*asterisked references discuss homosexuality/transgender
Clarke, A. S. (1991) “Sociosexual Behavior of Captive Sichuan Golden Monkeys (Rhinopithecus roxellana).” Zoo Biology 10:369-74.
Gorzitze, A. B. (1996) “Birth-related Behaviors in Wild Proboscis Monkeys (Nasalis larvatus).” Primates 37:75-78.
Kawabe, M., and T. Mano (1972) “Ecology and Behavior of the Wild Proboscis Monkey, Nasalis larvatus (Wurmb) in Sabah, Malaysia.” Primates 13:213-27.
Poirier, F. E., and H. Hongxhin (1983) “Macaca mulatta and Rhinopithecus in China: Preliminary Research Results.” Current Anthropology 24:387-88.
Qi, J.-F. (1988) “Observation Studies on Reproduction of Golden Monkeys in Captivity: I. Copulatory Behavior.” Acta Theriologica Sinica 8:172-75.
*Ren, R., K. Yan, Y. Su, H. Qi, B. Liang, W. Bao, and F. B. M. de Waal (1995) “The Reproductive Behavior of Golden Monkeys in Captivity.” Primates 36:135-43.
———(1991) “The Reconciliation Behavior of Golden Monkeys (Rhinopithecus roxellanae roxellanae) in Small Breeding Groups.” Primates 32:321-27.
Schaller, G. B. (1985) “China’s Golden Treasure.” International Wildlife 15:29- 31.
*Yeager, C. P. (1990a) “Notes on the Sexual Behavior of the Proboscis Monkey.” American Journal of Primatology 21:223-27.
———(1990b) “Proboscis Monkey (Nasalis larvatus) Social Organization: Group Structure.” American Journal of Primatology 20:95-106.
MACAQUES
IDENTIFICATION: A medium-sized (3-foot-long) monkey with a brownish gray coat, short tail, and red facial skin. DISTRIBUTION: Japan; endangered. HABITAT: Forests, including subalpine and snow-covered terrain. STUDY AREAS: Wild: near Arashiyama, Takasakiyama (Kyushu), Shiga Heights (Jigokundai), Koshima, Miyajima, and other areas of Japan; semiwild: near Laredo, Texas; the Oregon Regional Primate Research Center; captivity: Laboratory of Behavioral Primatology, St-Hyacinthe, Quebec; Calgary Zoo; Cavriglia Park (Italy).
Social Organization
Japanese Macaques live in cosexual troops of 20–100 individuals, subdivided into smaller matrilineal groups composed of numerous related females and several unrelated males. Males usually leave their birth group on reaching maturity and may even periodically transfer to different groups, live in all-male groups, or become solitary or peripheral. Females, in contrast, generally remain in their home group for life. As a result, the group is centered around the kinship and bonding between females (and some groups have only one regular adult male member).
Description
Behavioral Expression: Female Japanese Macaques form intense, exclusive pair-bonds with each other based on mutual sexual attraction. These pairings are known as CONSORTSHIPS and are characterized by a number of distinctive affectionate, sexual, and social activities. Female partners in a consortship typically sit together, huddling or in close physical contact, and often spend long periods grooming one another between their sexual interactions. They also synchronize their movements, including traveling in tandem and persistently following one another, and may make cooing sounds toward one another. Consorts become agitated when another animal approaches or intrudes, responding with threats, shudders, vocalizing, or withdrawal. They sometimes actively separate themselves from their relatives (temporarily forgoing the time they would usually spend grooming their young or other kin), even occasionally withdrawing from the main troop as a whole. Partners in homosexual consortships often forge strong supportive “alliances” with each other, defending their partner and intervening on her behalf when she is threatened by another individual. Most such interventions do not disrupt the traditional hierarchy or rank system of the troop. However, some interventions are termed REVOLUTIONARY because partners challenge higher-ranking individuals, while others are ARBITRATING interventions, involving mediation between individuals with more ambiguous rankings. Females also actively compete with individuals of both sexes for access to other females as sexual partners, sometimes even incurring severe injuries when they challenge intruding males. Consortships typically last from a few days to several weeks during the mating season, although partners often develop a strong friendship as a result of a consortship and remain bonded throughout the year (in contrast to heterosexual consortships, which generally do not extend beyond the mating season). Females are often serially monogamous, forming several exclusive consortships sequentially during the mating season, although they generally have fewer partners than individuals engaging in heterosexual or male homosexual activities. Homosexual pairings occur among females of all ages, from adolescents to the very old, and sometimes an adult female will pair with a pubescent female. Interestingly, an incest taboo is in effect for homosexual but not heterosexual consortships: females never choose close relatives (mothers, sisters, daughters, granddaughters, first cousins) as partners, whereas brother-sister and mother-son consorts do occasionally occur. Aunts and nieces, however, do not generally recognize each other as kin in this species, and so they sometimes consort together.
As signaled by the reddening of their faces and sexual skin (which also swells), females in consortships are usually “in heat,” and sexual activity is a regular and prominent feature of these homosexual pairings. Females engage in a variety of behaviors that involve genital stimulation, usually in the form of one female mounting another. Fully seven different positions are used for homosexual copulation (as well as for heterosexual mating, although with different frequencies). Most commonly, one female sits or lies on the back of the other, making pelvic thrusts and rubbing her clitoris against her partner’s rump. The mountee’s clitoris maybe stimulated by her partner’s thrusting or with her own tail. Two females also sometimes embrace each other in a face-to-face position and rub their genitals together, either lying down or “sitting in each other’s lap”; these postures are more common in lesbian interactions than heterosexual ones. Other positions include the “double-foot-clasp” (one female mounts from behind, grasping her partner’s ankles or thighs with her feet); rear mounts where the mounter keeps her feet on the ground; and thrusting against the partner in sideways or variable postures. During mounting, females sometimes make vocalizations such as hoarse, cackling ko-ko-ko-ko sounds. Females being mounted commonly reach back and grab their partner, gazing intensely into her eyes and grimacing, while the mounter clutches the fur on her partner’s back—indications that both females probably experience orgasm during homosexual interactions. Consorts usually mount each other in a series of three or more consecutive mounts (as in heterosexual mating), and mounting is often reciprocal (partners exchange positions). In addition to mounting and genital rubbing, consorts occasionally suck each other’s nipples. Sexual interactions are frequently initiated by the mountee, who performs a number of characteristic “courtship” behaviors to solicit or “demand” a mount from her consort. She may slap the ground while shrieking, run away from her partner and then return while presenting her hindquarters, or else display a variety of other behaviors such as head bobbing, vocalizing, arching of the back, lip quivering, intense staring, and even spasms and more aggressive pushing and grabbing. All of these are distinct from heterosexual courtship, in which the female usually invites the male to mount by slowly inching toward him in a sitting position, with mutual flipping of ears and eyebrows. A wide variety of sounds are used in both heterosexual and homosexual courtships, including cooing, whistling, warbling, squawking, chirping, barking, and squeaking—the latter is more typical of interactions between females, however.
Male Japanese Macaques also mount each other. They do not generally form homosexual consortships— indeed, some males mount up to 24 different partners—but some individuals do have certain preferred partners, usually of the same age. Males also sometimes engage in affectionate and playful activities in combination with their sexual interactions, including touching, huddling, and grooming. Unlike heterosexual and lesbian interactions, most mounts between males are single (rather than series) and are generally briefer, although they can still involve full erections, pelvic thrusting, penetration, and/or ejaculation. The double-foot-clasp is the preferred posture
