Biological Exuberance: Animal Homosexuality and Natural Diversity

(Gray, 1844)” and “Australian Sea Lion, Neophoca cinerea (Peron, 1816).” In S. H. Ridgway and R. J. Harrison, eds., Handbook of Marine Mammals, vol. 1, pp. 25–38, 99– 118. London: Academic Press.

Wilson, G. J. (1979) “Hooker’s Sea Lions in Southern New Zealand.” New Zealand Journal of Marine and Freshwater Research 13:373–75.

York, A. E., and V. B. Scheffer (1997) “Timing of Implantation in the Northern Fur Seal, Callorhinus ursinus.” Journal of Mammalogy 78:675–83.

IDENTIfICATION: A huge pinniped (up to 12 feet and 3,500 pounds in males) with sparsely furred, brownish orange skin, a bristled “mustache,” and prominent tusks in adult males. DISTRIBUTION: Throughout the

Arctic. HABITAT: Shallow waters, coastal areas, ice floes. STUDY AREAS: Round Island, Bristol Bay, Alaska; Coats Island, Hudson Bay; Bathurst and Dundas Islands, Northwest Territories, Canada; New York Aquarium; subspecies O.r. divergens, the Pacific Walrus, and O.r. rosmarus, the Atlantic Walrus.

Social Organization

From January to March (the breeding season), Walruses congregate far from shore on and around pack ice, where heterosexual courtship and mating take place. The mating system is polygynous—males generally copulate with several different females without forming long-lasting heterosexual bonds. During the summer and early fall, males gather together in large aggregations numbering in the thousands, typically on islands that are used for this purpose year after year. These “haul-outs” are sex-segregated. When both males and females are present, they tend to occupy distinct areas; more typically, however, the haul-outs are entirely male, since females and their young migrate to the far north to spend the summer.

Description

Behavioral Expression: In the shallow waters off the coast of the summer haul-out grounds, male Walruses engage in homosexual courtship, sexual, and affectionate activities. Pairs of males—sometimes as many as 50 animals at a time—float at the surface of the water by inflating special pouches in their throat, which act like the buoyant sacs in a life vest. While floating and swimming, the males— especially younger ones—rub their bodies against one another, clasp and embrace each other with their front flippers, touch noses, and loll together in groups. Males sometimes even sleep together in the water—pairs or groups of males float vertically at the surface (a posture known as BOTTLING), one behind the other, each male clasping the one in front of him in a “sleeping line.” Males also perform courtship displays for other males, employing a number of extraordinary behaviors and sounds that are also used in heterosexual courtship. Typically a younger male displays to an older one, and courting males often situate themselves in the water near a favorite cliff face, boulder, or rock formation at the water’s edge. The spectacular display consists of inflation of the throat pouches—often preceded by head bowing—interspersed with dives by one or both males, and an incredible series of vocalizations that form a courtship “song.” In homosexual encounters, at least three types of calls are used: KNOCKS, which are rapid, clicklike sounds resembling castanets, produced by “chattering” the cheek teeth underwater; a metallic BELL call, which is an eerie gonglike sound thought to be produced underwater by striking the throat pouches with the flippers to generate air pulses; and a short, piercing WHISTLE made through pursed lips when the Walrus resurfaces.

During same-sex courtship, sometimes one male rubs his erect, arm-sized penis with his front flipper. Overt sexual behavior between males takes the form of mounting (in the shallow water): one male clasps another with his flippers from behind, thrusting his pelvis and erect penis against the other male’s anal region. Younger males mount older ones and vice versa. Although most homosexual behavior is confined to the summer haul-outs, younger males sometimes also mount adults or other younger males during the breeding season. Groups of younger Walruses may crowd around an older male and roll on top of him; in addition, an adult male occasionally sings his courtship songs to a group of younger males or is accompanied by a younger attendant male while he sings. Often, his companion surfaces and dives in synchrony with him. This behavior, known as SHADOWING, occurs regardless of whether the adult is courting a female. In addition, male Walruses in captivity have been observed participating in cross-species homosexual encounters with male Gray Seals.

A male Walrus mounting another male off the coast of Round Island (Alaska), thrusting his erect penis (visible below the surface of the water) in the other male’s anal region.

Frequency: Male Walruses engage in homosexual activities frequently during the summer haul-outs: approximately a quarter of all social interactions of males in the shallow waters involve same-sex affectionate, courtship, or sexual behaviors. On average, each male participates in such activity roughly five times per hour when in the water, and contact between males (including sexual and courtship behaviors) occupies about 3 percent of the overall time spent by males in the water. Up to 2 percent of the total male population (which may number more than 3,000) may be in the water at any one time engaging in homosexual activities. During the breeding season, up to a third of mounting activity may take place between younger males or between adult and younger males, and 2-19 percent of singing males have a younger male companion “shadowing” them.

Orientation: Because male Walruses achieve sexual maturity approximately four years before they actually participate in breeding, a sizable segment of the population is probably involved exclusively in homosexuality for a portion of their lives. Roughly 40-60 percent of all sexually mature males—those between the ages of 10 and 14—do not participate in heterosexual mating, and a large proportion of these males engage in homosexual activity. Most older males are seasonally bisexual, courting and copulating with females during the mating season and participating in same-sex activities during the summer and early fall; some, however, also engage in same-sex mounting and companionships during the breeding season.

Nonreproductive and Alternative Heterosexualities

As noted above, male Walruses experience a delay in their breeding careers, since most do not begin mating heterosexually until about 40 percent of their maximum life span is over. Once they do begin procreating, often only a small percentage of males actually copulates each year, perhaps as few as a quarter of all adult males. Among females, about half of the individuals over the age of 23 are nonbreeding, experiencing a postreproductive or “menopausal” period that may last for 7 or more years. Because of their long pregnancy (15-16 months) and nursing period (two years), most breeding-age females do not reproduce every year. One reason the pregnancy is so long is because of DELAYED IMPLANTATION: after mating, the fertilized egg fails to implant in the uterus and temporarily stops developing, remaining in “suspended animation” for four to five months. This results in an even longer period that the two sexes are separated. Although their social life is characterized by extensive sex segregation (see above), Walruses do sometimes copulate outside of the mating season. Heterosexual matings have been recorded in nearly all months of the spring and fall. Because males have a distinct yearly sexual cycle— their testes are essentially dormant except during January-March—most of this sexual activity is nonreproductive. Even during the mating season, Walruses participate in a variety of nonprocreative sexual behaviors. REVERSE mountings, in which the female clasps the male from behind and mounts him, occur in the water, and younger, sexually immature individuals also mount each other. Females have also been observed sucking their partner’s penis as well as kneading it to erection with their flippers prior to copulation (in captivity). Outside of the breeding season, males masturbate by stroking their penis with a front flipper, sometimes accompanied by knock sounds or STRUM calls (the latter sounding something like the strumming of fingers on a guitar or zither). In addition, interspecies heterosexual copulations with Gray Seals have been seen in captivity.

As in many other polygamous mammals, female Walruses generally raise their young on their own, occasionally supplemented with a number of alternative parenting arrangements. Unrelated females and males may assist in the care and protection of calves, nursery groups of youngsters sometimes play together while their