Stumptail, and Pig-tailed Macaques being mounted; erection and masturbation by male mountee Rhesus, Pig-tailed, and Crested Black Macaques; thrusting by male Bonobos being mounted; and stimulation of the mountee’s clitoris by her partner’s thrusting in Hanuman Langurs and Japanese Macaques. In addition to direct and indirect genital stimulation during mounting, it is quite likely that male animals being penetrated during anal intercourse also experience stimulation of the prostate gland (which presses against the wall of the rectum). In human males, direct stimulation of the prostate—for instance, during anal intercourse—can be highly arousing and may precipitate or enhance orgasm. A similar capacity is probably present in all male mammals. Although direct evidence (in the form of firsthand accounts) of the pleasurable or arousing nature of this activity is, of course, lacking in nonhuman animals, there is some indirect evidence. A standard technique of inducing erection and ejaculation (for purposes of artificial insemination) in male mammals is through anal and/or prostate stimulation. Known as electroejaculation, this technique involves insertion of an anal probe and stimulation of the rectum—especially in the area of the prostate gland—with a mild electrical current as well as back-and-forth (thrusting) movements of the probe. This technique has proven effective in numerous species of mammals, including virtually all of those in which male homosexual mounting and /or anal penetration occur. For further information on electroejaculation, see Watson, P. F., ed. (1978) Artificial Breeding of Non-Domestic Animals, Symposia of the Zoological Society of London no. 43, especially pp. 109, 129, 208—10, 221, 295 (London: Academic Press).

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Hanuman Langur (Srivastava et al. 1991:506–7); for a similar assessment with regard to homosexual activity between males in this species, see Weber and Vogel (1970:77–78). See also Rowell (1967a:23), who states that “sexual” and “dominance” mounts in Savanna (Yellow) Baboons are virtually indistinguishable, and Enomoto (1990:473), who remarks on the difficulty of discriminating between sexual and ritualized dominance mounting in Bonobos because of the gradation between the two. Weinrich (Sexual Landscapes, p. 294), in discussing mounting between male Mountain Sheep, also points out how sexuality and dominance can both be part of the same behavior and suggests an analogy with human sexuality. Indeed, elements of consensual “dominance” or power-play, although rarely acknowledged, are often a part of human lovemaking and sexual pleasure, ranging along a continuum from gentle “love bites” to full sadomasochism (and nonconsensual dominance also figures prominently in many human sexual interactions, especially heterosexual ones).

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Japanese Macaque (Wolfe 1986:268); Rhesus Macaque (Akers and Conaway 1979:78); Greylag Goose (Lorenz 1991:206); Black-winged Stilt (Kitagawa 1989:65, 69) (see also the distinction between same-sex courtship and aggressive/appeasing kantling in Ostriches [Sauer 1972:731; Bertram 1992:15, 50–51]). For species such as these that have a clear distinction between mounts in sexual and nonsexual contexts, only the former are considered (in this book and in most sources) to be homosexual behavior. As noted in chapter 1, some species classified by Dagg (1984) as exhibiting homosexuality (e.g., bush squirrels and degus) are excluded from our roster on the basis of this criterion, because all same-sex mounting in these species appears to fall into this genuinely nonsexual category; see Viljoen, S. (1977) “Behavior of the Bush Squirrel, Paraxerus cepapi cepapi,” Mammalia 41:119—66; Fulk, G. W. (1976) “Notes on the Activity, Reproduction, and Social Behavior of Octodon degus,” Journal of Mammology 57:495–505.

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Walrus (Miller 1975:607); Gray Seal (Anderson and Fedak 1985); Oystercatcher (Ens, B. J., and J. D. Goss- Custard [1986] “Piping as a Display of Dominance by Wintering Oystercatchers Haematopus ostralegus,” Ibis 128:382—91). Early observers of this species (e.g., Makkink 1942) misinterpreted the piping display as a courtship activity because it often occurs between males and females.

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For details of the way that dominance is expressed in these species, see Savanna (Yellow) Baboon (Maxim and Buettner-Janusch 1963:169); Hamadryas Baboon (Stammbach, E. [1978] “On Social Differentiation in Groups of Captive Female Hamadryas Baboons,” Behavior 67:322-38); Bottlenose Dolphin (Samuels and Gifford 1997); Killer Whale (Rose 1992:108-9); Caribou (Espmark, J. [1964] “Studies in Dominance- Subordination Relationship in a Group of Semi-Domestic Reindeer (Rangifer tarandus L.),” Animal Behavior 12:420-26); Blackbuck (Dubost and Feer 1981:97-100); Wolf (Zimen 1976, 1981); Bush Dog (Macdonald 1996); Spotted Hyena (Frank 1986:1511); Grizzly Bear (Craighead et al. 1995:109ff); Black Bear (Stonorov and Stokes 1972:235, 242); Red-necked Wallaby (Johnson 1989:267); Canada Goose (Collias and Jahn 1959:500-501); Scottish Crossbill (Nethersole-Thompson 1975:53); Black-billed Magpie (Birkhead 1991); Jackdaw (Roell 1978); Acorn Woodpecker (Stanback 1994); Galah (Rowley 1990:57). In Pronghorns, mounting between males was originally claimed to represent a dominance activity (Kitchen 1974), yet more recent studies of dominance in this species have not included same-sex mounting (Bromley 1991).

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In some cases, sexual behaviors other than mounting can be correlated with dominance. For example, grooming between males in Nilgiri Langurs and Crested Black Macaques is often performed by a subordinate animal on a more dominant one. Nevertheless, it is apparent that this activity has a clearly sexual component as well: one or both males may become intensely aroused, developing an erection and even ejaculating during the grooming (see Poirier 1970a:334 for Nilgiri Langurs and Poirier 1964:146—47 for Crested Black Macaques). Similarly, adult (dominant) Bonobos often masturbate or massage the genitals of adolescent (subordinate) males, but again, the activity involves clear sexual stimulation (cf. de Waal 1987, 1995, 1997). Also, Squirrel Monkey genital displays are sometimes correlated with dominance, but there are also cases where the association is less than definitive, or where they occur in clearly sexual contexts between animals of the same sex (cf. Talmage-Riggs and Anschel 1973:70; Travis and Holmes 1974:55; Baldwin and Baldwin 1981:295-97; Castell and Heinrich 1971:187-88).

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One cannot help but surmise that it is the heterosexism of many biologists that has led them to focus on mounting behavior to the exclusion of other activities in their appeal to dominance factors—for only in mounting can the positions of the participants be clearly analogized to those of a male and female in a heterosexual interaction. As Fedigan (1982:101 [Japanese Macaque]) points out, underlying the entire discussion of dominance in same-sex interactions is the assumption that homosexual mounting is essentially a transposition from heterosexual copulation—and that males “dominate” females in such interactions. For further evidence against this view, see the discussion of homosexuality as a form of “pseudoheterosexuality” in chapter 4.

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Possible exceptions are same-sex courtship interactions in Mountain Sheep (Geist 1968, 1971), Musk-oxen (Reinhardt 1985), and Cavies (Rood 1972), which have been interpreted as reflecting dominance. Additionally, mounting or other sexual behaviors within a same-sex pair-bond—common in many bird species—does not fit easily into a dominance interpretation, since this usually involves ongoing interaction with only one other animal (rather than the establishment of hierarchical positions within a network of individuals).

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Giraffe (Pratt and Anderson 1985:774—75, 780—81); Crested Black Macaque (Dixson 1977:77-78; Reed et al. 1997:255); Stumptail Macaque (Bernstein 1980:40); Pig-tailed Macaque (Giacoma and Messeri 1992:187); Savanna (Olive) Baboon (Owens 1976:250-51); Squirrel Monkey (Baldwin and Baldwin 1981:295-97; Baldwin 1968:296, 311); Red Squirrel (Ferron 1980:136); Spinifex Hopping Mouse (Happold 1976:147); American Bison (Reinhardt 1985:222-23); Pukeko (Lambert et al. 1994); Sociable Weaver (Collias and Collias 1980:246, 248; in the latter instance, the inconsistency in dominance status was not one of the cases of temporary reversals of dominance that were occasionally seen in this species). In female Squirrel Monkeys, dominance hierarchies are not considered to be a salient feature of social organization in the wild (Baldwin and Baldwin 1981:294-95). However, even when dominance systems appear to develop (e.g., in some captive situations), investigators have found that the rank of females based on their homosexual activities does not agree with other measures of rank (Anschel and Talmage- Riggs 1978:602 [table 1]).

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For some reevaluation and/or critiques of the concept of dominance, see Gartlan, J. S. (1968) “Structure and Function in Primate Society,” Folia Primatologica 8:89-120; Bernstein 1970 (Crab-eating Macaque); Richards, S. M. (1974) “The Concept of Dominance and Methods of Assessment,” Animal Behavior 22:914— 30; Ralls, K. (1976) “Mammals in Which Females Are Larger Than Males,” Quarterly Review of Biology 51:245-76; Lockwood, R. (1979) “Dominance in Wolves:

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