matriarchal groups. Furthermore, in both African and Asiatic species males experience a periodic sexual cycle known as MUSTH. A male in musth exhibits a number of characteristic physiological and behavioral changes, including increased aggression, ear waving and head swinging, infrasonic rumbling calls, continuous urinal discharge, and secretion by the temporal glands (located on either side of the head). Musth can last a few days to several months; during this time, males tend to associate more with females, but once musth is over, they usually return to male-only groups. Among African Elephants, bulls frequent special areas during their nonmusth time where their interactions are exclusively with other males, and they sometimes form a stronger association with one or two particular bulls.
During heterosexual interactions, females may be overtly aggressive toward males. Female Asiatic Elephants, for example, often charge and chase males (especially younger ones) that are trying to mate with them. In addition, males sometimes try to mount juvenile animals, who squeal in protest until a female intervenes. In fact, it is thought that female aggressiveness toward males contributes to a significant delay in breeding in this species: although males are sexually mature when about 10 years old, most do not begin breeding until about 17 years old. Reproductive suppression also occurs in African Elephants: the onset of puberty in females may be delayed by up to 10 years in some populations as a result of social, nutritional, or physiological stresses. In addition, most males do not father their first offspring until they are 30—35 years old—this is 15—20 years after they become sexually mature. Other nonbreeding individuals include loner males (often older, postreproductive individuals), males in “companionships” (described above), as well as postreproductive or menopausal females (generally older than 50 years), and females who have nonbreeding intervals of up to 13 years between calves. Heterosexual relations are further complicated by an apparent mismatch in the structure of the male and female genitalia: unlike in most mammals, the female’s vaginal opening is placed far forward on her belly, hampering access by the male. Although his penis is able to assume a special S-shape to reach her vulva, penetration is often difficult to achieve. On occasion, the male’s organ will contact the female’s anus rather than her vulva, and he may ejaculate before achieving penetration. Heterosexual relations also often include touching and stimulation of the genitals with the trunk; in captivity, one female was also observed rubbing her clitoris against a male’s side while he was lying down.
Other Species
Homosexual mounting in the Indian Rhinoceros
asterisked references discuss homosexuality/transgender
*Buss, I. O. (1990)
Buss, I. 0,. and N. S. Smith (1966) “Observations on Reproduction and Breeding Behavior of the African Elephant.”
*Dixon, A., and M. MacNamara (1981) “Observations on the Social Interactions and Development of Sexual Behavior in Three Sub-adult, One-horned Indian Rhinoceros
Douglas-Hamilton, I., and O. Douglas-Hamilton (1975)
Eisenberg, J. F., G. M. McKay, and M. R. Jainudeen (1971) “Reproductive Behavior of the Asiatic Elephant
*Grzimek, B. (1990) “African Elephant.” In
*Jayewardene, J. (1994)
*Kuhme, W. (1962) “Ethology of the African Elephant
Laws, R. M. (1969) “Aspects of Reproduction in the African Elephant,
Lee, P. C. (1987) “Allomothering Among African Elephants.”
*McKay, G. M. (1973)
*Morris, D. (1964) “The Response of Animals to a Restricted Environment.”
Moss, C. (1988)
Moss, C., and J. H. Poole (1983) “Relationships and Social Structure of African Elephants.” In R. A. Hinde, ed.,
Poole, J. H. (1994) “Sex Differences in the Behavior of African Elephants.” In R. V. Short and E. Balaban, eds.,
———(1987) “Rutting Behavior in African Elephants: The Phenomenon of Musth.”
*Poole, T. B., V. J. Taylor, S. B. U. Fernando, W. D. Ratnasooriya, A. Ratnayeke, G. Lincoln, A. McNeilly, and A. M. V. R. Manatunga (1997) “Social Behavior and Breeding Physiology of a Group of Asian Elephants
*Ramachandran, K. K. (1984) “Observations on Unusual Sexual Behavior in Elephants.”
*Rosse, I. C. (1892) “Sexual Hypochondriasis and Perversion of the Genetic Instinct.”
*Shelton, D. J. (1965) “Some Observations on Elephants.”
*Sikes, S. K. (1971)
Other Mammals
CARNIVORES
LION
IDENTIFICATION: A large wild cat (up to 550 pounds) with a prominent mane in males. DISTRIBUTION: Throughout Africa and in Gujarat, northwest India; vulnerable. HABITAT: Plains, savannas, scrub, open forest. STUDY AREAS: Serengeti National Park, Tanzania; Gir Wildlife Sanctuary, India; Gay Lion Farm, California; subspecies
CHEETAH
IDENTIFICATION: A medium-sized wild cat with a sleek, greyhoundlike physique and a spotted coat DISTIBUTION-TION: Throughout Africa and sporadically in central Asia and the Middle East; vulnerable. HABITAT: Semidesert, grassland, steppes. STUDY AREAS: Serengeti National Park, Tanzania; Lion Country Safari, California; National Zoological Park, Washington, D.C.; Hogle Zoo, Utah; subspecies