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).
143
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).
144
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).
145
Lion (Eaton 1978; Bertram 1975:479); Raptors (Korpimaki et al. 1996).
146
Oystercatcher (Heg et al. 1993:256); Kob (Buechner and Schloeth 1965:218-19).
147
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).
148
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.
149
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.
150
As a rough measure of the overwhelming attention devoted to male as opposed to female genitalia, for example, the Zoological Record for 1978—97 lists 539 articles that deal with the penis, compared to only 7 for the clitoris (the Zoological Record is a comprehensive electronic database that indexes more than a million zoological source documents, including articles from over 6,000 journals worldwide; the following keywords/search terms were used in compiling this estimate: penis/penile/penial/penes, phallus/phallic, baculum, hemipenes, clitorislclitorallclitorides, (os) clitoridis).
151
Stumptail Macaque (Goldfoot et al. 1980); Rhesus Macaque (Zumpe, D., and R. P. Michael [1968] “The Clutching Reaction and Orgasm in the Female Rhesus Monkey [Macaca mulatta],” Journal of Endocrinology 40:117—23). In what is perhaps the most extreme “experiment” of this type, female Rhesus Macaques were strapped to an apparatus made of iron and wood and forced to undergo stimulation with a dildo or “penis substitute” while their responses were monitored with electrodes (Burton, F. D. [1971] “Sexual Climax in Female Macaca mulatta,” in J. Biegert and W. Leutenegger, eds., Proceedings of the 3rd International Congress of Primatology, vol. 3, pp. 180-91 [Basel: S. Karger]).
152
For a sample of some of this debate, see Allen, M. L., and W. B. Lemmon (1981) “Orgasm in Female Primates,” American Journal of Primatology 1:15—34; Rancour- Laferriere, D. (1983) “Four Adaptive Aspects of the Female Orgasm,” Journal of Social and Biological Structures 6:319-33; Baker, R., and M. A. Bellis (1995) Human Sperm Competition: Copulation, Masturbation, and Infidelity, pp. 234-49 (London: Chapman and Hall); Hrdy, S.B. (1996) “The Evolution of Female Orgasms: Logic Please but No Atavism,” Animal Behavior 52:851-52; Thornhill, R., and S. W. Gangstead (1996) “Human Female Copulatory Orgasm: A Human Adaptation or Phylogenetic Holdover,” Animal Behavior 52:853—55. For recent discussions that sidestep the question of sexual pleasure with regard to the “function” of the clitoris, as well as in relation to a variety of specific sexual behaviors (e.g., stimulatory movements such as thrusting during intercourse, multiple ejaculations, lengthy copulations, etc.), see Baker and Bellis, Human Sperm Competition, pp. 126—31;
