The Myth of Monogamy: Fidelity and Infidelity in Animals and People (42 page)

48 In another cooperatively breeding bird species: W. D. Koenig. 1990. Opportunity of parentage and nest destruction in polygynandrous acorn woodpeckers,
Melanerpes formicivorus. Behavioral Ecology
1: 55-61.

49 For example, male indigo buntings: D. F. Westneat. 1988. Parental care and extrapair copulations in the indigo bunting.
Auk
105: 149-160.

50 Their offspring get short shrift: D. P. Barash. 1975. Ecology of parental behavior in the hoary marmot
(Marmota caligata). Journal of Mammalogy
56: 613-618.

50 Attractive males usually provide: A. P. Mailer and R. Thornhill. 1998. Male parental care, differential parental investment by females and sexual selection.
Animal Behaviour 55:
1507-1515.

50 This tendency is captured: B. C. Sheldon, J. Merila, A. Qvarnstrom,

L. Gustafsson, and H. Ellegren. 1997. Paternal contribution to offspring condition is predicted by size of male secondary sexual characteristic.
Proceedings of the Royal Society of London, Series B
264: 297-302.

51 Incidentally, long-tailed male barn swallows: A. P. Mailer, A. Barbosa, J. J. Cuervo, F. de Lope, S. Merino, and N.. Saino. 1998. Sexual selection and tail streamers in the barn swallow.
Proceedings of the Royal Society of London, Series
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51 They simply have nothing: R. H. Wagner, M. D. Schug, and E. S. Morton. 1996. Confidence of paternity, actual paternity and parental effort by purple martins.
Animal Behaviour $2:
123-132.

52 I and others have documented: F. McKinney, S. R. Derrickson, and P. Mineau. 1983. Forced copulation in waterfowl.
Behaviour
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52 Since in nearly all cases: A. P. Mailer. 1987. House sparrow
Passer domesticus
communal displays.
Animal Behaviour
35: 203-210.

53 Males are sometimes aggressive: D. F. Westneat. 1987. Extrapair copulations in a predominatly monogamous bird: observations of behaviour.
Animal Behaviour
35: 865-876.

54 There is in fact: R. Thornhill and C. Palmer. 2000.
The Natural History of Rape.
Cambridge, MA: MIT Press.

54 This has subsequently been found: J. T. Burns, K. Cheng, and F. McKinney. 1980. Forced copulation in captive mallards: I. Fertilization of eggs.
Auk
97:

202
THE MYTH OF MONOGAMY

875-879; F. McKinney and P. Stolen. 1982. Extra-pair bond courtship and forced copulation among captive green-winged teal
(Anas carolinensis). Animal Behaviour
30: 461-474; F. McKinney, K. M. Cheng, and D. Bruggers. 1984. Sperm competition in apparently monogamous birds. In
Sperm Competition and the Evolution of Animal Mating Systems,
ed. R. L. Smith. New York: Academic Press; K. M. Cheng, J. T. Burns, and F. McKinney. 1983. Forced copulation in captive mallards: III. Sperm competition.
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100: 302-310.

54 Probably because, given "last male advantage": L. G. Sorenson. 1994. Forced extra-pair copulation and mate guarding in the white-cheeked pintail: timing and tradeoffs in an asynchronously breeding duck.
Animal Behaviour
48: 519-533.

54 Research conducted at the largest known goose colony: P. O. Dunn, A. D. Afton, M. L. Gloutney, and R. T. Alisauskas. 1999. Forced copulation results in few extrapair fertilizations in Ross's and lesser snow geese.
Animal Behaviour 57:
1071-1081.

54 It appears that species: G. Gauthier. 1988. Territorial behavior, forced copulations and mixed reproductive strategy in ducks.
Wildfowl
39: 102-114.

55 About one-third of spousal killings: M. Daly, M. Wilson, and S. Weghorst. 1982. Male sexual jealousy.
Ethology and Sociobiology
3: 11-27.

55 The frequency of infidelity-generated violence: J. M. Tanner. 1970.
Homicide in Uganda, 1964.
Uppsala, Sweden: Scandinavian Institute of African Studies; C. F. Lobban. 1972.
Law and Anthropology in the Sudan.
African Studies Seminar Series No. 13. Khartoum, Sudan: Sudan Research Unit, Khartoum University.

55 Furthermore, as we shall see: D. M. Buss and D. P. Schmitt. 1994. Sexual strategies theory: a contextual evolutionary analysis of human mating.
Psychological Review
100: 204-232.

chapter 3
Undermining the Myth: Females (Choosing Male Genes)

57 Clearly, there was some hanky-panky: O. Bray, J. Kennelly, and J. Guarlno. 1975. Fertility of eggs produced on territories of vasectomized red-winged blackbirds.
Wilson Bulletin
87: 187-195.

58 In such cases--and especially when genetic testing: S. M. Smith. 1988. Extra-pair copulations in black-capped chickadees: the role of the female.
Behaviour
107: 15-23; B. Kempenaers, G. R. Verheyen, M. Van den Broeck, T. Burke, C. Van Broeckhoven, and A. A. Dhondt. 1992. Extra-pair paternity results from female preference for high-quality males in the blue tit.
Nature
357: 494-496.

58 One possibility is that extra-pair copulations: T. Halliday and S. Arnold. 1987. Multiple mating by females: a perspective from quantitative genetics.
Animal Behaviour
35: 939-941.

59 An interesting idea, this: K. M. Cheng and P. B. Siegel. 1990. Quantitative genetics of multiple mating.
Animal Behaviour
40: 406-407.

NOTES
203

60 The upshot is that females who copulate: D. W. Pyle and M. H. Gromko. 1978. Repeated mating by female
Drosophila melanogaster:
the adaptive importance.
Experimentia
34: 449-450; T. R. Birkhead and A. P. Moller. 1992.
Sperm Competition in Birds: Evolutionary Causes and Consequences.
London: Academic Press.

60 This is suggested by the fact: J. Graves, J. Ortega-Ruano, and P. J. B. Slater. 1993. Extra-pair copulations and paternity in shags: do females choose better males?
Proceedings of the Royal Society of London, Series B
253: 3-7.

60 Females consistently have more EPCs: J. Wetton and D. Parkin. 1991. An association between fertility and cuckoldry in the house sparrow,
Passer domesticus. Proceedings of the Royal Society of London, Series B
245: 227-233.

60 Among those birds whose females: E. M. Gray. 1997. Do red-winged blackbirds benefit genetically from seeking copulations with extra-pair males?
Animal Behaviour
53: 605-623.

61 They do not initiate: D. F. Westneat. 1992. Do female red-winged blackbirds engage in a mixed mating strategy?
Ethology
92: 7-28.

61 Extra-pair copulations are frequent: J. H. Wetton and D. T. Parkin. 1991. An association between fertility and cuckoldry in the house sparrow
Passer domesticus. Proceedings of the Royal Society of London, Series B
245: 227-233.

61 In addition, litter size is larger: J. L. Hoogland. 1998. Why do Gunnison's prairie dogs copulate with more than one male?
Animal Behaviour 55:
351-359.

61 Not all mammals show this pattern: J. O. Murie. 1996. Mating behavior of Columbian ground squirrels: I. Multiple mating by females and multiple paternity.
Canadian Journal of Zoology
73: 1819-1826.

61 In fact, even in another prairie dog species: J. L. Hoogland. 1995.
The Black-Tailed Prairie Dog: Social Life of a Burrowing Mammal.
Chicago: University of Chicago Press.

61 In some mammals, there is actually a
reduction:
K. E. Wynne-Edwards and R. D. Lisk. 1984. Djungarian hamsters fail to conceive in the presence of multiple males.
Animal Behaviour
32: 626-628.

62 The more males a female adder mates with: T. Madsen, R. Shine, J. Loman, and T. Hakansson. 1992. Why do female adders copulate so frequently?
Nature
365: 440-441.

63 DNA fingerprinting has shown: M. Ollson, R. Shine, A. Gullberg, A. Madsen, and J. Tegelstrom. 1996. Female lizards control the paternity of their offspring by selective use of sperm.
Nature
383: 585.

63 By multiple mating, a questing female: J. A. Zeh and D. W. Zeh. 1996. The evolution of polyandry: I. Intragenomic conflict and genetic incompatability.
Proceedings of the Royal Society of London, Series B
263: 1711-1717; J. A. Zeh and D. W. Zeh. 1997. The evolution of polyandry: II. Post-copulatory defences

204
THE MYTH OF MONOGAMY

against genetic incompatability.
Proceedings of the Royal Society of London, Series B
264: 69-75.

63 In a bird species wonderfully called: M. G. Brooker, I. Rowley, M. Adams, and R Baverstock. 1990. Promiscuity: an inbreeding avoidance mechanism in a socially monogamous species?
Behavioral Ecology and Sociobiology 26:
191-199.

63 Interestingly, when female mice: W. K. Potts, C. J. Manning, and E. K. Wakeland. 1991. Mating patterns in semi-natural populations of mice influenced by MHC genotype.
Nature
352: 619-621.

63 Female primates, for their part: B. A. Smuts. 1987. Gender, aggression and influence. In
Primate Societies,
ed. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham, and T. T. Struhsaker. Chicago: University of Chicago Press.

64 For example, in one troop: Y. Takahata. 1982. The socio-sexual behavior of Japanese monkeys.
Zeitschrift fur Tierpsychologie
59: 89-108.

64 In one remarkable case: R. Sekulic. 1982. Behavior and ranging patterns of a solitary female red howler
(Alouatta seniculus). Folia Primatologica
38: 217-232.

64 A key summary point is that: S. B. Hrdy and P. L. Whitten. 1987. Patterning of sexual activity. In
Primate Societies,
ed. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham, and T. T. Struhsaker. Chicago: University of Chicago Press.

64 In this species, therefore: O. Ratti, M. Hovi, A. Lundberg, H. Tegelstrom, and R. Alatalo. 1995. Extra-pair paternity and male characteristics in the pied flycatcher.
Behavioral Ecology and Sociobiology
37: 419-425.

66 They also vocalize loudly: C. R. Cox and B. J. LeBoeuf. 1977. Female incitation of male competition: a mechanism in sexual selection.
The American Naturalist
111: 317-335; J. H. Poole. 1989. Mate guarding, reproductibve success and female choice in African elephants.
Animal Behaviour
37: 842-849.

66 Or females can advertise: R. H. Wiley and J. Poston. 1996. Indirect mate choice, competition for mates, and co-evolution of the sexes.
Evolution
50: 1371-1381.

66 In eight of twelve observed copulations: J. J. Perry-Richardson, C. S. Wilson, and N. B. Ford. 1990. Courtship of the garter snake,
Thamnophic marianus,
with a description of a female behavior for coitus interruption.
Journal of Herpetology
24: 76-78.

66 Interestingly, of ten such forced copulations: R. Thornhill. 1988. The jungle fowl hen's cackle incites male competition.
Verhalten Deutsche Zoologische Geselschaft
81: 145-154.

66 Once, after he forced such a copulation: T. R. Birkhead and A. P. M0ller. 1992.
Sperm Competition in Birds: Evolutionary Causes and Consequences.
London: Academic Press.

67 For example, a female pied flycatcher: M. Hovi and O. Ratti. 1994. Mate sampling and assessment procedures in female pied flycatchers
(Ficedula hypoleuca). Ethology 96:
127-137.

NOTES
205

67 In this situation, most females: C. T. Gabor and T. R. Haliday. 1997. Sequential mate choice by smooth newts: females become more choosy.
Behavioral Ecology
8: 162-166.

68 Not only that, but the offspring: P. J. Watson. 1998. Multi-male mating and female choice increase offspring growth in the spider
Neriene litigiosa
(Linyphiidae).
Animal Behaviour
55: 387-403.

68 For example, among waterfowl: F. McKinney, S. R. Derrickson, and P. Mineau. 1983. Forced copulation in waterfowl.
Behaviour
86: 250-294.

69 This seems to be a way for females: J. A. Zeh, S. D. Newcomer, and D. W. Zeh. 1998. Polyandrous females discriminate against previous mates.
Proceedings of the National Academy of Sciences
95: 13732-13736.

69 It had already been demonstrated: J. A. Zeh. 1997. Polyandry and enhanced ' reproductive success in the harlequin beetle-riding pseudoscorpion.
Behavioral Ecology and Sociobiology
40: 111-118.

69 In one type of beetle: M. S. Archer and M. E. Elgar. 1999. Female preference for multiple partners: sperm competition in the hide beetle,
Dermestes maculatus. Animal Behaviour
58:
669-675.

69 In some insects, the female deposits: J. A. Zeh, S. D. Newcomer, and D. W. Zeh. 1998. Polyandrous females discriminate against previous mates.
Proceedings of the National Academy of Sciences
95: 13732-13736.

69 Most commonly, however, it appears that: M. Petrie. 1994. Improved growth and survival of offspring of peacocks with more elaborate trains.
Nature
341: 598-599.

70 Male house sparrows, for example: A. P. Moller. 1990. Sexual behaviour is related to badge size in the house sparrow
Passer domesticus. Behavioral Ecology and Sociobiology 27:
23-29.

70 Among zebra finches: N. Burley and D. Price. 1991. Extra-pair copulation and attractiveness in zebra finches.
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20: 1367-1372.

71 This supports the hypothesis: R. Wagner. 1991. The role of extra-pair copulations in razorbill mating strategies. D. Phil, thesis, University of Oxford, Oxford, UK.

71 As a result of such choice: P. Dunn and A. Cockburn. 1996. Evolution of male paternal care in a bird with almost complete cuckoldry.
Evolution
50: 2542-2548.

72 Incidentally, these helpers: Ibid.

72 Accordingly, females are well advised: Allison Welch, R. Semlitsch, and H. C. Gerhardt. 1998. Call duration as an indicator of genetic quality in male gray tree frogs.
Science
280: 1928-1930.

72 It has been suggested that bright coloration: W. D. Hamilton. 1990. Mate choice near or far.
American Zoologist
30: 341-352.

206
THE MYTH OF MONOGAMY

73 Such a correlaion does not: A. P. Mailer. 1997. Immune defence, extra-pair paternity, and sexual selection in birds.
Proceedings of the Royal Society of London, Series B
264:
561-566.

73 It is then also revealing: B. Kempenaers, G. R. Verheyen, M. Van den Broeck, T. Burke, C. Van Broeckhoven, and A. A. Dhondt. 1992. Extra-pair paternity results from female preference for high-quality males in the blue tit.
Nature
357: 494-496.