitch (Davison, 1982). Also when walking quadrupedally the hands and feet are always plantigrade (Su et al., 1998). When walking on snow, this species will sometimes run on the knuckles (Ren et al., 1996/1997). In captivity, a male's typical movement off the ground consisted of 92% walking along horizontal supports, 2% vertical climbing, and 6% arboreal display rushes, and for the females it was 20% walking, 45% modified brachiation, 5% vertical leaping, and 30% vertical climbing (Davison, 1982). Arm swinging is used by the golden snub-nosed langur to assist in moving to lower parts of a tree trunk or lower branches (Su et al., 1998). This species has been observed walking bipedally for a short distance (Su et al., 1998).

SOCIAL BEHAVIOR:
The golden snub-nosed langur has a unimale social system (Ren et al., 1998). These groups come together in the summer time to form larger troops, which are composed of many one-male groups and several all-male groups (Ren et al., 1998). Troops may split and come together on a daily basis at some sites (Kirkpatrick, 1998). One-male groups are made up of an adult male, several adult females, and several immatures (Kirkpatrick et al., 1999). Males live solitarily or on the periphery, in all-male groups, to unimale group before they are able to obtain a group of their own (Kirkpatrick et al., 1999). When troops move, the all-male group will be in front followed by the one-male groups, which are lead by the adult male (Ren et al., 1998). All-male groups will stay in close contact with each other, embracing when sleeping and foraging together (Ren et al., 1998).

Spontaneous aggressive incidents between individuals are not high, but when aggression does occur, the fights are often reciprocal (Ren et al., 1992, 1991). Although some have found that in captivity adult males often fight in the presence of females (Kirkpatrick, 1998). Adult males tend to act as sort of a mediator between adult females when they fight, often trying to appease or reassure each of the females in order to calm them down (Ren et al., 1992, 1991). Open mouth is the most frequent behavior seen when a male acts as a mediator between two adult females that are fighting (Ren et al., 1991). Males will also show play, mounting, or embracing when acting as a mediator, and females will usually respond to the male with crouch (Ren et al., 1991). Ren et al. (1991) suggest "the maintenance of harmonious relation among females has a high adaptive value for males." Reconciliation behavior after fights include: open mouth, rapid grooming, embrace, and crouching (Ren et al., 1992, 1991). Reconciliation is defined as: "a nonagonistic contact between two former opponents not long after an agonistic confrontation between them" (de Waal and van Roosmalen, 1979; cited in Ren et al., 1991).

In captivity, males were found to be protective of his infant and the mother (Rapaport and Mellen, 1990). The male will position himself between himself and public (Rapaport and Mellen, 1990). The male will also direct aggression towards outside sources, in the form of hard stares, bow and lunge threats, and other displays (Rapaport and Mellen, 1990). Males will also go to the mother with an infant when the infant emits a squeak call (Rapaport and Mellen, 1990). Males will not direct aggression against their infants, except possibly mild threats when the infant engages in rough play (Rapaport and Mellen, 1990). Males may also groom their infants (Kirkpatrick, 1998). Females spend a large amount of time in an embrace with their infants, with embracing being mutual for the first ten days of life (Rapaport and Mellen, 1990, 1987). After ten days, the mother has the primary responsibility for maintaining contact with the infant (Rapaport and Mellen, 1990). Infants cling to their mothers against the armpits and waist (Qi, 1986). Although there has been reported that there is little aggression amongst members of this species, infanticide has occurred in captivity (Zhanget al., 1999a). Adult males were found to attack and bite infants with their canines (Zhanget al., 1999a).

VOCAL COMMUNICATION:
group contact call: This soft call sounds like "wu-aa" and is emitted by group members when foraging to keep in contact with each other (Happel and Cheek, 1986).

ga-ga call: This call is uttered when a group member finds a rich food source, and it serves to notify other group members (Happel and Cheek, 1986).

kuku: This call serves to communicate threat on the part of the sender (Ren et al., 1995). The call may be done with the mouth closed (Ren et al., 1995). This call is emitted sometimes when adult females are trying to interfere with another adult female who is soliciting mounting from an adult male (Ren et al., 1995). This call is also emitted when an individual is frightened (Happel and Cheek, 1986).

alarm call: This call sounds like "ou-ga, ou-ga" (Zhang et al., 1999b).

chuck: This call sounds like "ee-tcha" or "ah-tcha", with the "tcha" being a noise between 3.5 and 5.5 kilohertz (Tenaza et al., 1988). The "ee" or "ah" is clearer and of a lower pitch than the "tcha" (Tenaza et al., 1988). The mean duration of this call is 1.1 seconds (Tenaza et al., 1988). This call is emitted by both males and females (Tenaza et al., 1988).

shrill calls: These calls are comprised of various calls of shrill squeaks, squeals, and shrieks, which are between 1 and 7 kilohertz in frequency (Tenaza et al., 1988). This call is emitted by both males and females (Tenaza et al., 1988). Female shrill calls can occur without any body movement or no visible sign that the call is being made (Tenaza et al., 1988).

bawl: This call sounds like a loud "wah", "bah", "ahhh", or "aww" (Tenaza et al., 1988). This call resembles the cry made by a bawling human, Homo sapiens, baby (Tenaza et al., 1988). The mean duration of this call is 0.8 seconds (Tenaza et al., 1988). This call is emitted by both males and females, but is emitted more frequently by males (Tenaza et al., 1988). This call may be the loud contact call described by Davison (1982). Infants will emit a variation on this call when lost and searching for their mothers (Happel and Cheek, 1986).

whine: This call is a loud and wavering "waaaaa" sounding call (Tenaza et al., 1988). The mean duration of this call is 4.2 seconds (Tenaza et al., 1988). This call occurs during male-female vocal interactions, grooming, and eating (Tenaza et al., 1988). This call is emitted by both males and females, but is emitted more frequently by males (Tenaza et al., 1988). Male whines can occur without any body movement or no visible sign that the call is being made (Tenaza et al., 1988).

soft coo: This call is emitted by the adult male as he "stalks" the female during a mounting approach (Clarke, 1991). The adult female responds by presenting to the adult male (Clarke, 1991).

o-o: This call is given by the adult male when he is mounting an adult female during sexual encounters (Ren et al., 1995).

OLFACTORY COMMUNICATION:

VISUAL COMMUNICATION:
open mouth: This is when an individual opens the mouth wide which may include the retracting of the lips and baring of the teeth (Ren et al., 1991). This behavior is seen in reconciliation (Ren et al., 1991). This behavior is seen as a response to sexual solicitation by the adult female towards the adult male (Ren et al., 1995). This is described as a friendly behavior of the golden snub-nosed langur (Ren et al., 1995).

mild threat: This is when an adult male will cup an infant under the chin with the hand and administer a light slap to the face (Rapaport and Mellen, 1990). This serves to communicate a mild threat on the part of the performer of this behavior (Rapaport and Mellen, 1990).

grimace: This is where an individual will open the mouth exposing the unclenched teeth, with the corners of the mouth drawn back (Davison, 1982). Whether the lips are drawn back from the teeth depends on the intensity of the behavior (Davison, 1982). This behavior is performed by the male and sometimes by the female (Davison, 1982). This behavior serves to communicate aggression (Davison, 1982).

crouch: This is when an individual is in a sitting position with the shoulders drooping, the head hung low, the eyes staring at the ground, and the tail hanging down (Ren et al., 1991). The head during this display is lowered in a conspicuous movement (Ren et al., 1991). This behavior serves to communicate submission (Ren et al., 1991). This behavior is seen during reconciliation after a fight episode (Ren et al., 1991). The female will use this behavior to solicit copulations from the adult male, and the anogenital region may be shown towards the receiver in this case (Ren et al., 1995).

stalking: This is when an adult will approach the adult female in a slow walking manner while staring at the female and alternating open mouth and soft coo (Clarke, 1991). This behavior serves to communicate sexual solicitation on the part of the adult male (Clarke, 1991). The female responds to this behavior by presenting, and the adult female will start presenting until the adult male performs this behavior (Clarke, 1991).

pass by: This is when the adult female will approach the adult male from a far distance (about five meters) and pass by (at about one meter) the adult male without pausing or stopping (Clarke, 1991). This behavior communicates sexual solicitation on the part of the adult female (Clarke, 1991).

arboreal display rush: This behavior is performed by the male (Davison, 1982). This is where a male will leap from horizontal support to horizontal support in a zigzag fashion, which is accompanied by extreme flexure and extension of the limbs at each support leaped on to by the male (Davison, 1982). No call is heard when this display is performed (Davison, 1982). This behavior is similar to other branch-leaping type displays by other colobines (Davison, 1982).

TACTILE COMMUNICATION:
social grooming: This is when one individual grooms another and is used to reinforce the bonds between individuals. This behavior is seen during reconciliation after a fight episode (Ren et al., 1991).

clasping with head contact: This is described as when "an adult pulls the infant to its body and presses the chin, mouth, forehead, or nose against the infant" (Rapaport and Mellen, 1990). Both adult males and females perform this behavior after being reunited with the infant after separation (Rapaport and Mellen, 1990). This behavior may be performed top calm the infant (Rapaport and Mellen, 1990).

embrace: This is when one individual will place both arms around a conspecific (Ren et al., 1991). This behavior is seen during reconciliation after a fight episode (Ren et al., 1991).

solicit embrace: This is where an individual will walk up to a conspecific with the head lowered and butt the conspecific on the side, forearm, or chest with the crown of the head (Davison, 1982). This behavior is performed to solicit embrace (Davison, 1982).

contact sit: This is when one individual will sit huddle in close body contact to another individual (Ren et al., 1991). This behavior is seen during reconciliation after a fight episode (Ren et al., 1991).

hold-hand: This is when one individual will reach out to a conspecific and hold the other's hand (Ren et al., 1991). This behavior is seen during reconciliation after a fight episode (Ren et al., 1991).

hold-lumbar: This is when an individual will grasp a conspecific from behind with both hands at hip level (lumbar vertebrae) while sitting in a position where the front faces the conspecific's back (Ren et al., 1991). This behavior is seen during reconciliation after a fight episode (Ren et al., 1991).

REPRODUCTION:
The golden snub-nosed langur gives birth to a single offspring (Qi, 1986). Births for this species are concentrated in the warm spring, March to June, and this may be because of a reduction of the exposure to cold by the infant and a greater amount of high quality food for the lactating mother (Gao and Liu, 1995; Qi, 1986; Zhang et al., 2000). Golden snub-nosed langurs become sexually mature at the age of 6 years (Qi, 1986). Davison (1982) found that in captivity females mature at 3 or 4 years and males at 4 years old. Liang et al. (2000) found that males begin to copulate at 3.8 to 4.2 years and ejaculation begins at 6.3 to 6.7 years of age. The gestation period ranges from 193.5 to 203 days (Qi, 1986; Zhang et al., 2000). Births for this species usually take place at night (Qi, 1986).

Sexual solicitations and mating behavior was found to occur from September to December, suggesting that this species is a seasonal breeder (Zhu et al., 1994; Zhang et al., 2000; Davison, 1982). Mounting by the male is first preceded by the adult male approaching the female in a slow walking manner described as "stalking" by one author while staring at the female and alternating open mouth with soft coo (Clarke, 1991). The female will respond to this behavior by crouching the body with the tail raised and the ventrum pressed against the substrate, prostration (Clarke, 1991; Ren et al., 1995). The female is found only to start presenting behavioral patterns after the male begins "stalking" (Clarke, 1991). The duration of mounts are generally one minute or less and are accompanied by pelvic thrusts that can number anywhere from 3-47 per mount (Clarke, 1991). Mounts by the male consists of the male mounting from the rear, holding the female around the midsection with both hands, holding one of the female's rear legs with a foot, sometimes uttering o-o vocalization, and having the head tilted down to one side (Ren et al., 1995). During copulation the pair often looks at each other (Ren et al., 1995). Mounts are terminated by the male dismounting (Clarke, 1991). The pair will often perform social grooming at the termination of copulation (Ren et al., 1995). Ejaculations may or may not occur depending on the season, mating occurs year round for this species but ejaculations are only found to occur during the mating season (Clarke, 1991). In an experiment by Ren et al. (1995), ejaculations were only found to occur from the period of October to January, where mountings did occur during other times of the year but without ejaculations. Males have been also found to solicit mounting by performing open mouth display towards the female, then pushing the bottom of the female with hand, with the result being the female allows the male to mount (Ren et al., 1995). Females have also been found to mount males, which are preceded by males presenting towards the females (Clarke, 1991). Many times female mounts occur after the male starts his "stalking" behavior towards the female, and instead of the male mounting the female, the reverse occurs with the female mounting the male thrusting against his hindquarters (Clarke, 1991). In other instances of female mounting the female will mount right after the male completes a mounting episode (Clarke, 1991). Females are also found to solicit mounting from the male by approaching the male from some distance and then passing by the male closely without pausing or stopping (Clarke, 1991). Ren et al. (1995) found that females solicited males far more than males solicited females, and female solicitation starts with the female making eye contact with the male then running a short distance away, where the male responds with open mouth and the female counters with the crouch display.

Adult females have been found to interfere with other adult females when they are attempting to copulate with an adult male (Ren et al., 1995). Interruptive behavior consists of: a female soliciting a male who is already being solicited by another, a female mounts the female who is soliciting a male, or a group of females show threat behavior towards a female who is soliciting a male in the form of stares and kuku vocalizations with closed mouths (Ren et al., 1995). Males will respond to interfering females by: mounting the female doing the original solicitation, mounting the interfering female, mounting both females (Ren et al., 1995).

Males have been found to masturbate, and when this occurs the male will make facial and vocal displays towards the female (Clarke, 1991). During the one observed case of ejaculation during masturbation, both the male and the female were found to consume the ejaculate (Clarke, 1991).

REFERENCES:
Allen, G.M. 1938. The Mammals of China and Mongolia. American Museum of Natural History: New York.

Ankel-Simons, F. 2000. Primate Anatomy. Academic Press: San Diego.

Burton, F. 1995. The Multimedia Guide to the Non-human Primates. Prentice-Hall Canada Inc.

Clarke, A.S. 1991. Sociosexual behavior of captive Sichuan golden monkeys (Rhinopithecus roxellana). Zoo Biology. Vol. 10, 369-374.

Davison, G.W.H. 1982. Convergence with terrestrial cercopithecines by the monkey Rhinopithecus roxellanae. Folia Primatologica. Vol. 37, 209-215.

Fleagle, J. G. 1988. Primate Adaptation and Evolution. Academic Press.

Gao, Y. and Liu, S. 1995. Physiological adaptations of golden monkeys to high altitude habitat in the Qinling Mountains. Asian Primates. Vol. 5, 17-19.

Happel, R. and Cheek, T. 1986. Evolutionary biology and ecology of Rhinopithecus. in Current Perspectives in Primate Social Dynamics. eds. D.M. Taub and F.A. King. Van Nostrand Reinhold Co.: New York.

Hu, J.C. 1998. Rhinopithecus roxellanae. in China Red Data Book of Endangered Animals: Mammalia. ed. S. Wang. Science Press: Beijing.

Jablonski, N.G. and Ruliang, P. 1995. Sexual dimorphism in the snub-nosed langurs (Colobinae: Rhinopithecus). American Journal of Physical Anthropology. Vol. 96, 251-272.

Kirkpatrick, R.C. 1998. Ecology and behavior in snub-nosed and douc langurs. in The Natural History of the Doucs and Snub-nosed Monkeys. ed. N.G. Jablonski. World Scientific Publishing: Singapore.

Kirkpatrick, R.C., Gu, H.J., and Zhou, X.P. 1998. A preliminary report on feeding ecology in Rhinopithecus roxellana. XVII Congress of the International Primatological Society. University of Antananarivo, Madagascar, 262.

Kirkpatrick, R.C., Gu, H.J., and Zhou, X.P. 1999. A preliminary report on Sichuan snub-nosed monkeys (Rhinopithecus roxellana) at Baihe Nature Reserve. Folia Primatologica. Vol. 70, 117-120.

Li, B., Chen, C., Ji, W., and Ren, B. 2000. Seasonal home range changes of the Sichuan snub-nosed monkey (Rhinopithecus roxellana) in the Qinling Mountains of China. Folia Primatologica. Vol. 71, 375-386.

Li, B., Ren, B., and Gao, Y. 1999. A change in the summer home range of Sichuan snub-nosed monkeys in Yuhuangmiao, Qinling Mountains. Folia Primatologica. Vol. 70, 269-273.

Li, Y.M. 2001. The seasonal diet of the Sichuan snub-nosed monkey (Rhinopithecus roxellana) in Shennongjia Nature Reserve, China. Folia Primatologica. Vol. 72, 40-43.

Li, Y., Stanford, C.B., and Yang, Y. 2002. Winter feeding tree choice in Sichuan snub-nosed monkeys (Rhinopithecus roxellanae) in Shennongjia Nature Reserve, China. International Journal of Primatology. Vol. 23(3), 657-675.

Liang, B., Zhang, S., and Wang, L. 2000. Development of sexual morphology, physiology and behaviour in Sichuan golden monkeys, Rhinopithecus roxellana. Vol. 71, 413-416.

Qi, J.F. 1986. The Chinese golden monkey- Husbandry and reproduction. in Primates: The Road to Self-Sustaining Populations. ed. K. Benirschke. Springer-Verlag: New York.

Rapaport, L. and Mellen, J. 1987. Infant development and parental care in a family group of captive golden monkeys (Rhinopithecus roxellanae). (abstract) American Journal of Primatology. Vol.12(3), 366.

Rapaport, L. and Mellen, J.D. 1990. Parental care and infant development in a family group of captive Sichuan golden monkeys (Rhinopithecus roxellanae): First 20 days. Primates. Vol. 31(1), 129-135.

Ren, B., Zhang, S., Wang, L., Liang, B., and Li, B. 2001. Vertical distribution of different age-sex classes in a foraging group of Sichuan golden monkeys (Rhinopithecus roxellana). Folia Primatologica. Vol. 72, 96-99.

Ren, R.M., Yan, K.H., Su, Y.J., Qi, H.J., Liang, B., Bao, W.Y., and de Waal, F.B.M. 1991. Reconciliation behavior of golden monkeys (Rhinopithecus roxellanae roxellanae) in small breeding groups. Primates. Vol. 32(3), 321-327.

Ren, R.M., Yan, K.H., Su, Y.J., Qi, H.J., Liang, B., Bao, W.Y., and de Waal, F.B.M. 1992. Reconciliation behavior of golden monkeys. XIVth Congress of the International Primatological Society, Strasbourg, France.

Ren, R.M., Yan, K.H., Su, Y.J., Qi, H.J., Liang, B., Bao, W.Y., and de Waal, F.B.M. 1995. The reproductive behavior of golden monkeys in captivity (Rhinopithecus roxellana roxellana). Primates. Vol. 36(1), 135-143.

Ren, R.M., Kirkpatrick, R. C., Jablonski, N.G., Bleisch, W.V., and Le, X.C. 1996/1997. Conservation status and prospects for the snub-nosed langurs (Colobinae: Rhinopithecus). Primate Conservation. Vol. 17, 152-159.

Ren, R. Su, Y., Yan, K., Li, J., Zhou, Y., Zhu, Z., Hu, Z., and Hu, Y. 1998. Preliminary survey of the social organization of Rhinopithecus [Rhinopithecus] roxellana in Shennongjia National Natural Reserve, Hubei, China. in The Natural History of the Doucs and Snub-nosed Monkeys. ed. N.G. Jablonski. World Scientific Publishing: Singapore.

Su, Y., Ren, R., Yan, K., Li, J., Zhou, Y., Zhu, Z., Hu, Z., and Hu, Y. 1998. Preliminary survey of the home range and ranging behavior of golden monkeys (Rhinopithecus [Rhinopithecus] roxellana) in Shennongjia National Natural Reserve, Hubei, China. in The Natural History of the Doucs and Snub-nosed Monkeys. ed. N.G. Jablonski. World Scientific Publishing: Singapore.

Tenaza, R.R., Fitch, H.M., and Lindburg, D.G. 1988. Vocal behavior of captive Sichuan golden monkeys (Rhinopithecus r. roxellana). American Journal of Primatology. Vol. 14, 1-9.

de Waal, F.B.M. and van Roosmalen, A. 1979. Reconciliation and consolation among chimpanzees. Behavioral Ecology and Sociobiology. Vol. 5, 55-66.

Wang, Y., Jiang, X., and Li, D. 1998. Classification and distribution of the extant subspecies of golden snub-nosed monkey (Rhinopithecus [Rhinopithecus] roxellana). in The Natural History of the Doucs and Snub-nosed Monkeys. ed. N.G. Jablonski. World Scientific Publishing: Singapore.

Zhang, S., Liang, B., and Wang, L. 1999a. Infanticide within captive groups of Sichuan golden snub-nosed monkeys (Rhinopithecus roxellana). Folia Primatologica. Vol. 70, 274-276.

Zhang, S., Liang, B., and Wang, L. 2000. Seasonality of matings and births in captive Sichuan golden monkeys (Rhinopithecus roxellana). American Journal of Primatology. Vol. 51, 265-269.

Zhang, S., Ren, B., and Li, B. 1999. A juvenile Sichuan golden monkey (Rhinopithecus roxellana) predated by a goshawk (Accipiter gentilis) in the Qinling Mountains. Folia Primatologica. Vol. 70, 175-176.

Zhu, Z., Ren, R., Su, Y., and Hu, Z. 1994. Field observations of the reproductive ecology and behavior of golden monkeys (Rhinopithecus roxellana) in Shennongjia National Natural Reserve. (abstract) XVth Congress of the International Primatological Society.

Last Updated: October 12, 2003.
[The Primata] [Primate Fact Sheets] [Subfamily Colobinae] [Pygathrix Links]