Women After All: Sex, Evolution, and the End of Male Supremacy (43 page)

64
progesterone induces “the full suite of sexual behavior”:
Ibid., 4629.

66
Sex change fish:
J. Godwin, “Neuroendocrinology of Sexual Plasticity in Teleost Fishes,”
Frontiers in Neuroendocrinology
31, no. 2 (2010): 203–16.

66
Anglerfish with parasitic males:
Theodore W. Pietsch, “Dimorphism, Parasitism, and Sex Revisited: Modes of Reproduction Among Deep-Sea Ceratioid Anglerfishes (Teleostei: Lophiiformes),”
Ichthyological Research
52, no. 3 (2005): 207–36.

66
Reproduction in sea horses:
Kai N. Stölting and Anthony B. Wilson, “Male Pregnancy in Seahorses and Pipefish: Beyond the Mammalian Model,”
Bioessays
29, no. 9 (2007): 884–96; Camilla M. Whittington, K. Musolf, S. Sommer, and A. B. Wilson, “Behavioural Cues of Reproductive Status in Seahorses
Hippocampus abdominalis,
”
Journal of Fish Biology
83, no. 1 (2013): 220–26. A developmental gene that enables males in a closely related pipefish to have pregnancy pouches was found by April Harlin-Cognato and her colleagues: A. Harlin-Cognato, E. A. Hoffman, and A. G. Jones, “Gene Cooption Without Duplication During the Evolution of a Male-Pregnancy Gene in Pipefish,”
Proceedings of the National Academy of Sciences USA
103, no. 51 (2006): 19407–12.

67
a cloud of minute seahorses:
For a video of seahorse mating, see http://vimeo.com/76477081; seahorse males giving birth can be seen at https://www.youtube.com/watch?v=fDfRIIM5iHQ, at https://www.youtube.com/watch?v=2eIuXxp0sxM, and at https://www.youtube.com/watch?v=BPlnqckOPdY; all accessed Sept. 14, 2014.

Chapter 3: Picky Females, Easy Males

69
Darwin quotations on sexual selection:
Darwin,
Origin
, 97–99. For those consulting other editions than the one cited above, the quotes occur a few pages into chapter 4.

71
But tactless wording aside, was Darwin right?:
For a dissenting view, see Joan Roughgarden’s
Evolution’s Rainbow: Diversity, Gender, and Sexuality in Nature and People
(Berkeley: University of California
Press, 2013). Roughgarden is a distinguished evolutionary ecologist who makes a frontal (and, in the consensus view, greatly exaggerated) assault on the theory of sexual selection as presented by Darwin and greatly improved in the past half century. The book should be read with reviews by two other leading evolutionary biologists, Sarah Blaffer Hrdy (
Nature
429 [2004]: 19–21) and Alison Jolly (
Science
304 [2004]: 965–66). Hrdy cautions, “Competition between those of one sex for reproductive access to the other remains a robust explanatory framework, even though it is not the whole story” (p. 21); Jolly writes, “What Darwinian theory needs is not so much radical revision as a simple expansion to take sexual diversity much more seriously” (p. 966). Both admire, as do I, Roughgarden’s call for greater attention to the variety of sexual arrangements in the animal and human worlds; I try to pay that attention here.

71
Darwin went on to write a book about all this:
Charles Darwin,
The Descent of Man, and Selection in Relation to Sex
(1871; rpt.; New York: Penguin Classics, 2004).

72
“Where one sex invests considerably more”:
R. L. Trivers, “Parental Investment and Sexual Selection,” in
Sexual Selection and the Descent of Man
, 1871–1971, ed. B. Campbell (Chicago: Aldine, 1972), 173.

72
consider the cassowary:
Olivia Judson, “Big Bird,”
National Geographic
224, no. 3 (2013): 60–77. For more scientific detail, see L. A. Moore, “Population Ecology of the Southern Cassowary
Casuarius casuarius johnsonii
, Mission Beach, North Queensland,”
Journal of Ornithology
148, no. 3 (2007): 357–66. For a superb documentary, see David Attenborough’s
Cassowaries,
produced by Bianca Keeley for the BBC Natural World series.

73
“His chicks .
. .
make funny whistling-peeping sounds”:
Judson, “Big Bird,” 67–68.

73
“I’m coming back as a female cassowary”:
Ibid., 67.

73
Jacanas:
Stephen T. Emlen and Peter H. Wrege, “Division of Labour in Parental Care Behaviour of a Sex-Role-Reversed Shorebird, the Wattled Jacana,”
Animal Behaviour
68, no. 4 (2004): 847–55; see also the same authors’ “Size Dimorphism, Intrasexual Competition, and Sexual Selection in Wattled Jacana (
Jacana jacana
), a Sex-Role-Reversed Shorebird in Panama,”
Auk
121, no. 2 (2004): 391. A dramatic video of jacana reproductive life was available on Sept. 12, 2014, at https://www.youtube.com/watch?v=OitI6o076rY. It’s
an excerpt from episode 12 of
The Trials of Life,
narrated by David Attenborough, BBC, 1990.

74
Jacana female an egg-making machine:
Emlen quoted by the Evolution Library, PBS, http://www.pbs.org/wgbh/evolution/library/01/6/l_016_04.html, accessed Sept. 12, 2014.

74
Eight thousand species of pair-bonding birds:
A. Cockburn, “Prevalence of Different Modes of Parental Care in Birds,”
Proceedings of the Royal Society B: Biological Sciences
273, no. 1592 (2006): 1375–83.

77
peahens lay more eggs for males with larger trains:
M. Petrie, T. Halliday, and C. Sanders, “Peahens Prefer Peacocks with Elaborate Trains,”
Animal Behavior
41 (1991): 323–31; Marion Petrie, “Improved Growth and Survival of Offspring of Peacocks with More Elaborate Trains,”
Nature
371, no. 6498 (1994): 598–99; and M. Petrie, P. Cotgreave, and T. W. Pike, “Variation in the Peacock’s Train Shows a Genetic Component,”
Genetica
135, no. 1 (2009): 7–11.

78
Zahavi’s “handicap principle”:
A. Zahavi, “Mate Selection: A Selection for a Handicap,”
Journal of Theoretical Biology
53 (1975): 205–14; Amotz Zahavi,
The Handicap Principle: A Missing Piece of Darwin’s Puzzle
(New York: Oxford University Press, 1999).

78
Fisher’s “runaway selection” theory:
Ronald A. Fisher,
The Genetical Theory of Natural Selection
(1930; repr., New York: Dover, 1958).

78
Peahens’ preference for elaborate peacock tails:
Roslyn Dakin and Robert Montgomerie, “Eye for an Eyespot: How Iridescent Plumage Ocelli Influence Peacock Mating Success,”
Behavioral Ecology
24, no. 5 (2013): 1048–57.

78
Tail length in male barn swallows:
A. P. Møller, A. Barbosa, J. J. Cuervo, F. de Lope, S. Merino, and N. Saino, “Sexual Selection and Tail Streamers in the Barn Swallow,”
Proceedings of the Royal Society: Biological Sciences
265 (1998): 409–14; and Y. Vortman, A. Lotem, R. Dor, I. J. Lovette, and R. J. Safran, “The Sexual Signals of the East-Mediterranean Barn Swallow: A Different Swallow Tale,”
Behavioral Ecology
22, no. 6 (2011): 1344–52.

79
Palmate newt filament length:
Jérémie H. Cornuau, Margaux Rat, Dirk S. Schmeller, and Adeline Loyau, “Multiple Signals in the Palmate Newt: Ornaments Help When Courting,”
Behavioral Ecology and Sociobiology
66, no. 7 (2012): 1045–55.

79
The Trinidadian guppy:
Kenji Karino, Miho Kobayashi, and
Kazuhiro Orita, “Adaptive Offspring Sex Ratio Depends on Male Tail Length in the Guppy,”
Ethology
112, no. 11 (2006): 1050–55.

79
In the mandrill:
Joanna M. Setchell, “Do Female Mandrills Prefer Brightly Colored Males?”
International Journal of Primatology
26, no. 4 (2005): 715–35.

79
even in our relatively pair-bonding species:
Alan F. Dixson, Barnaby J. Dixson, and Matthew Anderson, “Sexual Selection and the Evolution of Visually Conspicuous Sexually Dimorphic Traits in Male Monkeys, Apes, and Human Beings,”
Annual Review of Sex Research
16 (2005): 1–19.

79
Roseate tern female streamers:
Brian G. Palestis, Ian C. T. Nisbet, Jeremy J. Hatch, Jennifer M. Arnold, and Patricia Szczys, “Tail Length and Sexual Selection in a Monogamous, Monomorphic Species, the Roseate Tern
Sterna dougallii,
”
Journal of Ornithology
153, no. 4 (2013): 1153–63.

79
Courtship feeding:
David Lack, “Courtship Feeding in Birds,”
Auk
57, no. 2 (1940): 169–78; I. Galvan and J. J. Sanz, “Mate-Feeding Has Evolved as a Compensatory Energetic Strategy That Affects Breeding Success in Birds,”
Behavioral Ecology
22, no. 5 (2011): 1088–95.

80
Ringdove courtship:
Daniel S. Lehrman, “The Reproductive Behavior of Ring Doves,”
Scientific American
211, no. 5 (1964): 48–54; Kristen E. Mantei, S. Ramakrishnan, P. J. Sharp, and J. D. Buntin, “Courtship Interactions Stimulate Rapid Changes in GnRH Synthesis in Male Ring Doves,”
Hormones and Behavior
54, no. 5 (2008): 669–75. A brief video, accessed Sept. 12, 2014, can be seen here: https://www.youtube.com/watch?v=kOoSDWVM8f8.

81
Rat courtship:
S. A. Barnett,
The Rat: A Study in Behavior
(Chicago: University of Chicago Press, 1975).

81
“Sex with Knockout Models”:
Emilie F. Rissman, Scott R. Wersinger, Heather N. Fugger, and Thomas C. Foster, “Sex with Knockout Models: Behavioral Studies of Estrogen Receptor Alpha,”
Brain Research
835, no. 1 (1999): 80–90.

82
Black grouse leks:
Anni Hämäläinen, Rauno V. Alatalo, Christophe Lebigre, Heli Siitari, and Carl D. Soulsbury, “Fighting Behaviour as a Correlate of Male Mating Success in Black Grouse
Tetrao tetrix
,”
Behavioral Ecology and Sociobiology
66, no. 12 (2102): 1577–86.

82
Uganda kob:
Andrew Balmford, “Social Dispersion and Lekking in Uganda Kob,”
Behaviour
120, no. 3–4 (1992): 177–91.

83
Consider the majestic red deer:
T. H. Clutton-Brock, F. E. Guinness, and S. D. Albon,
Red Deer: Behavior and Ecology of Two Sexes
(Chicago: University of Chicago Press, 1982). A video of this herd during the rutting season can be seen here: https://www.youtube.com/watch?v=SZCwezLt7Ks, accessed Sept. 12, 2014.

83
the more basso the croak:
David Lesbarrères, Juha Merilä, and Thierry Lodé, “Male Breeding Success Is Predicted by Call Frequency in a Territorial Species, the Agile Frog (
Rana dalmatina
),”
Canadian Journal of Zoology
86, no. 11 (2008): 1273–79.

84
Elephant seals:
B. J. Le Boeuf, “Male-Male Competition and Reproductive Success in Elephant Seals,”
American Zoologist
14 (1974): 163–76. For a video, see http://channel.nationalgeographic.com/channel/videos/elephant-seal-wars/, “Race to Survive: Elephant Seal Wars,” National Geographic Channel, accessed Sept. 12, 2014.

84
4 percent of the males:
B. Le Boeuf and R. Peterson, “Social Status and Mating Activity in Elephant Seals,”
Science
163 (1969): 91–93.

85
Mammals with larger females:
Katherine Ralls, “Mammals in Which Females Are Larger Than Males,”
Quarterly Review of Biology
51 (1976): 245–76.

86
“Once female mammals became committed”:
Ibid., 262.

87
Platypus milk proteins:
Wesley C. Warren and 101 other authors, “Genome Analysis of the Platypus Reveals Unique Signatures of Evolution,”
Nature
453, no. 7192 (2008): 175–83.

87
Oliver Krüger’s agility comparison:
Oliver Krüger, “The Evolution of Reversed Sexual Size Dimorphism in Hawks, Falcons and Owls: A Comparative Study,”
Evolutionary Ecology
19, no. 5 (2005): 467–86.

87
Bringing Ralls up to date on mammals:
P. Lindenfors and B. S. Tullberg, “Evolutionary Aspects of Aggression: The Importance of Sexual Selection,”
Advances in Genetics
75 (2011): 7–22.

88
Holekamp’s comprehensive 2011 review:
K. E. Holekamp, J. E. Smith, C. C. Strelioff, R. C. Van Horn, and H. E. Watts, “Society, Demography and Genetic Structure in the Spotted Hyena,”
Molecular Ecology
21, no. 3 (2012): 613–32.

88
“Adult females are larger”:
Ibid., 614.

89
“Assumption of the receptive stance”:
Micaela Szykman, Russell C. Van Horn, Anne L. Engh, Erin E. Boydston, and Kay E. Holekamp, “Courtship and Mating in Free-Living Spotted Hyenas,”
Behaviour
144, no. 7 (2007): 828.

89
“There is no external vagina”:
Ibid., 816.

90
Consider the small tropical fish:
M. E. Cummings, “Looking for Sexual Selection in the Female Brain,”
Philosophical Transactions of the Royal Society B: Biological Sciences
367, no. 1600 (2012): 2348–56.

92
The importance of oxytocin:
C. S. Carter, “Oxytocin Pathways and the Evolution of Human Behavior,”
Annual Review of Psychology
65 (2014): 17–39.

92
Male fidelity and fatherhood:
T. R. Insel, “The Challenge of Translation in Social Neuroscience: A Review of Oxytocin, Vasopressin, and Affiliative Behavior,”
Neuron
65, no. 6 (2010): 768–79.

92
How the gene controls receptors in the emotional brain:
Zoe R. Donaldson and Larry J. Young, “The Relative Contribution of Proximal 5' Flanking Sequence and Microsatellite Variation on Brain Vasopressin 1a Receptor (Avpr1a) Gene Expression and Behavior,”
PLoS Genetics
9, no. 8 (2013): e1003729.