Mammalian Sexuality Alan F. Dixson Index More Information Www

Cambridge University Press 978-1-108-42618-3 — Mammalian Sexuality Alan F. Dixson Index More Information

Index

aardvark (Orycteropus afer), 41,44–45, 60 annexins, 235 testicular descent, 185 Anomaluromorpha, 35 accessory reproductive glands, 187 antagonistic coevolution, 225 ampullary glands, 189–192, 273 and sperm survival, 228–229 anatomical arrangement, 189 invertebrates bulbourethral glands (Cowper's glands), 273 chemical cues, 226–227 evolution of, 228–229 paragenital structures, 226 functions, 274 anteaters, 47,48–49 importance of, 198 Antechinus sp. monotremes, 207 A. agilis, 124–125 phylogenetic distribution, 187, 187 A. stuartii, copulatory courtship, 105 prostate and vesicular glands, 273–274 mating systems, 23, 94, 122–123 Acp70A, 226–227 penile morphology, 123–125, 124 acrosome reaction, 243 semelparity, 94, 122 relationship to sperm capacitation, 239 testis size, 122 Africa wild dog (Lycaon pictus), 127–128 Aotinae, testes size, 158 African civet (Civettictis civetta), 29 apes Afrosoricida, 45 baculum, 135 copulatory locks, 70 copulatory posture, 88 copulatory patterns testes size, 157. See also Primates spiny species, 80 apyrene sperm, 178 intromission durations aquatic animals spiny species, 80 and copulatory locks, 71 Afrotheria, 41–42, 41 copulatory patterns, 81 Afrosoricida, 45 Archibald, David, 50, 52 Hyracoidea, 44 armadillos, 47 Macroscelidea, 45–46 mating systems, 25–48 Proboscidea, 42 ovulation control, 251 RXFP2 and INSL3 genes, 185, 185 penile muscles, 150, 152 Sirenia, 42–44, 43 polyembryony, 48 Tubilidentata, 44–45 sperm rouleaux, 182 Age of Mammals, 279–280 artiodactyls, 25 alpaca, sperm transport and storage, 235–236, copulatory postures, 85 236 Assam macaque (Macaca assamensis), vaginal Alston’s brown mouse (Scotinomys teguina), morphology, 221 189–190 Atelinae, testes size, 158 American bison (Bison bison), 27 Australasian marsupials amphibious animals Dasyuromorphia, 21–23, 22 and copulatory locks, 71 Diprotodontia, 16–21 copulatory patterns, 81–83 Notorictemorphia, 14–16 Amphicyon, 141 Peramelemorphia, 23–48 ampulla, 189–192 aye-aye (Daubentonia madagascariensis), ampullary glands, 189–192, 273 196 phylogenetic distribution, 187, 188 copulatory pattern, 300

371

372 Index

Azara’s agouti (Dasyprocta azarae), penile beta nerve growth factor (β-NGF), 259–262, 275 morphology, 131–132 binary sperm, 172 birds baboons. See Papio sp. ovulation control, 252 Bactrian camel (Camelus bactrianus) retention of the phallus, 112 ovulation induction factor, 259 birth canal, marsupials, 209 pelvic thrusting pattern, 106 birth sex ratios, 239, 275 baculum (os penis), 134–135, 223, 269 black rhinoceros (Diceros bicornis), 26 anatomy, 135–136 black-footed ferret (Mustela nigripes), 81–82 androgenic control, 136 black-handed spider monkey (Ateles geoffroyi) biomechanical function, 138 copulatory plugs, 201 canids, 71, 127–128 vaginal morphology, 221 fossil evidence, 141–142 black-tailed jackrabbit (Lepus californicus), fractures, 140 copulatory pattern, 65 morphological diversity, 135 black-tailed prairie dog (Cynomys ludovicianus), mountain beaver (Aplodontia rufa), 35 copulatory pattern, 77 phylogenetic distribution, 134–135 black-wristed deer mouse (Peromyscus pinnipeds, 83 melanocarpus), copulatory plug, 201 projection of, 140–141, 270 blind mole-rat (Spalax ehrenbergi), 78 protection of the urethra, 140 bobuck (Trichosurus cunninghami), 20 role in induced ovulation, 140 body size selective forces for, 138–141 inﬂuence on copulatory patterns, 83–84 size intromission duration, 267–268 relationship to body mass, 136, 137 relationship to SEC length, 164, 164 relationship to copulatory pattern, 137–138, relationship to sperm length, 174, 271 137 bonobo (Pan paniscus) Balaenidae, copulatory pattern, 289 baculum, 135 banded mongoose (Mungos mungo), 30 copulatory posture, 88 bandicoots, 23–48 ejaculation intervals, 157–170 banner-tailed kangaroo rat (Dipodomys spectabilis), mating system, 40 copulatory plugs, 200–201, 201 testes size, 158 banteng (Bos javanicus), seminal pH, 231 vesicular glands, 194 Barbary macaque (Macaca sylvanus), copulatory bontebok (Damaliscus pygargus), copulatory pattern, 102 pattern, 85 Bathyergidae bottlenose squid (Sepioloidea magna), copulatory patterns, 77, 298 spermatophore transfer, 115 intromission duration, 77 Bovidae, 27 bats, 31–33 copulatory patterns, 63 baculum, 135 multiple partner matings, 98 copulatory courtship, 108 multiple paternity, 98 neck grips, 214 urethral processes, 130, 270 gloomy tube-nosed, 280 Bradipodidae, copulatory patterns, 302 hoary (Lasiurus cinereus), penile morphology, Brazilian tapir (Tapirus terrestris), penile 112 morphology, 112 long-term sperm storage, 246–247, 246–247 broad-footed marsupial mouse (Antechinus sp.), M. lucifugus copulatory pattern, 105 copulatory locks, 127 brown four-eyed opossum (Metachirus penile morphology, 126 nudicaudatus) M. myotis, pelvic thrusting pattern, 106 glans penis, 123 M. nattereri, 32 penile muscles, 153 mating systems, 97–98 brush-tail possum (Trichosurus vulpecula), 18–20, 19 ovulation control, 156, 251 ovulation control, 249 sperm storage, 98 brush-tailed bettong (Bettongia penicillata), testes size, 97–98, 156, 156 ovulation control, 249 beaked whales, testes size, 161–162, 162 bubonic plague (‘Black Death’), 280 beaver (Castor canadensis), 34 bulbocavernosus (BC) muscles, 149–152, 150, 270 bed bugs, paragenital structures, 226–227, 226 marsupials, 154

Index 373

bulbourethral glands (Cowper’s glands), 121, 273 cervical mucus anatomy, 189–190 effect of multiple intromissions, 102 phylogenetic distribution, 187, 187 effect on sperm transport, 231–233 role in seminal coagulation, marsupials, Cervidae 197–198 copulatory patterns, 62 bulbus glandis,71 multiple partner mating, 98 Burramyidae, 20 multiple paternity, 98 cervix, protective function, 225, 228 cactus mouse (Peromyscus eremicus), 107 cetaceans, 25 Callitrichidae accessory reproductive glands, 201 testes size, 158 copulatory patterns, 63–64, 81 Camelidae copulatory posture, 64, 87,87–88 copulatory patterns, 285 lumbar rigidity, 72 ovulation induction factors, 259–261, 261 mating systems, 98–99 Canadian river otter (Lontra canadensis), 82 pelvic bones, 119, 119 Canidae penile morphology, 119–120, 120 copulatory locks, 70,71 penile muscles, 149 copulatory pattern, 61,61 sperm mitochondria, 172 testes size, 159, 160 testes size, 119, 161–162, 162, 164 Caniformia, 28, 29 vaginal folds, 221–223, 222 copulatory locks, 127–128 Cetartiodactyla, 25, 27 intromission durations, 75 copulatory patterns, 62–64 capacitation. See sperm capacitation induced ovulation, 251 Cape ground squirrels (Xerus inauris), copulatory mating systems, 98 patterns, 77–78 pelvic thrusting, 72 capuchin (Cebus sp.), 39 chacma baboon (Papio ursinus), copulatory pattern, capybara (Hydrochoerus hydrochairis), testicular 66–67, 101–102 organization, 163 chacoan pecary (Catagonus wagneri), seminal pH, Carnivora, 27–30, 30 231 baculum, 135 Cheirogaleidae fossil evidence, 142 copulatory patterns, 299 size, relationship to copulatory pattern, 138 testes size, 158 copulatory locks, 70, 127–130 chelonians, ovulation control, 251–252 copulatory patterns, 63–64 chemical cues, 227–228 female reﬂexive responses, 214 invertebrates, 226–227 induced ovulation, 249 chemotaxis, 241–243 intromission durations, 75–76,75–76, 84 chevrotains (mouse deer), copulatory pattern, 63 neck grips, 214 chimpanzee (Pan troglodytes) testes size, 159–161, 160 baculum, 135 Castorimorpha, 35 copulatory posture, 88 cattle (Bos taurus), urethral process, 130 ejaculation intervals, 157–170 cavies mating system, 40, 168–169 copulatory patterns, 297–298 sexual skin swellings, 219–221, 220 mating systems, 97. See also guinea pig social organization, 97 Cebidae sperm counts, effect of frequent ejaculation, testes size, 158 168 Cercopithecinae testes size, 158 testes size, 158 vesicular glands, 194 cervical crypts, 232–233 chinchilla (Chinchilla lanigera) cervical grooves, 232 copulatory pattern, 298 cervical morphology, 223, 225 ovulation control, 256 and intrauterine insemination, 223 seminal pH, 231 and the baculum, 223 Chinese rock squirrel (Sciurotamias davidianus), and urethral processes, 223 penile morphology, 112 bovines, 225, 232 chinning behaviour, 214–216 macaques, 224, 232 Chiroptera, 31–33 relationship to penile morphology, 224 copulatory locks, 70

374 Index

Cingulata pattern no. 8, 68 copulatory patterns. See also armadillos phylogenetic distribution, 69–72 circadian rhythms, 257 copulatory patterns, 267–268 civets, 28, 30 and copulatory locks, 71 female reflexive responses, 214 classification, 58,60 clitoris, role of, 255 data sources, 59–60 clitoro-pelvic reflex, 212 Dewsbury scheme, 57 coati (Nasua nasua), seminal pH, 231 modified Dewsbury scheme, 57–59 coevolution, 217–219 female influences, 60 antagonistic, 225–229 influencing factors cervix, 223–225 aquatic vs. terrestrial copulations, female sexual skin swellings, 219–221, 220 57–83, 82 vaginal morphology, 221–223 body size, 83–84 Colobinae carnivorous vs. herbivorous mode of life, testes size, 158 75–76, 75–76 Colombian ground squirrels (Spermophilus induced ovulation, 257–259, 263, 268 columbianus), copulatory pattern, 76–77 intromission duration, 267–268 colugo (Galeopterus variegatus), 36–37, 37 multiple-partner matings, 268 common noctule bat (Nyctalus noctula), sperm predation risk, 268 storage, 156 spines, 79–81 common shrew (Sorax araneus), secondary sperm subterranean mammals, 76–79 storage, 193, 193 major patterns, 60–61 common wombat (Vombartus ursinus), 17,21 distribution in placental mammals and coprophagy, 36 marsupials, 66 copulatory courtship, 105, 211–212, 269 no. 3, 61–62,61–62 cervical morphology, 223–225 nos. 10, 11 and 12, 65–67 dasyurid marsupials, 105, 124–125 no. 16, 62,62–65 female influences, 108 minor patterns, 67 female reflexive responses, 212 no. 6, 67 carnivores, 214 no. 7, 67–68 human, 212–214 no. 8, 68 koala, 214 no. 9, 68 non-human primates, 213 no. 14, 68 mechanical fit, 217–219, 219 no. 15, 69 and female sexual skin swellings, 219–221, 220 primates, 99–100, 101–102 multiple intromission, 107–108 interspecies variability, 102 non-genital stimulation intraspecies variability, 101–102 immobilization behaviour, 214–215, 215 Macaca sp., 102 tactile stimulation, 216–217 prolonged intromission, 268–269 visual communication, 217 relationship to baculum size, 137, 137–138 patterns of pelvic thrusting, 106, 106 sperm competition strategies, 99 vaginal morphology, 221–223 frequent copulation, 99–100 copulatory frequency, human, 279 multiple intromission, 100–102 copulatory locks, 61, 104–105 primates, 100–102 and penile morphology, 125, 270 prolonged intromission, 100 bats, 127 rodents, 102–105 carnivores, 127–130 transition between, 67 Cricetidae, 125–127 variation between males, 74 platypus, 130 copulatory plugs, 151, 187, 223, 270, 274 spines, 145 functions, 201–203 and vesicular glands, 196 functions, 211 aquatic and amphibious animals, 71 importance of, 198 associated copulatory patterns, 71 masked palm civet, 68 brief locks, 72 murid rodents, 198 domestic dog, 61 primates, 198–199 pattern no. 6, 67 relationship to mating systems, 198, 200 pattern no. 7, 67–68 relationship to sexual dimorphism, 198

Index 375

removal of, 148 domestic dog, 232–234, 234 role of penile spines, 147–148 effect of cervical mucus, 231–233 rodents, 104 oviduct, 235 role of vesicular glands, 196 oviducts, 238–239 copulatory postures rapid phase, 229–231 cetaceans, 64 uterotubal junction, 234–235 dorsoventral vaginal, 209 marsupials, 84 strategies, 99 placental animals, 85–87, 86 Ctenomyidae ventro-ventral, 87 copulatory patterns, 77, 298 primates, 88 intromission duration, 77 courtship blind mole rat, 78 daily sperm production, 164, 165 crested porcupine, 79 Darwin, Charles, 3 echidnas, 79, 93 Dasypodidae, 47–48 naked mole rat, 78 dasyurids, copulatory courtship, 105 North American porcupine, 79–80. See also Dasyuromorphia, 21–22, 22 copulatory courtship intromission durations, 304 COVID-19, 280–281 semelparity, 23 Cowper, William, 121 Tasmanian devil, 22–23 Cowper’s glands. See bulbourethral glands Delphinidae coyote (Canis latrans), pelvic thrusting pattern, 106 copulatory patterns, 288 crested porcupine (Histrix cristata) mating systems, 27, 162. See also dolphins copulatory pattern, 80 Dermoptera, 36–37, 37 courtship, 79 intromission durations, 308 Cretaceous mammals, fossil evidence, 253–255 desmans, 33 Cricetidae, 35 Dewsbury, Donald, 57, 58 copulatory locks, 81, 125–127 dhole (Cuon alpinus), pelvic thrusting pattern, penile morphology, 125–127 106 cryptic female choice, 116, 211, 274 didelphids, 13 and copulatory courtship, 211–212 intromission durations. See also Virginia opossum cervical morphology, 223–225 (Didelphis virginiana) female reflexive responses, 212–214 Didelphimorphia, 12–14 mechanical fit, 217–221, 219 intromission durations, 304 non-genital stimulation, 214–217 Diprotodon, 24 vaginal morphology, 221–223 Diprotodontia, 16 and induced ovulation, 275 Burramyidae, 20 antagonistic coevolution, 225–227 feathertail glider, 20 associated traits, 278–279 intromission durations, 75–76 in fertilization, 243 koala, 16 masked palm civet, 68 macropods, 16–18 polyovulation, 245 Petauridae, 20–21 relationship to sperm competition, 89–90,89 possums, 18–20 rodents, 103 Pseudocheiridae, 21 role of oviducts, 275 Tarsipedidae, 20 sex-specific responses, 239, 275 Vombatidae, 21 sperm storage, 231 dire wolves (Canis dirus), baculum, 141, 142 cervical crypts, 232–233 dispersed mating systems, 18 long-term, 246–247 diurnality, 93 oviducts, 235–237, 238 dolphins uterotubal junction, 235–236 copulatory patterns, 81 sperm survival copulatory posture, 87 immune defences, 228 Ganges river dolphin, 119 pH challenges, 229 mating systems, 162 sperm transport penile morphology, 120 bovines, 232 seminal pH, 231 cervical grooves, 232 sperm morphology, 173

376 Index

dolphins (cont.) embryo resorption, 244, 245 testes size, 162, 162 Enaliarctos mealsi, 141 Yangtze River dolphin, 280 endangered species, 119, 280 domestic cat (Felix catus) use in traditional Chinese medicine, copulatory posture, 86 281 ovulation control, 256 Eomaia scansoria,51 somatosensory feedback, epididymis 257–259, 258 morphology, 165, 165 domestic dog (Canis familiaris) smooth muscle thickness, 170–171, 171 copulatory pattern, 61, 61–62 sperm maturation in, 165 male reproductive tract, 189–190 sperm storage, 167 penile morphology, 127 sperm transit times, 165–167, 167 sperm transport and storage, 232–234, 234 terminal segment cooling, 184 dorsolateral copulatory postures, 86 Equidae, 26–27 dorsoventral copulatory postures, 85 ergothioneine secretion, 189–192. See also horse marsupials, 84 ergothioneine, 189–192 Douglas squirrel (Tamiasciurus douglasii), vaginal Erinaceidae, 33, 74 morphology, 221 copulatory patterns, 294 dragging, and copulatory locks, 67–68 Euarchontoglires, 34 drill (Mandrillus leucophaeus) Dermoptera, 36–37, 37 male reproductive tract, 194 Lagomorpha, 35–36, 36 vesicular glands, 194 Primates, 37–41, 39 dromedary (Camelus dromedarius), copulatory Rodentia, 34–35, 34 posture, 85, 86 Scandentia, 37 dugong (Dugong dugon), 41,42–43 Eulipotyphla, 33–34 mating system, 43 copulatory patterns vaginal morphology, 221 spiny species, 79–80 induced ovulation, 250 eastern grey kangaroo (Macropus giganteus), 17, intromission durations 24, 75, 95 spiny species, 79–80 Eberhard, William, 3, 114, 116, 228 Eupleridae, 29–30 echidnas, 9, 9–10, 11, 93 copulatory pattern, 292 courtship, 79 eupyrene sperm, 178 epididymal morphology, 165 European river otter (Lutra lutra), 82 mating system, 93 extinction events, 3, 52, 253, 279–280 pelvic thrusting, 72, 106 extinctions penile morphology, 130 endangered species, 280 reproductive system, 11–12 human-related, 24, 280–281 female, links to male morphology, 209 recent species, 280 sperm bundles, 179, 179–180, 272 extra-pair matings, 92 ectospermalege, 226 ejaculation, 149, 152 fallopian tubes. See oviducts cavernosal pressures and electromyograms, 151 feathertail glider (Acrobates pygmaeus), 20 sperm transport, 169–171 mating system, 94 ejaculation intervals, 169 Felidae, 28–29 primates, relationship to mating systems, copulatory patterns, 64 157–170, 169–170 copulatory postures, 85, 86 elephant seal (Mirounga sp.), 82 female reﬂexive responses, 214 elephant shrews (sengis), 45–46 induced ovulation, 64 epididymal morphology, 165 360.110, 257–259 polyovulation, 245 ovulation control, 256 testicular positioning, 183 somatosensory feedback, 257–259, 258 elephants, 42 seminal pH, 231 copulatory patterns, 75 testes size, 159, 160 levator penis muscle, 152 Feliformia, 28–30, 30 mating system, 96 female behavioural responses, effect of penile penile movements, 72 spines, 147 seminal pH, 231 female defence polygyny, 91

Index 377

female reflexive responses, 212 duration of female receptivity, 146 carnivores, 214 penile morphology, testosterone-dependent human, 212–214 development, 136 koala, 214 lesser galago (Galago moholi), mating system, lordotic, 212, 218 96 role of penile spines, 147 testes size, 158 marmoset, effects of local anaesthesia, Ganges river dolphin (Platanista gangetica), 119 213 gelada (Teropithecus gelada), ejaculation intervals, non-human primates, 213 170 female reproductive system, 205–207, 208 genets, 29 antagonistic coevolution, 225–227 genital locks. See copulatory locks embryonic development, 210 Geomyidae links to male morphology, 207–211 copulatory patterns, 77, 298 female reproductive tract depths, intromission duration, 77 relationship to penile length, 217–219, 219 giant anteater (Myrmecophaga tridactyla), 47 effect of sexual skin swellings, 219–221 giant otter (Pteronura brasiliensis), 82 female sexual receptivity, effect of copulation, giant pangolin (Smutsia gigantea), 30–31 145–146 gibbons, 41 fertilization, 243–244, 244 intromission duration. See also Hylobates sp. fertilization sites, 237–238 giraffe, subspecies, 6–7 shrews, 237 Giraffidae, copulatory patterns, 286 fimbriae, 241 glans penis, marsupials, 121–122, 122 fish species, ovulation control, 251 gliders, 20–21 Flack, Gabriella, 60 gloomy tube-nosed bat (Munna tenebrosa), 280 Flannery, Tim, 14 Glyptodon, 50 flying foxes (fruit bats), 32,32 goat (Capra aegagrus hircus) follicular fluid, sperm chemotaxis, 241–243 seminal pH, 231 fossa (Cryptoprocta ferox), 29–30 urethral process, 130, 270 copulatory pattern, 68 golden lion tamarin (Leontopithecus rosalia), 100 penile morphology, 128 golden moles, 45 spines, 145 gorilla (Gorilla gorilla), 39 fossil evidence baculum, 135 baculum, 141–142 copulatory courtship, 108 bats, 31–32 ejaculation intervals, 170 Carnivora, 28 mating system, 40 Eulipotyphla, 33 penile morphology, 118 placental animals, 50–52, 253–255 testes size, 158 Sirenians, 43,43 vesicular glands, 194 Xenarthra, 49 Gracilinanus agilis, glans penis, 122 fossorial mammals, copulatory patterns, 76–79, 77 grasshopper mouse (Ochrotomys torridus), 67 frequent ejaculation greater bilby (Macrotis lagotis), 23 effect on sperm counts mating system, 94 chimpanzee, 168 greater glider (Petauroides volans), 21 human males, 168, 168 greater white-toothed shrew (Crocidura russula), sheep, 169 sperm storage, 237 recovery of non-ejaculated sperm, 193 green possum (Pseudocheirops archeri), 19 role of scrotum, 184 grey kangaroo (Macropus giganteus), 17 fruit bats (flying foxes), 32,32 grey seal (Halichoerus grypus), 83 fruit flies (Drosophila melanogaster), seminal grey short-tailed opossum (Mondelphis domestica) proteins, 226–227 copulatory posture, 84 fur seal (Arctocephalus forsteri), 106 ovulation control, 248–249 copulatory courtship, 216 ground squirrels, copulatory patterns, 76–79 fusion–fission social organization, 97 guinea pig (Cavia porcellus) copulatory plugs, 201, 201 galagos, 39 duration of female receptivity, 146 greater galago (Otolemur garnettii), 100 intromittent sac, 133–134, 146, 270 copulatory lock, 126 sperm rouleaux, 182

378 Index

Hamadryas baboon (Papio hamadryas), 100–101 human males, 192 harbour porpoise (Phocoena phocoena) ejaculation intervals, 170 copulatory pattern, 119 penile morphology, 117, 278 penile morphology, 120 prostate and vesicular gland sizes, 196 testis size, 119 sperm count, effect of frequent ejaculation, 168, harbour seal (Phoca vitullina), testicular 168 positioning, 183 testes size, 158, 158 hares, superfoetation, 247 traits associated with polygynandry, 279 hedgehogs vas deferens wall thickness, 191 Atelerix algirus, copulatory behaviour, human mating systems, 276–277 106 human migration, impact of, 281 Erinaceus europaeus, intromittent sac, 134 human population growth, 280–282 helper sperm (spermatozeugma), 178 human sexual dimorphism, 277 herbivores, intromission durations, 75–76,75–76, humpback whale (Megaptera novaeangliae) 84 mating system, 161 Herpestidae, 29–30 sperm length, 174 Heterocephalidae, copulatory patterns and testes size, 161, 162 intromission duration, 77 Hunter, Ronald, 205 Heteromyidae, 74 Hyaenidae, 29, 74 multiple-partner matings, 97 copulatory pattern, 62, 292 Hill, W.C. Osman, 116–117 spotted hyena (Crocuta crocuta), 30, 62, 128, 129 hippopotamus, 25, 27 copulatory lock, 129–130 copulatory pattern, 63, 285 female genitalia, 128–129, 129 Histricomorphia, 35 mating mechanics, 129 HIV/AIDS, 280 pelvic thrusting pattern, 106 hoary bat (Lasiurus cinereus), penile morphology, penile morphology, 129 112 testes size, 159, 160 Hoffman’s two-toed sloth (Choleopus hoffmanni), Hylobates sp. 47 ejaculation intervals, 170 Hominoidea, 40–41 testes size, 158. See also gibbons copulatory patterns, 302 hypodermic insemination, 226 testes size, 158 Hyracoidea, 44 honey possum (Tarsipes rostratus), 20 mating systems, 96 epididymal morphology, 165 sperm storage site, 193 mating system, 93–94 urethral process, 130 sperm length, 174 Hystricidae, copulatory patterns, 298 hopping mouse (Notomys alexis) Hystricognathi, intromittent sacs, 131–134, 133 copulatory lock, 126 copulatory pattern, 68 immobilization reflexes genitalic coevolution, 145 in hoofed animals, 214–216 penile morphology, 125–127, 126 neck grips, 214, 215 spines, 145 immune defences, female reproductive system, 228 horse (Equus caballus) Indarctos arctoides, baculum, 141 ergothioneine secretion, 189–192 Indian rhinoceros (Rhinoceros uncornis), 28 intrauterine insemination, 223 induced ovulation, 211, 248, 263 seminal pH, 231 and cryptic female choice, 275 testicular positioning, 183 and female reflexive responses, 214 house shrew (Suncus murinus), sperm storage, and oestrus stimulation, 252 236–237 and post-copulatory sexual selection, 262–263 human evolution research, 276 and social organization, 252–253 comparative studies, 278–279 mole-rats, 253–254 mating systems, 276–277 copulatory patterns, 268 sexual dimorphism, 277 evolutionary origins, 248, 255, 263–264, human females 274–275 reflexive responses, 212–214 felids, 64 role of orgasm, 214 lagomorphs, 36, 65 vaginal pH, 229, 230 multiple paternity, 262, 262

Index 379

neuroendocrine mechanisms, 256 lactobacilli, vaginal, 229 overlap with spontaneous ovulation, 256–257, Lagomorpha, 35–36, 36 263 copulatory patterns, 62, 62 ovulation induction factors, 85, 259–262, induced ovulation, 249 275 large vesper mouse (Calomys callosus), copulatory phylogenetic distribution, 249–250, 264 plug, 201 marsupials, 248–249 lateral vaginae, marsupials, 207–209 non-mammalian vertebrates, 251–252 sperm transport, 209 placental animals, 249–251 tammar wallaby, 209 uncertainties, 249–251 Laurasiatheria relationship to testes size, 159–160 Carnivora, 27–30, 30 rodents, 105 Cetartiodactyla, 25, 27 somatosensory feedback, 258 Chiroptera, 31–33, 32 multiple cues, 257 Eulipotyphla, 33–34 multiple intromission, 257–259 Perissodactyla, 25–27, 28 prolonged intromission, 259–260 Pholidota, 30–31, 31 role of baculum, 140 least shrew (Cryptotis parva), sperm storage, role of clitoral stimulation, 255 237 role of penile spines, 146 leeches, paragenital structures, 226 Insectivora, 33 lemurs, 39,40 INSL3 gene, 185 ejaculation intervals, 170 internal courtship device, phallus as, 116, 125, 149, Eulemur fulvus, penile morphology, 118 211–212, 223, 269–271 testes size, 158 International Union for the Conservation of Nature Leonard, Janet, 3 (IUCN) Red List, 280 Leporidae, 35–36, 36 intrauterine insemination, 211, 223, 270, 274 lesser hedgehog tenrec (Echinops telfairi), baculum, cost to females, 225–226 134 intromission duration, 57–59 lesser mouse deer (Tragulus sp.), 27 and induced ovulation, 259–260 penile morphology, 112 carnivores vs. herbivores, 75, 75 levator penis (LP) muscle, 152–153 influencing factors Leydig cells, 163 body size, 83–84, 267–268 lion, ovulation control, 256 predation risk, 268 llamas, ovulation induction factor, 259–261, 261 relationship to baculum size, 137, 137–138 lock-and-key hypothesis, penile morphology, 114 intromittent sacs, 131–134, 270 long-beaked common dolphin (Delphinus morphological variability, 133 capensis), penile morphology, 120 invertebrates long-nosed bandicoot (Perameles nasuta), chemical cues, 226–227 copulatory posture, 84 paragenital structures, 226 long-tailed macaque (Macaca fascicularis), inverted copulatory postures, 86–87 copulatory pattern, 66–67, 102 ischiocavernosus (IC) muscles, 149, 150, 270 long-tailed pangolin (Phataginus tetradactyla), 31 marsupials, 154 long-term sperm storage, 246, 246–247 lordotic responses, 212, 218 jaguar (Panthera onca), 30 role of penile spines, 147 Javan rhinoceros (Rhinoceros sondaicus), 26 lorises copulatory courtship, 217 Kafka, Franz, 76 copulatory patterns, 300 kangaroos, 16–18 mating system, 96 copulatory posture, 84 penile morphology, 131 mating systems, 95 sperm rouleaux, 182 penile muscles, 153 testes size, 158 kinkajou (Potos flavus), copulatory courtship, 214–217 luteinizing hormone (LH), 255–257 kisspeptins, 256, 261 lynx (Lynx pardinus), seminal pH, 231 koala (Phascolarctos cinereus), 16, 17 female reflexive responses, 214 macaques induced ovulation, 211 cervical morphology, 224, 232 ovulation control, 248–249 cervical sperm migration, 232–233, 233

380 Index

macaques (cont.) pelvic thrusting, 72 copulatory patterns, 102 penile morphology, 121 phylogenetic distribution, 89–103 Antechinus sp., 123–125, 124 ejaculation intervals, 170 biﬁd glans penis, 122 M. assamensis, vaginal morphology, 221 penile muscles, 153 M. nigra, sexual skin swellings, 220 phylogeny, 15 Macropodidae, 16–18 seminal coagulation, 197–198 copulatory patterns, 283 size, 24 mating systems, 94–95 South American, 12 prolonged intromission, 106. See also kangaroos; Didelphimorphia, 12–14, 13 wallabies Microbiotheria, 14 Macroscelidea, 45–46 Paucituberculata, 14 copulatory patterns, 303 sperm morphology, 172 male urogenital mating protuberance (MUMP), sperm pairing, 180 131–132 masked palm civet (Paguma larvata), copulatory Mammalia, 6, 50 pattern, 68, 73 number of species, 6–7, 6–7 mating season duration, relationship to testes size, subclasses, 7–8 159 Marsupialia, 12 mating systems, 90–91 Monotremata, 9–12 and copulatory plugs, 198 Placentalia, 25–53 armadillos, 25–48 manatee classiﬁcation pitfalls, 91–92 copulatory posture, 87 dispersed, 18 mating system, 95 female multiple partner matings, 92–93 mandrill (Mandrillus sphinx) copulatory patterns, 268 ejaculation intervals, 170 monotremes, 93 sexual skin swellings, 220 placental animals, 95–99 vesicular glands, 194 Hominoidea, 40–41 Marcus Aurelius Antonius, 267 human, 276–277 marmoset (Callithrix jacchus) parentage studies, 92 epididymal morphology, 165 polygyny, 91–92 female sexual responses, 213 relationship to ejaculation interval, 169–170, 170 penile morphology, 118 relationship to penile morphology, 116–117 Marmosidae relationship to seminal coagulation, 198 copulatory patterns, 285 relationship to sperm counts, 205 penile morphology, 122, 122 relationship to sperm storage, 167 marsupial moles (Notorictes spp.), 14–16 relationship to testes size, 271 Marsupialia, 8 bats, 156 ancestral migrations, 12 cetaceans, 161–162 Australasian primates, 156–159, 158 Dasyuromorphia, 21–23, 22 terrestrial carnivores, 160–161 Diprotodontia, 16–21 relationship to vas deferens morphology, 191, Notorictemorphia, 14–16 192 Peramelemorphia, 23–48 relationship to vesicular gland size, 194–196, 195 copulatory courtship relationship to white cell counts, 226 neck grips, 215 Rodentia, 35 tactile stimulation, 216 Sirenians, 43 copulatory interactions, 271 sloths, 49 copulatory locks, 70 Mayr, Ernst, 114–116 copulatory patterns, 65 meerkat (Suricata suricatta), 29–30 copulatory postures, 84–85 Megachiroptera, 32,32 extinct species, 12–24 Megalictis ferox, 141 female reproductive system, 207–208, 208 Metatheria, 7–8, 12 links to male morphology, 209–211 Microbiotheria, 14 induced ovulation, 248–249, 250 Microchiroptera, 32,32 intromission durations, 75–76 mink (Mustela vison), 81–82 mating systems, 93–95 pelvic thrusting pattern, 106

Index 381

Miss Waldron’s red colobus monkey (Piliocolobus multiple intromission, 268 waldronae), 280 and induced ovulation, 257–259 mole-rats and sperm competition, 100–102 blind mole-rat, 78 blind mole-rat, 78 naked mole-rat, 34,78–79 copulatory plug removal, 147, 147 penile spines, 146 courtship role, 107–108 social organization and ovulation control, 254 number of mounts, 73 molecular studies, 52–53 phylogenetic distribution, 73–74 moles, 33 rodents, 102–105 Molina’s grass mouse (Akodon molinae), 67 sperm recovery studies, 104 copulatory pattern, 73 multiple-partner matings mongooses, 29 Antechinus sp., 122–123 monito del monte (Dromiciops gliroides), 14, 180 artiodactyls, 98 monkeys, 40 Cape ground squirrels, 78 Monodelphis domestica, pelvic thrusting pattern, females, 92–98 106 and penile morphology, 117, 118 Monotremata, 8–9, 9–10 copulatory patterns, 268 ancestors, 11 marsupials, 93–95 comparative anatomy, 11 monotremes, 93 copulatory locks, 70 placental animals, 95–99 copulatory patterns relationship to penile morphology, 269 spiny species, 79 multiple paternity, 262, 262 echidnas, 9, 11 Muridae, 35 female multiple partner matings, 93 copulatory locks, 70 female reproductive system, 206–207, 208 copulatory plugs, 198 links to male morphology, 207–209 muriqui (Brachyteles sp.), 149 intromission durations, 304 penile muscles, 150 spiny species, 79–80 Muroidea, copulatory patterns, 102–105 ovulation control, 248 musk deer (Moschidae), copulatory pattern, 63 platypus, 9–11, 10–11 musk shrew (Suncus murinus), sperm morphology, reproductive organs, 11–12 172, 173 sperm bundles, 179–180, 179 Mustelidae sperm morphology, 171–172 copulatory patterns, 81–82 montane vole (Microtus montanus), 259 and induced ovulation, 259–260 Moore, Carl, 182 testes size, 159, 160 moose (Alces alces), chinning behaviour, 216 musth, elephants, 42 mouﬂon (Ovis orientalis), 27 Myomorpha, 35 mountain beaver (Aplodontia rufa), 35, 138 mountain possum (Trichosurus cunninghami), 20 naked mole-rat (Heterocephalus glaber), mountain pygmy possum (Burramys parvus), 20 34,78–79 mouse lemur (Microcebus murinus) neck grips, 85, 214, 215 epididymal morphology, 165 nerve growth factor (NGF), 259–262, 275 mating system, 96 new species, 7 mouse (Mus musculus) New World monkeys, 39–40 baculum, 139 nine-banded armadillo (Dasypus novemcinctus), 25 copulatory pattern, 139 nocturnal primates penile morphology, 131 prolonged intromission, 100 sperm hooks, 181 testes size, 96 mouse opossum North American porcupine (Erithizon dorsatum) M. mexicana, glans penis, 122–123 copulatory pattern, 80 M. robinsoni, copulatory posture, 84 courtship, 79–80 mouse shrew (Myosurex varius), sperm storage, 237 northern pygmy mouse (Baiomys taylori), penile multimale–multifemale groups, 91–92 morphology, 125–127 primates, 96–97 northern right whale (Eubalaena glacialis) multiple brief intromissions, 65 copulatory posture, 87 multiple ejaculations, 59, 85 mating system, 162, 162 relationship to pregnancy rate, 107 testes size, 98

382 Index

northern short-tailed shrew (Blarina brevicauda) neuroendocrine mechanisms, 255–256 copulatory lock, 126, 221 phylogenetic distribution vaginal morphology, 221 marsupials, 248–249 Notoryctemorphia, 14–16 non-mammalian vertebrates, 251–252 copulatory patterns, 284 placental animals, 249–250 numbat (Myrmecobius fasciatus), 22 uncertainties, 249–251 ovulation induction factors, 259–262, 275 Obdurodon, 9–10 llamas, 261 Ochotonidae, 35–36, 36 ovulation, induced. See induced ovulation Octodontidae, copulatory patterns and intromission ovulation, spontaneous. See spontaneous duration, 77 ovulation oestrogen, role in sperm transport, 229 oxytocin, role in sperm transport, 229–230 oestrus stimulation, 252 okapi (Okapia johnstoni), 27 paca (Cuniculus paca), copulatory plugs, 201 Old World monkeys, 39 palm civet (Nandinia binotata), 28 orang-utan (Pongo sp.), 40–41 pandemics, 280 copulatory courtship, 108 COVID-19, 280–281 copulatory posture, 88 pangolins, 30–31, 31 penile muscles, 150 copulatory pattern, 74–75 subspecies, 7 copulatory posture, 86,86 orca (Orcinus orca), 27 ovulation control, 251 orgasm, female Papio sp., 39,64–65 human, 213–214 copulatory patterns, 66–67 non-human primates, 213 ejaculation intervals, 170 role of, 214, 255 P. hamadryas, sexual skin swellings, oriental short-clawed otter (Aonyx cinerea), 220 82 testes size, 158 Otariidae, 28, 74 paragenital structures, invertebrates, 226, 226 otter civet (Cynogale bennettii), 29 Parker, Geoffrey, 3 otters, copulatory patterns, 82 patas monkey (Erythrocebus patas) ova ejaculation intervals, 170 fertilization, 243–244, 244 penile morphology, 118 transport by fimbriae, 241 paternities, induced ovulators, 262, 262 ovaries, monotremes, 206–207 Paucituberculata, 14 overlap promiscuity, 18 pelvic thrusting oviduct length, 239–240, 241 functions of, 106–107 human, 279 patterns of, 106, 106 oviductal morphology, 240–241 Antechinus stuartii, 105 oviducts phylogenetic distribution, 72–73 cryptic female choice pinnipeds, 83 research issues, 275 penile erection sex-specific responses, 239, 275 cavernosal pressures and electromyograms, 151 sperm binding, 235, 237 morphological changes, 152 sperm storage, 235–237, 238, 246 penile lengths fertilization sites, 237–238 relationship to body weight, 278 shrews, 237 relationship to female anatomy, 217–219, 219 fimbriae, 241 effect of sexual skin swellings, 219–221 sperm transport, 235 penile morphology, 118, 211, 269 post-copulatory sexual selection, 238–239 and copulatory locks, 68, 71, 104–105, 125, 270 thermotaxis, 241 bats, 127 ovulation control, 263 carnivores, 127–130 and oestrus stimulation, 252 Cricetidae, 125–127 and social organization, 252–253 platypus, 130 mole-rats, 254 and intrauterine insemination, 270 continuum of, 256–257, 263 androgenic control, 136 fossil evidence, 253–255 baculum, 134–135 monotremes, 248 anatomy, 135–136

Index 383

biomechanical function, 138 photoperiodic cues, role in ovulation control, 257 fossil evidence, 141–142 Phyllostomidae, copulatory patterns, 294 phylogenetic distribution, 134–135 pica (Ochotona hyperborea), 36 selective forces for, 138–141 pig (Sus scrofa) size, 136–138, 137 copulatory pattern, 285 carnivores, 161 immobilization reﬂexes, 216 cetaceans, 119–120, 120 intrauterine insemination, 223 harbour porpoise, 120 mating mechanics, 130 changes during erection and ejaculation, 152 oviduct studies, 239–240 divergent evolution, 114–116, 211–212 seminal pH, 231 diversity, 112 Pilosa, 48–49 effect of bulbocavernosus contractions, 151–152 pinnipeds elongation, 72, 81, 119 aquatic vs. terrestrial copulations, 82–83,82–83 ﬁbroelastic and vascular types, 112–114 baculum, 138 human, 277–279 pelvic thrusting, 83 intromittent sacs, 131–134, 133 Pipistrellus sp., 31 marsupials, 121–125 Placentalia, 8 Antechinus sp., 124 Afrotheria, 41,41–46 links to female morphology, 209 copulatory postures, 85–87 monotremes, 207 dating origins of, 52 phylogenetic factors, 111–114, 113 fossil evidence, 50–52 primates, 116–117 molecular studies, 52–53 relationship to cervical morphology, 224 Euarchontoglires, 34 testosterone-dependent development, 136 Dermoptera, 36–37, 37 urethral processes, 130, 130, 152, 270 Lagomorpha, 35–36, 36 penile movements, 72 Primates, 37–41, 39 penile muscles, 149, 270 Rodentia, 34,34–35 bulbocavernosus, 149–152 Scandentia, 37 ischiocavernosus, 149 female multiple partner matings, 95–99 levator penis, 152–153 Laurasiatheria, 25–34 marsupials, 153 phylogeny, 25, 26 placental mammals, 150 Xenarthra, 46–49, 47, 50 penile spines, 128, 131–132, 132, 142, 269–270 plains mouse (Pseudomys australis), sperm associated receptors, 143–144 morphology, 172, 173 effects of removal, 143, 144, 149 plains viscacha (Lagostomus maximus) functions embryo resorption, 245 copulatory plug and semen removal, 147–148 polyovulation, 243–244 effect on female behavioural responses, 147 plains zebra (Equus quagga), 28 female tactile stimulation, 145–146 Planistidae, copulatory pattern, 288 role in copulatory locks, 145 platypus (Ornithorhynchus anatinus), 9–10,9–10, sensory feedback, 143 10–11 hormonal effects, 143 copulatory lock, 130 intromittent sacs, 131–132, 270 copulatory pattern, 67–68, 283 morphological categories, 142 female reproductive system, 206–207, 208 role in sexual selection, 145–148 links to male morphology, 207–209 testosterone-dependent development, 136 penile morphology, 130–131 Peramelemorphia, 23–48 reproductive organs, 11–12 Peramelidae, copulatory patterns, 284 pleiotropism hypothesis, penile morphology, 114–115 Perissodactyla, 25–27, 28 Plesiogulo marshalli, 141 Petauridae, 20–21 polyembryony, 48 Phalangeridae, copulatory patterns, 284 polygynandry, 91–92 Phascolarctidae and penile morphology, 117, 118 copulatory patterns, 284. See also koala Antechinus sp., 122–123 (Phascolarctos cinereus) associated traits, 278–279 Phocoenidae, copulatory pattern, 288 marsupials, 93–95 Pholidota, 30–31, 31 monotremes, 93 intromission durations, 307 placental animals, 95–99

384 Index

polygyny, 91–92 monkeys, 40 human, 277 penile morphology, 116–117 polyovulation, 46 phylogeny, 38 and cryptic female choice, 245 prosimians, 40 explanatory hypotheses social organization, 96–97 bet hedging, 245 sperm competition strategies selective abortion, 245 frequent copulation, 99–100 twinning, 244 multiple intromission, 100–102 occurrence, 243 prolonged intromission, 100 plains viscacha, 243–244 testes size, 156–159, 158 porcupines, 34 vesicular glands, 194, 196 copulatory patterns, 80 Proboscidea, 42 courtship, 79–80 copulatory patterns, 303 porpoises progesterone, induced secretion, 103, 107, 151 copulatory patterns, 81 prolactin, induced secretion, 103, 256–257, 269 penile morphology, 120 prolonged intromission, 59, 71, 105–106, 268–269 testes size, 119, 162, 162 and induced ovulation, 259–260 possums, 19 and sperm competition, 100 honey, 20 benefits, 105–106 pygmy, 20 patterns of pelvic thrusting, 106 ringtail, 21 Antechinus stuartii, 105 striped, 20 phylogenetic distribution, 74–75,74–75 post-copulatory sexual selection, 3, 155, 268 relationship to baculum size, 137–138, 137 and induced ovulation, 262–263 promiscuity, 92 baculum, 138–141, 139 and sperm competition. See also multiple-partner insect research, 4 matings penile spines, 145–146 prosimians, 38–39,39–40 sperm transport, 238–239. See also cryptic penile morphology, 131 female choice; sperm competition prostate gland, 273–274 post-ejaculatory intromission, 107 anatomy, 189–190 Potoroidae, copulatory patterns, 283 importance of, 198 potto (Perodicticus potto), 39 phylogenetic distribution, 187, 187 penile morphology, 131 role in seminal coagulation, 196–197 prairie vole (Microtus ochrogaster), induced size ovulation, 252 human, 196, 279 predation risk, relationship to intromission duration, relationship to testes size, 196, 197 75–76, 268 Pseudocheiridae, 21 Primates, 39 pseudo-vagina (birth canal), marsupials, 209 baculum, 135–140, 135 Pteropodidae, 32, 32, 70 size, relationship to copulatory pattern, 137 pygmy hippopotamus (Choeropsis liberiensis), classification schemes, 37–39 copulatory pattern, 63 copulatory courtship pygmy mice (Baiomys taylori), copulatory pattern, neck grips, 215 68 tactile stimulation, 217 pygmy possums, 20 copulatory locks, 70 pygmy rabbits (Brachylagus idahoensis), copulatory patterns, 66–67, 101–102 copulatory pattern, 65 interspecies variability, 102 pygmy spotted skunk (Spillogale pygmaea), pelvic intraspecies variability, 101–102 thrusting, 106 copulatory plugs, 198–199 copulatory postures, 87–88,87–88 rabbit (Oryctolagus cuniculus), 35–36 defining traits, 39–40 copulatory patterns, 62–64,62–64 ejaculation intervals, 169–170, 170 induced ovulation, 65 female behavioural responses, 147 somatosensory feedback, 257, 258 female orgasmic responses, 213 nerve growth factor, 261–262 Hominoidea, 40–41 seminal pH, 231 levator penis muscle, 153 sperm storage site, 193 mating systems, 96 rapid mating (pattern no. 16), 62–65

Index 385

rapid sperm transport, 229–231 ruffed lemur (Varecia variegata), 38–39 rat kangaroos, mating systems, 94 RXFP2 gene, 185, 185 rat (Rattus norvegicus) copulatory pattern, 68 scaly-tailed squirrels, 35 number of intromissions, 73 Scandentia, 37 copulatory plugs, 198 Sciuridae, 35 ‘cups’, 151 copulatory patterns, 77 lordotic response, 147, 218 intromission duration, 77 ovulation control, 256–257 mating systems, 97 penile muscles, 150 multiple-partner matings, 97 sperm morphology, 172 vaginal morphology, 221 red deer (Cervus elaphus), seminal pH, 231 scrotum, 184, 272–273 red kangaroo (Macropus rufus) evolutionary origins and functions, 184 copulatory posture, 84 hair loss, 184 mating system, 95 phylogenetic distribution, 184–185 red panda (Ailurus fulgens), 106 Scuiromorpha, 35 red uakari (Cacajao calvus rubicundus), 148 sea otter (Enhydra lutris), 82 copulatory courtship, 108 seacows, 42–44 red-necked wallaby (Macropus rufogriseus), 17 seals red-sided garter snake (Thamnophis sirtalis) aquatic vs. terrestrial copulations, 82–83 ovulation control, 251–252 copulatory pattern, 69 penile spine ablation, 149 fur seal (Arctocephalus forsteri), 106 reptiles, ovulation control, 251–252 copulatory courtship, 216 rhesus macaque (Macaca mulatta) seasonal breeding, tammar wallaby, 18 cervical sperm migration, 232, 233 secondary structures, involvement in gamete levator penis muscle, 153 transfer, 115, 115 mating system, 91–92 selective abortion, 245 rhinoceros, 26, 28 semelparity, 23 chinning behaviour, 214–216 Antechinus sp., 94, 122 mating systems, 96 semen ovulation control, 251 buffering action, 200, 200, 229, 274 pelvic thrusting pattern, 106 collection methods, 169 population decline, 280 immunosuppressant qualities, 228 seminal pH, 231 pH, 200–201, 231, 274 ringtail possums, 21 semenogelin 2 gene evolution, 202–203, 202–203 ring-tailed lemur (Lemur catta), 148 seminal coagulation, 187, 196–197, 274 copulatory plugs, 198 and semenogelin 2 gene evolution, 202–203. See duration of female receptivity, 146 also copulatory plugs mating system, 198 functions, 201–203 rock hyrax (Procavia capensis), 41 human, 279 Rodentia, 34, 34 importance of, 198 baculum, 135 marsupials, 197–198 copulatory courtship, 217–218 placental animals, 198 copulatory locks, 81, 125–127 primates, 198–199 copulatory patterns, 102–105 relationship to mating systems, 199 copulatory plugs, 198 role of vesicular glands, 196 cryptic female choice, 103 seminal vesicle secretory (SVS) proteins, 203, 203 induced ovulation, 249–250 seminal vesicles. See vesicular glands mating systems, 35, 97 seminiferous epithelium cycle (SEC) lengths, 163, 163 multiple intromission, 73 human, 279 penile morphology, 125–127 relationship to sperm length, 174 sperm competition strategies, 103–104 seminiferous tubules, 163 sperm recovery studies, 104 diameter, relationship to sperm length, 174–175 spiny species, copulatory patterns and sengis (elephant shrews), 45–46 intromission duration, 79–80 sensory traps suborders, 35 hoofed animals, 214–216 rosettes, echidna, 130 neck grips, 214, 215

386 Index

sex peptide, fruit ﬂies, 226–227 hazards encountered, 205–207, 206 sexual dimorphism immune responses to, 228–229 human, 277 maturation in epididymis, 165 relationship to copulatory plugs, 198 non-ejaculated, recovery from vas deferens, sexual skin swellings, 40, 219–221, 220 192–193 sheep sperm binding, oviducts, 237 cervical morphology, 225–226 sperm bundles, 179, 179–180, 272 sperm counts, effect of frequent ejaculation, 169 relationship to acrosome reaction and short-beaked echidna (Tachyglossus aculeatus), 93–95 hyperactivation, 239 pelvic thrusting pattern, 106 sperm competition, 3–4, 155 shrew opossums (Caenolestidae), 14 and cryptic female choice, 89–90,89 shrews and relative testes size, 156, 271 neck grips, 215 bats, 156 sperm storage, 236–237, 238 cetaceans, 161–162 Simpson, George Gaylord, 46 primates, 156–159 single intromission, phylogenetic distribution, 73 terrestrial carnivores, 159–161 Sirenia, 42–44 and semenogelin 2 gene evolution, 202–203 copulatory patterns, 303 and vas deferens morphology, 191, 192 fossil evidence, 43 associated traits, 278–279 lumbar rigidity, 72 bulbocavernosus muscle size, 149–151 skunks insect research, 4 Mephitis mephitis kangaroos, 95 copulatory courtship, 216 sperm competition strategies, 99 intromission duration, 82 bundle formation, 180 Spillogale pygmaea, pelvic thrusting pattern, 106 frequent copulation, 100 slender loris (Loris tardigradus) multiple intromission copulatory courtship, 217 primates, 100–102 copulatory posture, 86–87, 86 rodents, 102–105 sloths, 47,49 primates, 100–102 fossil records, 50 prolonged intromission, 100 snow-shoe hare (Lepus americanus), 36 rodents, 102–105 social organization seminal coagulation, 201–203 and ovulation control, 252–253 sperm counts mole-rats, 254 effect of frequent ejaculation bats, 98 chimpanzee, 168 somatosensory feedback, 256–257 human, 168, 168, 279 role in induced ovulation, 258, 263 sheep, 169 multiple cues, 257 inﬂuencing factors, 169 multiple intromission, 257–259 relationship to mating systems, 205 prolonged intromission, 259–260 relationship to sperm length, 174, 271 Soricidae, 33, 70, 74 semen collection methods, 169 copulatory patterns, 71 sperm energetics, 177, 178 South American marsupials, 12 sperm hyperactivation, 241 Didelphimorphia, 12–14, 13 relationship to capacitation, 239 Microbiotheria, 14 sperm length, 164, 173–174, 271 Paucituberculata, 14 relationship to body size, 174 Spalacidae relationship to seminiferous tubule diameter, copulatory pattern, 77 174–175 intromission duration, 77 relationship to sperm numbers, 174 species, number of, 6–7, 6–7 relationship to swimming speed, 175, 271–272 sperm sperm midpiece cooperative behaviour, 178–179 energy production, 176 bundle formation, 179–180, 179, 272 JC-1 studies, 177, 178 pairing, 180, 272 size, 271–272 rouleaux formation, 182 relationship to swimming speed, 177 train formation, 180–181, 272 relationship to testes size, 176, 176–177 epididymal transit times, 166–167, 167 sperm mitochondria, 172

Index 387

sperm morphology, 173 spiny bandicoots, 23 apical hooks, 180–182, 181, 272 spiny species diversity, 171–172, 172 copulatory patterns, 79–80,80–81 heads, 172 courtship, 79–80 human, 279 penile morphology, 81 midpiece, 172 spontaneous ovulation, 248 sperm pairing, 180, 272 and social organization, 253 sperm production, 163–164 evolutionary origins, 248–251, 255, 263–264 augmentation methods, 155, 271 neuroendocrine mechanisms, 256 effect of high temperature, 182 overlap with induced ovulation, 256–257, 263 human, 279 phylogenetic distribution, 251 placental animals, 165 non-mammalian vertebrates, 251–252 sperm quality, 155 uncertainties, 249–251 sperm storage, 231 spotted hyena (Crocuta crocuta), 29–30,29 epididymis, 167 copulatory lock, 129–130 female reproductive tract copulatory pattern, 61–62 alpaca, 235–236 female genitalia, 128–129, 129 bats, 156 mating mechanics, 129 bats, 98 pelvic thrusting pattern, 106 cervical crypts, 232–233 penile morphology, 129 domestic dog, 232–234 squirrel monkeys (Saimiri sciureus), copulatory long-term, 246, 246–247 pattern, 66, 102 oviducts, 235–237, 238 stickleback, sperm morphology studies, 177 uterotubal junction, 235–236 striped possum (Dactylopsila sp.), 19,20 sites of, 193 striped skunk (Mephitis mephitis), 82 vas deferens, 193 stump-tail macaque (Macaca arctoides) sperm survival, 155 copulatory lock, 71, 126 effect of female immune defences, 228 copulatory pattern, 102 pH challenges, 229 female orgasmic responses, 213 role of seminal coagulation, 201 effects of local anaesthesia, 213 sperm swimming speed, 271–272 penile morphology, 118 relationship to length, 175 subspecies, 6–7 relationship to midpiece length, 177 subterranean mammals, copulatory patterns, 76–79 sperm train formation, 180–181, 272 sugar gliders (Petaurus breviceps), 20–21 sperm transport Sumatran rhinoceros (Dicerorhinus sumatrensis), ejaculation, 169–170 ovulation control, 251 in female reproductive tract Sunda pangolin (Manis javanica), 86 alpaca, 235–236, 236 superfoetation, hares, 247 cervical grooves, 232 swamp wallaby (Wallabia bicolor), 17 chemotaxis, 241–243 domestic dog, 232–234, 234 talapoin (Miopithecus talapoin), sexual skin effect of cervical mucus, 231–233 swellings, 220 hyperactivation, 241 tammar wallaby (Macropus eugenii), 18 marsupial vaginae, 209 female reproductive system, 208, 209–210 oviducts, 235, 238–239 mating system, 94–95 rapid phase, 229–231 sperm storage, 232 thermotaxis, 241 tapir, 26, 28 uterotubal junction, 234–235 copulatory pattern, 289 recovery of non-ejaculated sperm, 192–193 penile morphology, 112 role of seminal coagulation, 201 seminal pH, 231 vas deferens smooth muscle function, 192–193 tarsiers sperm whale (Physeter macrocephalus) Tarsius spectrum, 38–39 copulatory posture, 87–88 Tarsipedidae, 20 testes size, 161, 162 Tasmanian devil (Sarcophilus harrisii), 22,22–23 spinal reﬂexes intromission duration, 75 domestic dog, 61 Tayassuidae, 98 felids, 64 copulatory pattern, 72, 285

388 Index

Tenrecidae, 41,45 uterine contractions, role in sperm transport, baculum, lesser hedgehog tenrec, 134 229–230, 234 copulatory patterns, 80 uterotubal junction ovulation control, 251 sperm storage function, 235–236 testes size, 156 bats, 246 and semenogelin 2 gene evolution, structure, 234–235 202–204 valve function, 235 Antechinus sp., 122 uterus bats, 97, 156–157 protective function, 228 cetaceans, 119, 161–162, 164 sperm storage, 246 feathertail glider, 94 honey possum, 94 vagina, marsupials, 207–209 human, 279 sperm transport, 209 intraspecies variation, 156 tammar wallaby, 209 nocturnal primates, 96 vaginal folds, 221–223, 222 northern right whale, 98 vaginal morphology, 221 primates, 156–159, 158 cetaceans, 221–223, 222 relationship to accessory reproductive gland dugong, 221 weights, 196, 197 primates, 221 relationship to mating systems, 271 vaginal pH, 200, 229 relationship to oviduct length, buffering action of seminal fluid, 229, 274 239–240, 241 human, comparison with other mammals, 230 relationship to sperm midpiece volume, 176, vagino-cavernosus reflex, 212 176–177 human females, 212–213 terrestrial carnivores, 159–161, 160 vagino-levator reflex, 213 testicular descent, 183–184, 272–273 vas deferens evolutionary origins and functions, 184 ampulla, 189–192 phylogenetic distribution, 184–185 epithelial structure, 189 RXFP2 and INSL3 genes, 185 length, 191, 192 testicular organization, 163 recovery of non-ejaculated sperm, 192–193 testicular positioning, 183 smooth muscle function, 192–193 testosterone-dependent development, penile smooth muscle thickness, 189, 191, 192 morphology, 136 human, 191, 279 therapsids, 3 sperm storage, 193 Theria, 7–8 ventro-ventral copulatory postures, 87–88 thermotaxis, 241 cetaceans and sirenians, 87–88 Thylacosmilus,25 primates, 88 Tidarren fordum, sperm transfer, 115–116 vesicular glands, 273–274 tiger quoll (Dasyurus maculatus), 22 drill, 194 Titanotaria orangensis, 141 importance of, 198 traditional Chinese medicine, 281 phylogenetic distribution, 187, 187–188 Tragulidae, 74 secretions, 194, 196–197 copulatory patterns, 63, 72, 286 pH, 200 treeshrews, 37 SVS proteins, 203, 203 mating system, 96 size Trivers, Robert, 178 human, 196, 279 tuatara (Sphenodon punctatus), relationship to mating systems, 194–196, 195 111–112 relationship to testes size, 196, 197 Tubilidentata, 44–45 structure, 194 Tupaiidae, 37, 74 Vespertilionidae, 32, 70, 74 twinning, 244 copulatory locks, 127 sperm storage, 98, 246–247 urethral grooves, 122 testes size, 98 urethral processes, 130, 152, 223, 270 Virginia opossum (Didelphis virginiana), 12 urethral protection, role of baculum, 140 male reproductive tract, 122 Ursidae, 74 sperm morphology, 172 testes size, 160 sperm pairing, 180, 272

Index 389

visual communication, 217 sperm whale (Physeter macrocephalus) Viverridae, 29, 74 copulatory posture, 87–88 testes size, 160 testes size, 155–162, 162, 164. See also cetaceans vocalization white cell counts, relationship to mating system, blind mole-rat, 78 205–226 masked palm civet, 68 white-nosed coati (Nasua narica), pelvic thrusting Vombatidae, 21 pattern, 106 copulatory patterns, 283 woodmouse (Apodemus sylvaticus), 34 sperm train formation, 180, 272 wallabies, 16,16 woolly mouse opossum (Micoureus cinereus), glans female reproductive system, 208, 209–210 penis, 122–123 mating system, 94–95 sperm storage, 232 Xenarthra, 46, 47 walrus (Odobenus rosmarus), baculum, 138 armadillos (Dasypodidae), 46–48 Walton, Arthur, 111 fossil evidence, 49 warthog (Phacochoerus africanus), chinning lumbar rigidity, 72 behaviour, 216 Pilosa, 48–49 water shrew (Neomys fodiens), copulatory lock, 71 Weddell seals (Leptonychotes weddelli), 82–83 yak (Bos grunniens), urethral process, 130–131 western grey kangaroo (Macropus fulginosus), Yangtze River dolphin (Lipotes vexillifer), 280 penile muscles, 153 yapok (Chironectes minimus), 12–13 wet markets, 281 male reproductive tract, 189–190 whale lice, 161 yellow spotted hyrax (Heterohyrax brucei), urethral whales process, 130 humpback (Megaptera novaeangliae) mating system, 161 zoo research, 59–60 sperm length, 174 zoonotic infections, 280