Bruce Bagemihl - Biological Exuberance

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As noted earlier, females also avoid STDs by refraining from genital contact during such mountings. For both Razorbills and spotted sandpipers, see Wagner, R. H. (1991) “The Use of Extrapair Copulations for Mate Appraisal by Razorbills, Alca torda” Behavioral Ecology 2:198—203. See also Koala for an example of a species with high rates of STDs in wild populations (Brown et al. 1987; Weigler et al. 1988). For other species in which significant portions of nonmonogamous matings are nonprocreative, see the profiles of Snow Goose, Lesser Scaup Duck, Common Murre, Oystercatcher, Silver Gull, and Swallows.

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In addition to pair-bonding species in which nonmonogamous or alternate parenting arrangements are adopted by some individuals, the opposite situation also occurs. In some species in which the mating arrangement is typically polygamous or in which males do not usually participate in parenting, some individuals deviate from this pattern. Monogamous pair-bonding occurs in some Gray Seals (Amos et al. 1995) and Ruffs (Cramp and Simmons 1983:391), for example, even though most individuals are polygamous in these species, while some male Mallards (Losito and Baldassarre 1996:692) and Lyrebirds (Smith 1988:37— 38) occasionally parent their offspring even though males of these species generally do not contribute to parental duties.

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Based on data from 140 populations of 76 different bird species, the average divorce rate is about 20 percent; only about 11 percent of these populations have no heterosexual divorce at all or rates of less than 1 percent. See appendix 19.1 in Ens, B. J., S. Choudhury, and J. M. Black (1996) “Mate Fidelity and Divorce in Monogamous Birds,” in J. M. Black, ed., Partnerships in Birds: The Study of Monogamy , pp. 344-401 (Oxford: Oxford University Press). For further discussion of divorce, see Choudhury, S. (1995) “Divorce in Birds: A Review of the Hypotheses,” Animal Behavior 50:413-29; Rowley, I. (1983) “Re-Mating in Birds,” in P. Bateson, ed., Mate Choice , pp. 331-60 (Cambridge: Cambridge University Press).

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Oystercatcher (Harris et al. 1987:47, 55); Ocellated Antbird (Willis 1973:35-36); Warthog (Cumming 1975:89—90); White-tailed Deer (Gerlach, D., S. Atwater, and J. Schnell, eds. [1994] Deer , pp. 145, 150 [Mechanicsburg, Pa.: Stackpole Books]); Snow Goose (Prevett and MacInnes 1980:25, 43).

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Siamang (Fox 1977:409, 413—14).

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Common Murre (based on figures in Hatchwell 1988:161, 164, 168); Kleiman, D. G., and D. S. Mack (1977) “A Peak in Sexual Activity During Mid-Pregnancy in the Golden Lion Tamarin, Leontopithecus rosalia (Primates: Callitrichidae),” Journal of Mammalogy 58:657-60; Proboscis Monkey (Gorzitze 1996:77).

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Rhesus Macaque (Rowell et al. 1964:219); Mountain Goat (Hutchins 1984:45); addax antelope (Manski, D.A. [1982] “Herding of and Sexual Advances Toward Females in Late Stages of Pregnancy in Addax Antelope, Addax nasomaculatus,” Zoologische Garten 52:106-12; wildebeest (Watson, R. M. (1969) “Reproduction of Wildebeest, Connochaetes taurinus albojubatus Thomas, in the Serengeti Region, and Its Significance to Conservation,” p. 292, Journal of Reproduction and Fertility, supp. 6:287-310. One scientist (Loy 1970:294) goes so far as to suggest that the term estrus (meaning, roughly, the period when the female is “in heat”) should be redefined for Rhesus Macaques so as to make no reference to ovulation, since nonreproductive heterosexual behaviors are so prevalent in this species (traditionally, estrus is defined strictly in relation to the “reproductive” event of ovulation).

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See the index for examples of profiled species that engage in these activities. For cross-species surveys and additional examples, see also Rose et al. 1991 (Northern Elephant Seal); Robinson, S. K. (1988) “Anti-Social and Social Behavior of Adolescent Yellow-rumped Caciques (Icterinae: Cacicus cela ),” Animal Behavior 36:1482-95; Thornhill, N. W. (1992) The Natural History of Inbreeding and Outbreeding: Theoretical and Empirical Perspectives (Chicago: University of Chicago Press); Krizek, G. O. (1992) “Unusual Interaction Between a Butterfly and a Beetle: ‘Sexual Paraphilia’ in Insects?” Tropical Lepidoptera 3(2):118; Ishikawa, H. (1985) “An Abnormal Connection Between Indolestes peregrinus and Cercion hieroglyphicum ,” Tombo (Tokyo) 28(1—4):39; Matsui, M., and T. Satow (1975) “Abnormal Amplexus Found in the Breeding Japanese Toad,” Niigata Herpetological Journal 2:4-5; Riedman, M. (1990) The Pinnipeds: Seals, Sea Lions, and Walruses , pp. 216-17 (Berkeley: University of California Press).

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Lion (Eaton 1978; Bertram 1975:479); Raptors (Korpimaki et al. 1996).

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Oystercatcher (Heg et al. 1993:256); Kob (Buechner and Schloeth 1965:218-19).

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Such mounts are often described as “incomplete” or are viewed as nothing more than a component or prelude to “full” copulations. This implies that the “goal” of all sexual mounting is penetration, ejaculation, and ultimately, fertilization—certainly true for a great deal of mounting behavior, but by no means a uniform characterization of all sexual activity. For further discussion of what one biologist has aptly termed “fertilization myopia”—i.e., the narrowness and bias of most scientific descriptions of animal copulation, which focus only on “successful” matings (those that lead to fertilization)—see Eberhard, Female Control, pp. 28—34. For an example of “display” copulations in a bird species not profiled in part 2, as well as examples from other species, see Eberhard, Female Control, pp. 94-102; Strahl, S. D., and A. Schmitz (1990) “Hoatzins: Cooperative Breeding in a Folivorous Neotropical Bird,” p. 145, in P. B. Stacey and W. D. Koenig, eds., Cooperative Breeding in Birds: Long-term Studies of Ecology and Behavior , pp. 131—56 (Cambridge: Cambridge University Press).

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For a survey of mammal species where reverse mounting occurs, see Dagg (1984). Reverse mounting usually involves the female climbing on top of the male (and rarely includes penetration [in mammals] or cloacal contact [in birds]). Because heterosexual mating in Dolphins typically occurs with the male in an upside-down position underneath the female, however, “reverse” mounting in these species involves the female assuming a position underneath the male.

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In addition to the references for species profiled in part 2, descriptions and discussion of masturbation in a wide variety of other animals can be found in the following articles: Shadle, A. R. (1946) “Copulation in the Porcupine,” Journal of Wildlife Management 10:159—62; Ficken, M. S., and W. C. Dilger (1960) “Comments on Redirection with Examples of Avian Copulations with Substitute Objects,” Animal Behavior 8:219-22; Snow, B. K. (1977) “Comparison of the Leks of Guy’s Hermit Hummingbird Phaethornis guy in Costa Rica and Trinidad,” Ibis 119:211-14; Buechner, H. K., and S. F. Mackler (1978) “Breeding Behavior in Captive Indian Rhinoceros,” Zoologische Garten 48:305-22; Harger, M., and D. Lyon (1980) “Further Observations of Lek Behavior of the Green Hermit Hummingbird Phaethornis guy at Monteverde, Costa Rica,” Ibis 122:525—30; Wallis, S. J. (1983) “Sexual Behavior and Reproduction of Cercocebus albigena johnstonii in Kibale Forest, Western Uganda,” International Journal of Primatology 4:153—66; Poglayen-Neuwall, I., and I. Poglayen-Neuwall (1985) “Observations of Masturbation in Two Carnivora,” Zoologische Garten 1985 55:347—348; Frith, C. B., and D. W. Frith (1993) “Courtship Display of the Tooth-billed Bowerbird Scenopoeetes dentirostris and Its Behavioral and Systematic Significance,” Emu 93:129-36; Post, W. (1994) “Redirected Copulation by Male Boat-tailed Grackles,” Wilson Bulletin 106:770-71; Frith, C. B., and D. W. Frith (1997) “Courtship and Mating of the King of Saxony Bird of Paradise Pteridophora alberti in New Guinea with Comment on their Taxonomic Significance,” Emu 97:185—93.