(less than 1 in 200 such matings result in insemination), and during group promiscuous matings, males often mount on any part of the female’s body, including her head. In addition, about 15—30 percent of promiscuous copulations occur outside of the female’s fertile period. The same is also true for sexual activity between mated partners: copulation begins as long as four to five months before the start of egg laying, and half of all heterosexual matings in some populations occur during nonfertilizable periods. In addition, almost a quarter of pair copulations do not involve genital contact. In Common Murres—as in most other birds—females have the remarkable ability to store sperm in special ducts in their reproductive tract, allowing them to inseminate their eggs even when not directly engaging in reproductive copulations.
Other forms of nonprocreative sexuality also occur. Nonbreeding female Common Murres often solicit promiscuous matings from males, while nonbreeding pairs or those who have lost their young (which can make up as much as a third of all pairs) frequently continue to copulate throughout the season. Nonbreeding Laysan Albatross pairs also sometimes engage in copulation. Birds in this species do not reproduce until they are 6—16 years old, even though they mature at one year old and may form pairs fully two years before they actually breed. Similarly, younger Common Murres usually delay breeding until they are five years old, congregating in CLUBS on the tidal rocks beneath the breeding colonies. Such nonbreeders make up approximately 13 percent of the population; among adults, 5—10 percent of birds do not breed each year, and more than a third skip breeding for at least one season during their life. In addition, masturbatory activity—birds mount and “copulate” with clumps of grass—was recently discovered in a closely related species, the thick-billed murre (
A variety of alternative parenting arrangements are also found in these species. About 8 percent of all Common Murre chicks have “baby-sitters”—a pair of birds other than their parents who help brood (keep warm), protect, and sometimes feed the chick (even when the youngster’s parents are not away). Most such helpers are nonbreeders; others have tried but failed to breed, while some have finished raising their family or are also taking care of their own chick. In addition, pair separation and single parenting is routine in Common Murres: when a chick is old enough to leave the colony, only its father accompanies it to sea, feeding and chaperoning it for up to 12 weeks without his female partner. In Laysan Albatrosses, heterosexual parents are together at the nest for a remarkably short time—only 5—10 days out of the 230-day breeding season. Eggs are sometimes temporarily “adopted” by other birds who incubate them when the parents are away from the nest. Nonbreeding females have even been known to “join” existing pairs and regularly take turns with the parents incubating their egg. Sometimes females also lay a second egg in a stranger’s nest. Reproduction in this species is often fraught with difficulties, however. More than 20 percent of parents (both males and females) desert their nests—often when their partner fails to return for an incubation shift on time—and couples also occasionally divorce (2 percent of all pairs). Once the chicks have hatched, they are often subjected to abuse from neighboring birds, who may savagely peck, stab, bite, and occasionally even kill the youngsters if they stray too close.
Other Species
Homosexual copulations are common in another species of auk, the Razorbill (
Birkhead, T. R. (1993)
*———(1978a) “Behavioral Adaptations to High Density Nesting in the Common Guillemot
———(1978b) “Attendance Patterns of Guillemots
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Birkhead, T. R., S. D. Johnson, and D. N. Nettleship (1985) “Extra-pair Matings and Mate Guarding in the Common Murre
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Fisher, H. I. (1975) “The Relationship Between Deferred Breeding and Mortality in the Laysan Albatross.” Auk 92:433—41.
*———(1971) “The Laysan Albatross: Its Incubation, Hatching, and Associated Behaviors.”
———(1968) “The ‘Two-Egg Clutch’ in the Laysan Albatross.”
Fisher, H. I., and M. L. Fisher (1969) “The Visits of Laysan Albatrosses to the Breeding Colony.”
Fisher, M. L. (1970)
*Frings, H., and M. Frings (1961) “Some Biometric Studies on the Albatrosses of Midway Atoll.”
Gaston, T., and K. Kampp (1994) “Thick-billed Murre Masturbating on Grass Clump.”
Harris, M. P., and S. Wanless (1995) “Survival and Non-Breeding of Adult Common Guillemots
*Hatchwell, B. J. (1988) “Intraspecific Variation in Extra-pair Copulation and Mate Defence in Common Guillemots
Hudson, P. J. (1985) “Population Parameters for the Atlantic Alcidae.” In D. N. Nettleship and T. R. Birkhead, eds.,
Johnson, R. A. (1941) “Nesting Behavior of the Atlantic Murre.”
Meseth, E. H. (1975) “The Dance of the Laysan Albatross,
Rice, D. W., and K. W. Kenyon (1962) “Breeding Cycles and Behavior of Laysan and Black-footed Albatrosses.”
Tuck, L. M. (1960)
*Wagner, R. H. (1996) “Male-Male Mountings by a Sexually Monomorphic Bird: Mistaken Identity or Fighting Tactic?”
———(1991) “Evidence That Female Razorbills Control Extra-Pair Copulations.”
GREAT CORMORANT
IDENTIFICATION: A large (3 foot), black, web-footed bird with a white throat and white filamentary plumes on the nape. DISTRIBUTION: Throughout Europe, Australasia, Africa, and Atlantic North America. HABITAT: Seacoasts, lakes, rivers. STUDY AREAS: Shinobazu Pond, Tokyo, Japan; Amsterdam Zoo, the Netherlands; subspecies