54

Bonobo (Savage-Rumbaugh and Wilkerson 1978:334; Roth 1995:75, 88).

55

Bonobo (Savage-Rumbaugh et al. 1977:108).

56

Linguists studying the structure of American Sign Language, for example, have identified a continuum of iconicity in signs, ranging from transparent signs (quasi-mimetic gestures whose meaning is readily identifiable from their form, even to nonsigners) to translucent signs (gestures in which a connection between meaning and form can be discerned but not automatically identified without knowing the meaning of the sign) to opaque signs (gestures in which all form-meaning correspondences have been lost). According to these criteria, the Bonobo gestures would fall primarily in the transparent-translucent range. For further discussion see Klima, E.S., and U. Bellugi (1979) The Signs of Language, especially chapter 1, “Iconicity in Signs and Signing” (Cambridge: Harvard University Press).

57

This gestural system has only been observed in captivity, albeit in “untrained” Bonobos. Studies of wild Bonobos have so far revealed a less elaborate communicative repertoire associated with sexual interactions, although researchers have identified similar types of communicative exchanges prior to some episodes of sexual activity (e.g., Kitamura 1989:54-55; Enomoto 1990:473-75). It must also be remembered that many behaviors are easily missed in the field (especially given the particular difficulties of observing wild Bonobos; cf. de Waal 1997:12, 63—64, 70, 76—77); hence it is possible that more elaborate gestural repertoires do occur in wild Bonobos but have yet to be observed. For more on the issue of behaviors that are only observed in captivity as opposed to the wild, see chapter 4.

58

Hewes, G. W. (1973) “Primate Communication and the Gestural Origin of Language,” Current Anthropology 14:5-24; Hewes, G. W. (1976) “The Current Status of the Gestural Theory of Language Origin,” in S. Harnad, H. Steklis, and J. Lancaster, eds., Origins and Evolution of Language and Speech. Annals of the New York Academy of Science, vol. 280, pp. 482-504 (New York: New York Academy of Sciences).

59

Bonobo (Roth 1995:4-45).

60

Beck, B. B. (1980) Animal Tool Behavior: The Use and Manufacture of Tools by Animals (New York: Garland); Goodall 1986:545—48, 559 (Common Chimpanzee); van Lawick- Goodall, J., H. van Lawick, and C. Packer (1973) “Tool-Use in Free-living Baboons in the Gombe National Park, Tanzania,” Nature 241:212-13; McGrew, W.C. (1992) Chimpanzee Material Culture: Implications for Human Evolution (Cambridge: Cambridge University Press); Berthelet, A., and J. Chavaillon, eds., (1993) The Use of Tools by Human and Nonhuman Primates (Oxford: Clarendon Press); Weinberg, S.M., and D.K. Candland (1981) “‘Stone-Grooming’ in Macaca fuscata,” American Journal of Primatology 1:465—68.

61

Orang-utan (Rijksen 1978:262-63; Nadler 1982:241; Harrison 1961:61).

62

Common Chimpanzee (Bingham 1928:148-50; Kollar et al. 1968:456-57; Goodall 1986:559-60; McGrew, Chimpanzee Material Culture, p. 183); Bonobo (Takeshita and Walraven 1996:428; Walraven et al. 1993:28, 30; Becker, C. [1984] Orang-Utans und Bonobos im Spiel: Untersuchungen zum Spielverhalten von Menschenaffen [Orang-utans and Bonobos at Play: Investigations on the Play Behavior of Apes], pp. 149, 152, 193-94 [Munich: Profil-Verlag]). Female Japanese Macaques are also reported to use inanimate objects for masturbation (Rendall and Taylor 1991:321), although it is not clear whether this involves use of “tools” or simply rubbing of genitals against a surface (as is found in many other species). Masturbatory tool use is also occasionally reported for animals other than primates; see, for example, Shadle’s description of male and female Porcupines holding sticks in their forepaws while straddling the object in order to stimulate their genitals (Shadle, A. R. [1946] “Copulation in the Porcupine,” Journal of Wildlife Management 10:159—62; Shadle, A. R., M. Smelzer, and M. Metz [1946] “The Sex Reactions of Porcupines (Erethizon d. dorsaturn) Before and After Copulation,” Journal of Mammalogy 27:116—21). Objects or “tools” are also sometimes employed by Common Chimpanzees and Bonobos during heterosexual courtship and solicitations (cf. McGrew, Chimpanzee Material Culture, pp. 82, 188; Nishida 1997:385, 394 [Common Chimpanzee]; de Waal 1997:120 [Bonobo]).

63

Bonnet Macaque (Sinha 1997). Sinha (1997:23) believes that this female was using the tools to “scratch” her vagina, possibly because of “some irritation,” whose presence, however, was never confirmed. Sexual stimulation is also compatible with the observed behaviors (instead of, or along with, “scratching”), especially considering that masturbation without the use of tools occurs regularly in Bonnet Macaques of both sexes (cf. Makwana 1980:11; Kaufman and Rosenblum 1966:221; Rahaman and Parthasarathy 1969:155).

64

See, for example, (Rawson, P. (1973) Primitive Erotic Art, especially pp. 20, 71 (New York: G. P. Putnam’s Sons); Kinsey, A. C., W. B. Pomeroy, C. E. Martin, and P. H. Gebhard (1953) Sexual Behavior in the Human Female, p. 136 (Philadelphia: W. B. Saunders). Examples of tools utilized for sexual stimulation of partners (rather than self-stimulation) have yet to be reported for any nonhuman species. For a recent discussion of the role of sexual pleasure in the evolution of tool use among both nonhuman primates and early humans, see Vasey, P. L. (1998) “Intimate Sexual Relations in Prehistory: Lessons from Japanese Macaques,” World Archaeology 29:407—25.

65

For further discussion of these (and other) examples as well as cultural variation in the occurrence of incest and its taboos, see Leavitt, G. C. (1990) “Sociobiological Explanations of Incest Avoidance: A Critical Review of Evidential Claims,” American Anthropologist 92:971-93; Arens, W. (1986) The Original Sin: Incest and Its Meaning (New York and Oxford: Oxford University Press); Livingstone, F. B. (1980) “Cultural Causes of Genetic Change,” in G. W. Barlow and J. Silverberg, eds., Sociobiology: Beyond Nature/Nurture? pp. 307—29, AAAS Selected Symposium, no. 35 (Boulder: Westview Press); Schneider, D. M. (1976) “The Meaning of Incest,” Journal of the Polynesian Society 85:149—69.

66

For an overview of a variety of kinship restrictions on Melanesian homosexual relations, see Schwimmer, E. (1984) “Male Couples in New Guinea,” pp. 276-77, in G. H. Herdt, ed., Ritualized Homosexuality in Melanesia, pp. 248—91 (Berkeley: University of California Press); Murray, S. O. (1992) “Age-Stratified Homosexuality: Introduction,” pp. 10-12, in Murray, Oceanic Homosexualities, pp. 293- 327. For more on New Guinean homosexualities, see chapter 6.

67

Leavitt, “Sociobiological Explanations,” pp. 974-75; Livingstone, “Cultural Causes,” p. 318. For arguments that in animals some forms of inbreeding (such as between cousins) may actually have beneficial genetic and social effects and are preferred in some species (e.g., Vervet monkeys, Japanese quail), see Moore and Ali 1984 (Bonnet Macaque); Bateson, P. (1982) “Preferences for Cousins in Japanese Quail,” Nature 295:236—37; Shields, W. M. (1982) Philopatry, Inbreeding, and the Evolution of Sex (Albany: State University of New York Press); Cheney, D. M., and R. M. Seyfarth (1982) “Recognition of Individuals Within and Between Groups of Free-Ranging Vervet Monkeys,” American Zoologist 22:519-30.

68

Japanese Macaque (Wolfe 1979; Chapais and Mignault 1991; Vasey 1996:543; Chapais et al. 1997);

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