Frederick, H., and C. N. Johnson (1996) “Social Organization in the Rufous Bettong,
Ganslosser, U. (1993) “Stages in Formation of Social Relationships—an Experimental Investigation in Kangaroos (Macropodoidea: Mammalia).”
*———(1979) “Soziale Kommunikation, Gruppenleben, Spiel- und Jugendverhalten des Doria-Baumkanguruhs
*Ganslosser, U., and C. Fuchs (1988) “Some Quantitative Data on Social Behavior of Rufous Rat-Kangaroos
George, G. G. (1977) “Up a Tree with Kangaroos.”
*Hutchins, M., G. M. Smith, D. C. Mead, S. Elbin, and J. Steenberg (1991) “Social Behavior of Matschie’s Tree Kangaroo
Jarman, P. J. (1991) “Social Behavior and Organization in the Macropodoidea.”
*Johnson, P. M. (1980) “Observations of the Behavior of the Rufous Rat-Kangaroo,
IDENTIFICATION: A bearlike marsupial with woolly brown or gray fur, large black nose, white chest, and long claws. DISTRIBUTION: Eastern and southeastern Australia. HABITAT: Eucalyptus forests. STUDY AREAS: Lone Pine Sanctuary, Brisbane, Australia; San Diego Zoo; subspecies
Social Organization
Koalas are largely nocturnal and solitary, although in some populations they tend to live in scattered clusters of two to six females with several males. The mating system is probably promiscuous or polygamous (animals mate with multiple partners), and males take no part in raising their young.
Description
Nonreproductive and Alternative Heterosexualities
Heterosexual relations in Koalas are marked by a striking amount of aggression and violence: more than two-thirds of fights are between males and females rather than between males. Females are sometimes “pestered” by males that persistently follow, touch, bite, or snap at them; if the female returns the bites, the encounter can escalate into a severe fight. Males have been known to brutally attack females—including pregnant and nursing mothers—knocking them from the trees and savagely mauling them. In fact, it is typical for males to nip females on the neck during mating, and for heterosexual copulations to end with the male attacking the female. Females also fight with males (though less violently), and aggressiveness toward males is considered to be a defining feature of estrus for female Koalas. Occasionally adults are also abusive toward babies: mothers sometimes bite their young, while males have been observed attacking infants that interrupt them during a mating with their mother. Many heterosexual interactions are nonprocreative, since males often try to mount females who are not in heat. Although the females typically rebuff their advances, in some cases the males are able to mount them, often thrusting against the female and ejaculating on her without any penetration. Females in heat also sometimes mount males (REVERSE mounts).
Many wild populations of Koalas have particularly high rates of female infertility (and significantly reduced reproductive rates) due to venereal disease. More than half of all females in some areas are infected with genital chlamydia, a bacteria that causes a number of reproductive tract diseases and, ultimately, sterility. This pathogen has apparently been present in Koala populations for a relatively long time, as records of the associated diseases date back to at least the 1890s. Although the exact mode of its transmission is not yet fully understood, two routes have been implicated: sexual and mother-to-young. The latter may be due to the infant Koala’s habit of eating its mother’s feces directly from her anus during weaning, since she produces a special form of excrement known as PAP especially for feeding her young (this practice is also found in a number of other marsupials).
Other Species
In another marsupial, the Common Brushtail Possum
Brown, A. S., A. A. Girjes, M. F. Lavin, P. Timms, and J. B. Woolcock (1987) “Chlamydial Disease in Koalas.”
*Gilmore, D. P. (1965) “Gynandromorphism in
Lee, A., and R. Martin (1988)
Phillips, K. (1994)
*Sharman, G. B., R. L. Hughes, and D. W. Cooper (1990) “The Chromosomal Basis of Sex Differentiation in Marsupials.”
Smith, M. (1980a) “Behavior of the Koala,
*———(1980b) “Behavior of the Koala,