The Myth of Monogamy: Fidelity and Infidelity in Animals and People (41 page)

34 The possibility arises that adult males: I. U. Reichard. 1995. Extra-pair copulations in a monogamous gibbon
(Hylobates lar). Ethology
100: 99-112.

34 As a result, the younger males fathered: E. S. Morton, L. Forman, and M. Braun. 1990. Extrapair fertilizations and the evolution of colonial breeding in purple martins.
Auk
107: 275-283.

198
THE MYTH OF MONOGAMY

35 Males that are cuckolded: P. C. Frederick. 1987. Extrapair couplations in the mating system of white ibis
(Eudocimus albus). Behaviour
100: 170-201; D. F. Westneat. 1988. Parental care and extrapair copulations in the indigo bunting.
Auk
105: 149-160.

35 The general pattern is concisely described: A. Johnsen and J. T. Lifjeld. 1995. Unattractive males guard their mates more closely: an experiment with bluethroats (Aves, Turdidae:
Luscinia s. svecica). Ethology
101: 200-212.

35 Several studies have confirmed: 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; B. Kempenaers, G. R. Verheyen, and A. A. Dhondt. 1995. Mate guarding and copulation behaviour in monogamous and polygynous blue tits: do males follow a best-of-a-bad-job strategy?
Behavioral Ecology and Sociobiology
36: 33-42.

35 And not only birds: S. W. Gangestad and R. Thornhill. 1997. An evolutionary psychological analysis of human sexual selection: developmental features, male sexual behavior, and mediating features. In
Evolutionary Social Psychology,
ed. J. A. Simpson and D. T. Kenrick. Hillsdale, NJ: Erlbaum.

35 Several times this was seen: A. P. M0ller. 1990. Deceptive use of alarm calls by male swallows
Hirundo rustica:
a new paternity guard.
Behavioral Ecology
1: 1-6.

36 By ruining her web: P. J. Watson. 1986. Transmission of a female sex pheromone thwarted by males in the spider
Linyphia litigiosa
Keyserling (Linyphiidae).
Science
233:219-221.

36 Eberhard reviewed the sweaty details: W. G. Eberhard. 1994. Evidence for widespread courtship during copulation in 131 species of insects and spiders, and implications for cryptic female choice.
Evolution
48: 711-733.

37 Because of their basic biology: D. Dewsbury. 1982. Ejaculate cost and male choice.
The American Naturalist
119: 601-610.

37 Sure enough, when male rats: R. R. Bellis, M. A. Baker, and M. J. G. Gage. 1990. Variation in rat ejaculates consistent with the kamikaze sperm hypothesis.
Journal of Mammalology
71: 479-480.

37 Plains zebra stallions copulate less: J. R. Ginsberg and D. I. Rubenstein. 1990. Sperm competition and variation in zebra mating behavior.
Behavioral Ecology and Sociobiology
26: 427-434.

38 Males are absent: T. Birkhead and C. M. Lessels. 1988. Copulation behaviour of the osprey
Pandion haliaetus. Animal Behaviour
36: 1672-1682.

38 This has been especially well established: A. Poole. 1989.
Ospreys: A Natural and Unnatural History.
Cambridge: Cambridge University Press.

38 Among orioles, males will copulate: B. B. Edinger. 1988. Extra-pair courtship and copulation attempts in northern orioles.
Condor
90: 546-554.

NOTES
199

39 Very quickly afterward: D. P. Barash. 1977. Sociobiology of rape in mallards
(Anas platyrhinchos):
responses of the mated male.
Science
197: 788-789.

39 As soon as one male copulates: J. Faaborg and J. C. Bednarz. 1990. Galapagos and Harris' hawks: divergent causes of sociality in two raptors. In
Cooperative Breeding in Birds,
ed. P. B. Stacey and W. D. Koenig. Cambridge: Cambridge University Press.

39 Among rats, males mate: M. K. Matthews and N. T. Adler. 1977. Systematic interrelationship of mating, vaginal plug position, and sperm transport in the rat.
Physiology and Behavior
20: 303-309.

39 Among nonhuman primates: C. D. Busse and D. Q. Estep. 1984. Sexual arousal in male pigtailed monkeys
(Macaca nemestrina):
effects of serial matings by two males.
Journal of Comparative Psychology
98: 227-231; D. Q. Estep, T. P. Gordan, M. E. Wilson, and M. L. Walker. 1986. Social stimulation and the resumption of copulation in rhesus
(Macaca mulatto)
and stumptail
(M. arctoides)
macaques.
International Journal of Primatology
7: 507-517.

40 In at least one rat species: D. Q. Estep. 1988. Copulations by other males shorten the post-ejaculatory intervals of pairs of roof rats,
Rattus rattus. Animal Behaviour
36: 299-300.

40 The males of such species: A. P. Mailer. 1988. Testis size, ejaculate quality, and sperm competition in birds.
Biological Journal of the Linnean Society
33: 273-283.

40 It has been found for mammals generally: A. P. Moller. Ejaculate quality, testis size and sperm production in mammals.
Functional Ecology
3: 91-96; J. R. Ginsburg and D. I. Rubenstein. 1990. Sperm competition and variation in zebra mating behavior.
Behavioral Ecology and Sociobiology 26:
427-434; R. L. Brownell and K. Ralls. 1986. Potential for sperm competition in baleen whales.
Reports of the International Whale Commission
(special Issue) 8: 97-112.

40 Ditto for primates: A. P. Moller. 1988. Ejaculate quality, testis size and sperm competition in primates.
Journal of Human Evolution
17: 479-483.

41 The answer is pretty clear: T. Birkhead and A. P. Moller. 1992.
Sperm Competition in Birds.
San Diego: Academic Press.

42 One researcher tried injecting: P. Sugawara. 1979. Stretch reception in the bursa copulatrix of the butterfly
Pieris rapae crucivora,
and its role in behaviour.
Journal of Comparative Physiology
130: 191-199.

42 It is interesting to note: R. E. Silberglied, J. G. Sheperd, and J. L. Dickinson. 1984. Eunuchs: the role of apyrene sperm in lepidoptera?
The American Naturalist
123: 255-265; P. A. Cook and M. J. G. Gage. 1995. Effects of risks of sperm competition on the numbers of eupyrene and apyrene sperm ejaculated by the moth
Plodia interpunctella. Behavioral Ecology and Sociobiology
36: 261-268.

43 It is interesting that ring doves: C. J. Erickson and P. G. Zenone. 1976. Courtship differences in male ring doves: avoidance of cuckoldry?
Science
192: 1353-1354.

200
THE MYTH OF MONOGAMY

44 Copulating males use their penis: Jonathan K. Waage. 1979. Dual function of the damselfly penis: sperm removal and transfer.
Science
203: 916-918; see also R. L. Smith, ed., 1984.
Sperm Competition and the Evolution of Animal Mating Systems.
New York: Academic Press.

44 Male pygmy octopuses: J. A. Cigliano. 1995. Assessment of the mating history of female pygmy octopuses and a possible sperm competition mechanism.
Animal Behaviour
49: 849-851.

44 Prior to mating: P. L. Miller. 1990. Mechanisms of sperm removal and sperm transfer in
Orthetrum coerulescens
(Fabricus) (Odonata: Libellulidae).
Physiological Entomology
15: 199-209.

44 The more time a female dunnock: N. B. Davies. 1983. Polyandry, cloaca-pecking and sperm competition in dunnocks.
Nature
302: 334-336.

45 If and when the victim copulates: J. Carayon. 1974. Insemination traumatique heterosexualle et homosexualle chex
Xylocoris maculipennis. Comptes Rendues Academie de Sciences de Paris, Series D
278: 2803-2806.

45 To put it bluntly: Except if that someone is a close relative, in which case such behavior can be favored via "kin selection." For our purposes, however, it is accurate to say that any tendency to rear someone else's offspring will be strongly selected against.

45 It may surprise some readers: F. Cezilly and R. G. Nager. 1995 Comparative evidence for a positive association between divorce and extra-pair paternity in birds.
Proceedings of the Royal Society of London, Series B
262: 7-12.

46 Among indigo buntings: D. F. Westneat and P. W. Sherman. 1990. When monogamy isn't.
The Living Bird Quarterly
9: 24-28.

47 So-called cooperative breeding: P. P. Rabenold, K. N. Rabenold, W. H. Piper, et al. 1990. Shared paternity revealed by genetic analysis in cooperative breeding tropical wrens.
Nature
348: 538-542.

47 It turned out that it didn't matter: O. Svensson, C. Magnhagen, E. Forsgren, and C. Kvarnemo. 1998. Parental behaviour in relation to the occurrence of sneaking in the comon goby.
Animal Behaviour
56: 175-179.

47 It is not invariant: L. A. Whittingham and J. T. Lifjeld. 1995. High paternal investment in unrelated young: extra-pair paternity and male parental care in house martins.
Behavioral Ecology and Sociobiology
37: 103-108.

48 Or maybe in certain cases: I. P. F. Owens. 1993. When kids just aren't worth it: cuckoldry and parental care.
Trends in Ecology and Evolution
8: 269-271.

48 The male behaved aggressively: D. P. Barash. 1976. The male response to apparent female adultery in the mountain bluebird,
Sialia currucoides:
An evolutionary interpretation.
The American Naturalist
110: 1097-1101.

48 In any event, there have since been: E.g., N. B. Davies, B. J. Hatchwell, T. Burke, and T. Robson. 1992. Paternity and parental effort in dunnocks
Prunella modularis:

NOTES 201

how good are male chick-feeding rules?
Animal Behaviour
43: 729-745; L. A. Whittingham, P. D. Taylor, and R. J. Robertson. 1992. Confidence of paternity and male parental care.
The American Naturalist
139: 1115-1125; B. C. Sheldon, K. Rasanen, and P. C. Dias. 1997. Certainty of paternity and parental care in collared flycatchers: an experiment.
Behavioral Ecology
8: 421-428; see review by Jonathan Wright. 1998. Paternity and paternal care. In
Sperm Competition and Sexual Selection,
ed. T. R. Birkhead and A. P. Loller. San Diego: Academic Press.