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Nature's Nether Regions Page 29


  ant female sperm storage, 83

  apes. See also humans

  clitoris, 76, 188–89

  emasculation of rivals, 185

  homosexuality, 188–89

  mating plugs, 141, 143–44

  penile spines, 155, 156

  Argentine lake ducks. See ducks

  argonaut autonomous hectocotylus, 23

  Arnqvist, Göran, 152–54, 185

  artificial insemination of livestock, 73, 133, 183–84

  Australian Veterinary Journal, 79

  bacterial sex, 9

  Bagemihl, Bruce, 126

  Baker, Robin, 59–61, 78, 82

  banana flies

  cheap sperm/expensive egg principle, 48

  female reproductive maze, 71, 72

  lock-and-key genital match, 40–41

  manipulative seminal proteins, 145–47, 149

  rare-male effect, 94

  Barker, David, 63–64

  barnacle penis length, 44, 45

  Bateman’s cheap sperm/expensive egg principle, 47–48, 64, 81, 115, 161

  bat female sperm storage, 82–83

  bedbug traumatic insemination, 130–33, 189

  beetles

  cryptic female choice, 73

  erratic genital evolution, 98

  female reproductive maze, 71, 72

  fossil studies of, 96–98

  manipulative seminal proteins, 147, 149, 154

  negative allometry of penis, 101–2, 102

  parameres, 56

  paternity by proxy, 118

  penile diversity, 30–31, 33, 36, 39–40, 43

  penile spines, 150–54, 151

  potential speciation events, 188

  sexually antagonistic coevolution, 120–23, 124

  sperm scooping, 117–18

  Bellis, Mark, 59–61, 78, 82

  Bernasconi, Giorgina, 86–87

  Biological Exuberance (Bagemihl), 126

  birds. See also ducks

  artificial insemination of chickens, 133

  cloacal-kiss mating, 58

  colorful male plumage, 42–43, 49

  natural versus sexual selection, 16–17

  sensory drive, 51–53, 52, 58

  sperm dumping, 63

  Birkhead, Tim, 77, 126–28

  Blanchard, Katrina, 167

  Brennan, Patricia, 127–28, 183

  Brillard, Jean-Pierre, 127

  broadcast spawners in marine environment, 19

  Bruce effect, 87–88

  Burch, Rebecca, 116–17, 147–48

  Burger, Matthias, 61–62

  Burley, Nancy, 51–52

  Burton, Frances, 78

  by-product hypothesis of female orgasm, 81

  caltrop cornuti, 151, 152

  “Can Shoe Size Predict Penile Length?” (Shah and Christopher), 102–3

  Cardoso, Maria Fernanda, 35

  Carnahan, Sarah, 143

  Carosi, Monica, 79–80

  Case of the Female Orgasm, The (Lloyd), 81

  cave beetle penis diversity, 30–31, 39–40

  Chapman, Tracey, 6, 149

  Chase, Ronald, 166–67

  cheap sperm/expensive egg principle, 47–48, 64, 81, 115, 161

  chickens

  artificial insemination of, 133

  sperm dumping, 63

  chimpanzees

  clitoris, 76

  emasculation of rivals, 185

  genitalia of, in contrast to humans’, 5

  mating plugs, 141, 143–44

  penile spines, 155, 156

  Christopher, Nimal, 102–3

  Civetta, Alberto, 149

  Clark, Andrew, 149

  clitoris

  bonobo, 188–89

  chimpanzee, 5

  debate concerning function of, 73–74

  embryonic development of, 76

  human, 5, 74–75, 75, 77

  nonhuman mammals, 76–77

  spotted hyena, 76–77

  clonal reproduction, 11–12

  Coope, Russell, 97–98

  copulation

  within array of cryptic female choices, 64–66

  dry, 66–73, 69

  internal fertilization in land-based animals, 18–19

  copulatory courtship, 66–73, 69

  copulatory stridulation, 55, 56

  copulatory vigor, 70

  Córdoba-Aguilar, Alex, 113–14

  cost-of-eggs principle, 47–48, 64, 81, 115, 161

  cow uterine contractions, 77–78

  crane fly sensory drive, 54–55, 56

  Craze, Paul, 182

  cricket and katydid sperm flushing, 119–20

  Crudgington, Helen, 152–53

  cryptic choice in hermaphrodites, 175–76

  cryptic female choice

  as agent of genital evolution, 114–15, 131–33, 185–86

  cost-of-eggs principle, 47–48, 64, 81, 115, 161

  dry copulation, 66–73, 69

  embryo abortion, 87–88

  female collaboration in sperm scooping, 62, 119–20, 187

  female orgasm and upsuck hypothesis, 61, 78–80

  female reproductive maze, 71–73, 72

  internal courtship, 57, 62, 63, 66–73

  limitation by natural selection, 104–7

  manipulative seminal proteins and, 148

  overriding by male, female’s benefit from, 116

  as range of selection filters, 64–66, 122

  rare-male effect, 93–95

  sensory drive, 42, 51–58, 52, 56, 73, 94, 144

  sexually antagonistic coevolution and, 113–15

  sperm dumping, 59–64, 78, 120, 144, 187

  sperm storage and selective sperm use, 82–87, 86

  cryptic thrusting, 57

  Cuvier, Georges, 23, 31, 35–36

  damselflies

  acrobatic mating, 99–100, 109–10

  egg-mimicking penis, 113–14

  erratic genital evolution, 99–100

  female sperm storage, 113

  sexually antagonistic coevolution, 113–14

  sperm scooping, 2, 109–12, 111

  vaginal sense organ, 58, 113

  dart shooting

  chemical manipulation of female, 165, 166–67

  deployment of darts, 163–64

  diversity of darts, 164, 165, 167–68

  nuptial gift hypothesis, 164–66

  Darwin, Charles

  Descent of Man, and Selection in Relation to Sex, The, 2, 15, 45–46, 161

  on evolution as gradual process, 99

  on female preference for male beauty, 49

  on hermaphroditic sex, 161–62

  on male wooing versus bullying, 48

  on natural versus sexual selection, 2, 16, 43

  on primary and secondary sexual characteristics, 15–16, 17

  silence on genital diversity, 43–46

  study of beetles and barnacles, 44, 45

  Victorian constraints on, 46, 49

  Darwin, Henrietta “Etty,” 45–46

  Dawkins, Richard, 3, 115

  Dawn of the Deed, The (Long), 96

  Dean, Rebecca, 63

  de Graaf, Reinier, 74

  Descent of Man, and Selection in Relation to Sex, The (Darwin), 2, 15, 45–46, 161

  Dewsbury, Donald, 66

  di Palma, Antonella, 135–36

  Dixson, Alan, 141, 155–56

  DNA

  cellular organelles and sexual reproduction, 13–14

  in determination of pace of genital evolution, 99–100

&
nbsp; harmful mutations and genetic defects, 12, 88

  human evolutionary loss of penile spines, 156

  methods for exchange of, 9–10, 18–19

  of seminal proteins, 146

  in synthesis of mating plugs, 143–44

  Dr. Tatiana’s Sex Advice to All Creation (Judson), 82

  dry copulation, 66–73, 69

  ducks

  forced copulation, 126

  homosexuality and necrophilia, 125–26

  matched complexity of penis and vagina, 127–28

  penis length, 126–27

  sexually antagonistic coevolution, 127–28, 129, 180–82

  Düngelhoef, Susanne, 56

  dung-fly female sperm storage and selective sperm use, 84–87, 86

  Eberhard, Bill

  on copulatory courtship, 67

  on cryptic female choice, 64, 65, 185–86

  on cryptic thrusting, 57

  on Darwin’s failure to address genital diversity, 44

  definition of genitalia, 18

  Female Control, 65

  on genital allometry, 103

  on genital complexity and rapid evolution, 3–6, 95

  on lock-and-key mating hypothesis, 39–40

  on penis as sensory courtship device, 53, 54–55

  on potential benefit of genitalia research, 184

  Sexual Selection and Animal Genitalia, 4–5, 39–40

  ebony jewelwing damselfly sperm scooping, 109–12, 111

  elephant shrew penile diversity, 32–34

  embryo abortion, 87–88, 144

  evolution, genital. See genital evolution

  Evolution and Human Behavior journal, 117

  Evolution of Human Sexuality, The (Symons), 80

  female choice. See cryptic female choice

  Female Control (Eberhard), 65

  female orgasm. See orgasm, female

  female sperm storage. See sperm storage

  Field Guide to the Birds of West Malaysia and Singapore, A (Jeyarajasingam), 42

  “First Case of Homosexual Necrophilia in the Mallard . . . , The” (Moeliker), 125–26

  Fisherian versus Good-Gener debate, 49–51

  flagella, penile, 121–23, 124

  flies. See insects

  flour beetle sperm scooping, 117–18

  flowback. See sperm dumping

  Fooden, Jack, 37

  fossil studies, 96–98

  Fox, C. A., 79

  Freude, Heinz, 43

  fruit flies. See banana flies

  Gack, Claudia, 121–23

  Gallup, Gordon, 116–17, 147–48

  Gelhaus, Jon, 54–55

  General Outline of the Organisation of the Animal Kingdom (Jones), 163

  genital evolution. See also cryptic female choice; sexually antagonistic coevolution; sexual selection; sperm competition

  fossil studies on, 96–98

  genital allometry, 103

  gradual versus erratic pace of, 98–100, 107

  possible role of homosexuality, 188–89

  potential speciation events, 188

  rapid process of, 5, 95, 160

  reproductive selfishness of individual, 3

  genitalia. See also specific genital features; specific genital functions

  definition of, 18

  diversity and complexity of, 3–6, 30–35, 33

  embryonic development, 76

  homologous clitoris and penis, 81

  human fascination with, 1, 6, 184–85

  internal fertilization and, 18–19

  lock-and-key mating hypothesis, 36–41

  negative allometry, 101–3, 102

  substitutes, 19–26, 26, 132

  genitalia research

  cultural biases and stereotypes, 38, 186

  dearth of studies on females, 38, 81–82

  insights afforded by, 186–89

  practical applications, 183–84

  ridicule of, 183

  as scientific discipline, 2–3, 6, 112

  taxonomy, 5, 34, 37, 159

  genital stridulation, 55, 56

  genital thrusting and movement, 57–58, 62, 63, 67

  Gerhardt, Ulrich, 172–75

  Ghiselin, Michael, 17

  Ginsberg, Joshua, 62–63

  Gittenberger, Edmund, 177

  gonochorist animals, definition of, 156–57

  Gosse, Philip Henry, 36

  Gould, Stephen Jay, 81, 98–99

  Gowaty, Patricia Adair, 186

  guinea pigs

  artificial insemination of, 133

  mating plugs, 140–41

  guppies and related fish

  positive allometry of penis, 104–5

  rare-male effect, 93–95

  Haase, Martin, 169–70

  hamster female reproductive maze, 71

  Harde, Karl Wilhelm, 43

  Hays, Ray, 77–78

  hectocotylus, function of, 22–23

  Hellriegel, Barbara, 86–87

  Heredity journal, 47

  hermaphrodites

  cheap sperm/expensive egg principle, 161

  cryptic choice, 175–76

  dart shooting, 163–68, 165

  exaggerated genitalia and complex sexual behaviors, 159–63, 171–75, 174, 182

  mirror-image genitals, 176–82, 181

  mutual insemination, 158–60, 159

  penis detachment, 170–71

  preference for male role, 163, 175, 182

  self-fertilization, 158

  sexually antagonistic coevolution, 167–68, 175–76, 182

  sperm storage, 83

  sperm sucking, 170

  Historia Animalium Angliae (Lister), 172

  Holwell, Greg, 39

  homosexuality, 125–26, 131, 188–89

  Hotzy, Cosima, 153–54

  Hubbell, Stephen, 4

  Huber, Bernhard, 103

  Hubweber, Lasse, 150–51

  humans

  clitoris, 5, 74–75, 75, 76, 77

  evolutionary loss of penile spines, 154, 156

  female orgasm, 75–76, 77, 79, 80

  free-swimming sperm in female body cavity, 133–34

  genitalia of, in contrast to chimpanzees’, 5

  manipulative seminal proteins, 147–48

  mating plug consistency, 141, 143

  miscarriage, 88

  negative allometry of penis, 102

  possible evolutionary role of homosexuality, 188–89

  potential applications of genitalia research for, 184

  semen flowback, 59–61, 78

  sperm scooping, 116–17

  vaginal sense organs, 58

  viability of ejaculated sperm in vagina, 82, 116

  Human Sexual Response (Masters and Johnson), 77

  Hunter, John, 15

  hyena clitoris, 76

  hypodermic (traumatic) insemination, 128–34, 130, 163

  insects. See also beetles

  acrobatic mating, 99–100, 110

  cheap sperm/expensive egg principle, 48

  chiral genitalia, 39

  erratic genital evolution, 99–100

  female reproductive maze, 71, 72

  female sperm storage and selective sperm use, 83, 84–87, 86

  fossil studies of, 96–98

  homosexuality, 131, 189

  internal courtship, 58

  lock-and-key genital match, 40–41

  manipulative seminal proteins, 145–49

  penile complexity and diversity, 4, 31–32, 37, 38

  penile thorns, 151, 152

  rare-male
effect, 94

  sexually antagonistic coevolution, 113–14

  sexual selection, 90–93

  sperm flushing, 119–20

  sperm scooping, 2, 109–12, 111

  stridulation, 42, 55, 56

  traumatic insemination, 130–33

  vaginal sense organs, 58, 113

  internal courtship

  dry copulation, 66–73, 69

  female orgasm and, 77

  female sensory receptors, 58

  genital thrusting and movement, 57–58, 62, 63, 67

  mating plugs for, 144

  penile features for, 54, 57

  International Journal of Tropical Biology, 54–55

  Iwasa, Yoh, 92–93

  Jeannel, René, 29–31

  Jeyarajasingam, Allen, 42

  Johnson, Virginia, 77, 101

  Jones, Thomas Rymer, 163

  Journal of Evolutionary Biology, 182

  Journal of Experimental Biology, 166

  Journal of Molluscan Studies, 166

  Journal of Zoology, 25, 33–34

  Judson, Olivia, 82

  Käfer Mitteleuropas, Die (Freude, Harde, and Lohse), 43

  Kamimura, Yoshitaka, 40–41

  Karlsson, Anna, 169–70

  Kew, H. Wallis, 172

  Kingan, Sarah, 143

  Kingsley, David, 156

  Knoflach, Barbara, 140

  Kobelt, Georg Ludwig, 74, 75

  Kočárek, Petr, 36

  Koene, Joris, 158, 161–62, 166, 168, 176

  Kokko, Hanna, 94

  Koprowski, John, 145

  Krebs, John, 115

  Langerhans, Brian, 104–5

  lek paradox, 92

  leptodirin cave beetle penis diversity, 30–31, 39–40

  Leuckart, Karl Georg Friedrich Rudolf, 140

  Limax slug penis length, 171–75, 174

  Lister, Martin, 172

  Lloyd, Elisabeth, 81

  lock-and-key mating hypothesis, 36–41

  Lohse, Gustav Adolf, 43

  Long, John, 96

  longhorn beetle penis spines, 150–52

  love darts. See dart shooting

  Maan, Martine, 50

  macaques

  female orgasm, 79–80

  matched shapes of penis and vagina, 37

  miscarriage, 88

  McCracken, Kevin, 126

  Macgregor, Elizabeth Ann, 35

  MacLean, Paul, 53–54

  McPeek, Mark, 99–100

  Male and Female Lust Organs . . . (Kobelt), 74

  “Male Nipples and Clitoral Ripples” (Gould), 81

  mallard ducks. See ducks

  Marian, José Eduardo, 21

  Masters, William, 77, 101

  mating plugs

  consistency, 141, 143–44

  female cooperation with, 139, 145

  male body as, 140

  obstruction of future copulation, 136, 140

  pedipalps as, 136–40, 138