Biological Exuberance: Animal Homosexuality and Natural Diversity

York: Crown Publishers.

Wells, S., and L. F. Baptista (1979) “Displays and Morphology of an Anna X Allen Hummingbird Hybrid.” Wilson Bulletin 91:524–32.

Wells, S., L. E Baptista, S. F. Bailey, and H. M. Horblit (1996) “Age and Sex Determination in Anna’s Hummingbird by Means of Tail Pattern.” Western Birds 27:204–6.

Wells, S., R. A. Bradley, and L. E Baptista (1978) “Hybridization in Calypte Hummingbirds.” Auk 95:537–49.

Williamson, E S. L. (1956) “The Molt and Testis Cycle of the Anna Hummingbird.” Condor 58:342–66.

BLACK-RUMPED FLAMEBACK

IDENTIFICATION: A medium-sized, red-crested woodpecker with a golden back, black rump, and black-and-white patterning on the face and neck. DISTRIBUTION: India, Pakistan, Sri Lanka. HABITAT: Woodland, scrub, gardens. STUDY AREA: Near Chittur, India; subspecies D.b. puncticolle.

ACORN WOODPECKER

IDENTIFICATION: A red-capped woodpecker with a striking black-and-white face, black upperparts, white underparts, and a black breast band. DISTRIBUTION: Pacific and southwest United States, Mexico through Colombia. HABITAT: Oak and pine woodland. STUDY AREAS: Hastings Natural History Reservation (Monterey) and near Los Altos, California.

Social Organization

Acorn Woodpeckers have an extraordinarily varied and complex social organization. In many populations, birds live in communal family groups containing up to 15 individuals—typically there are as many as 4 breeding males and 3 breeding females in such groups (though nonbreeding groups also occur—see below). The remaining birds in a group are nonbreeding “helpers” that may share in the parenting duties. Within groups, the mating system is known as POLYGYNANDRY, that is, each male mates and bonds with several females and vice versa. In other populations, monogamous pairs as well as other variations on polygamy occur. Little is known about the social organization of Black-rumped Flamebacks, although it is thought that they form monogamous mated pairs.

Description

Behavioral Expression: Male Black-rumped Flamebacks sometimes copulate with each other. One male mounts the back of the other (as in heterosexual mating), bending his tail down and thrusting it under the belly of the other male to make cloacal (genital) contact. Reciprocal mounting may occur, in which a male copulates with a male that has just mounted him. A bird involved in same-sex mounting activity may adopt a distinctive posture, in which his body is perpendicular to the branch he is perching on and his wing tips are arched toward the ground and hanging below his feet. Males also sometimes drum against a tree trunk prior to homosexual mounting.

Acorn Woodpeckers participate in a fascinating group display that involves ritualized sexual and courtship behavior, including homosexual mounting. At dusk, the members of a group gather together prior to roosting in their tree holes. As more and more birds arrive, they begin mounting each other in all combinations—males mount females and other males, females mount males and other females, young Woodpeckers mount older ones and vice versa. The mounting behavior resembles heterosexual mating, except it is usually briefer and cloacal contact is generally not involved (although genital contact does sometimes occur). Reciprocal mountings are common, and sometimes two Woodpeckers will try to mount the same bird simultaneously. Following the display, group members fly off to their roost holes to sleep. Ritualized mounting may also occur at dawn when the birds emerge from their roost holes. Because many group members are related to each other, at least some of this mounting is incestuous. Female Acorn Woodpeckers often coparent together, both laying eggs in the same nest cavity. Such “joint nesters” are often related (mother and daughter, or sisters), but sometimes two unrelated females nest and parent together as well. Joint-nesting females may continue to associate even if they happen not to breed in a particular year.

Frequency: Homosexual behavior in Black-rumped Flamebacks probably occurs only occasionally; however, heterosexual mating has never been observed in this species in the wild, so much remains to be learned about the behavior of this Woodpecker. The mounting display of Acorn Woodpeckers—including homosexual mounting—is a regular feature of the social life of this species at all times of the year, occurring daily in most groups. More than a third of all female Acorn Woodpeckers nest jointly, and about a quarter of all groups have joint-nesting females; 14 percent of these joint nests involve unrelated females as coparents.

Orientation: To the extent that they mount both males and females in the group display, Acorn Woodpeckers are bisexual (although it must be remembered that such mounting is often ritualized, i.e., it may not always involve genital contact). Not enough is known about the life histories of individual Black-rumped Flamebacks to make any generalizations about their sexual orientation.

Nonreproductive and Alternative Heterosexualities

As described above, Acorn Woodpeckers have an unusual communal family organization that can involve different forms of polygamy. In addition, many birds are nonbreeding: more than a third of all groups may not reproduce in a given year, and one-quarter to one-half of all adult birds do not procreate. In some populations the proportion of nonbreeders may be as high as 85 percent. Many of these are birds who remain with their family group for several years after they become sexually mature, helping their parents raise young; some delay reproducing for three or four years. Other nonbreeders (as many as one-quarter) do not in any way help to raise young. Some groups are nonreproductive because all their adult members are of the same sex: nearly 15 percent of nonbreeding groups have no adult females and nearly 4 percent have no adult males. In addition to the nonprocreative heterosexual behaviors mentioned above (REVERSE mounting, group sexual activity, mounting without genital contact), female Acorn Woodpeckers also sometimes copulate with more than one male in quick succession. About 3 percent of families contain offspring that result from promiscuous matings with males outside the group. Incestuous heterosexual matings occasionally occur as well, although they seem to be avoided—in fact, incest avoidance may lead to a group’s forgoing breeding for an extended time. Parenting in this species is notable for a variety of counterreproductive and violent behaviors. Egg destruction is common—particularly among joint- nesting females, who often break (and eat) each other’s and their own eggs until they begin laying synchronously. Males also sometimes destroy eggs of their own group. In addition, infanticide and cannibalism occur regularly in Acorn Woodpeckers. A common pattern seems to be for a new bird in a group—often a female—to peck the nestlings to death and eat some of them in order to breed with the other adults in the group. Parents also regularly starve any chicks that hatch later than a day after the others do.

Sources

*asterisked references discuss homosexuality/transgender

*Koenig, W. D. (1995–96) Personal communication.

Koenig, W. D., and R. L. Mumme (1987) Population Ecology of the Cooperatively Breeding Acorn Woodpecker. Princeton, N.J.: Princeton University Press.

Koenig, W. D., R. L. Mumme, M. T. Stanback, and F. A. Pitelka (1995) “Patterns and Consequences of Egg Destruction Among Joint-Nesting Acorn Woodpeckers.” Animal Behavior 50:607–21.

Koenig, W. D., and P. B. Stacey (1990) “Acorn Woodpeckers: Group-Living and Food Storage Under Contrasting Ecological Conditions.” In P. B. Stacey and W. D. Koenig, eds., Cooperative Breeding in Birds: Long-Term Studies of Ecology and Behavior, pp. 415–53. Cambridge: Cambridge University Press.

Koenig, W. D., and F. A. Pitelka (1979) “Relatedness and Inbreeding Avoidance: Counterploys in the