The Genetics and Biology of Sexual Conflict

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Index

A C Acp36DE, 55 Caenorhabditis elegans, 186–187, 191, 292 Aedes aegypti, 56–57, 166, 168–169, 317 Caenorhabditis remanei, 103 Aging Callosobruchus, 87–88, 94, 107, 120–122, 396 evolution of longevity and aging Callosobruchus maculata, 362, 386 sex-specific selection, 104–105 Cannibalism, sexual potential for sexual conflict over longevity and female counterstrategies in monogynous or aging, 105 bigynous mating systems, 376–379 evidence, 105–106 male mating strategies in monogynous or bigynous susceptible systems, 107 mating systems, 373–375 prospects for study, 109–111 overview, 368–370 interlocus sexual conflict effects paternal investment, 379 environmentally mediated changes in degree prospects for study, 379–380 of sexual conflict, 103 responses to threat, 371–373 evolutionary consequences, 103 Cardinium, 312–313, 316 proximate effects, 101–103 Ceratitis capitata,57 sexual dimorphism, 106 Cetorhinus maximus, 388 variation effects on sexual conflict, 107–109 Cheilomenes sexmaculata, 320 Akodon azarae, 167 Chromodoris reticulata, 279 Anopheles gambiae, 185–186, 191 CI. See Cytoplasmic incompatibility Antichiropus variabilis, 388 CMS. See Cytoplasmic male sterility Antimicrobial peptides. See Seminal fluid Coevolution. See Genital coevolution; Seminal fluid proteins proteins; Sexual antagonism; Sexually Apis mellifera, 292–293 antagonistic traits Apoanagyrus diversicornis, 318 Copulatory plug, 333–334 Aquarius,93 Copulatory wounding (CW). See also Traumatic Argiopelobata, 375, 377–379 insemination Armidillidium vulgare, 311–312, 320 bleeding prevention, 124 Arsenophonus nasoniae, 314 case studies Arygoneta aquatica, 370 Drosophila, 118–119 Asobara japonica, 318 Siphopteron, 118, 120 Asobara tabida, 312 bed bug, 120–121 Callosobruchus, 120–122 evolutionary consequences B costs of mating, 126–129 Bathyergus suillus, 389 reproductive isolation Bean weevil. See Callosobruchus barrier against hybridization between species, Bed bug 131 copulatory wounding, 120–121 major evolutionary trends, 131–132 genital coevolution, 387 wounding as an isolating factor, 131 Birds. See Extrapair paternity; Genital coevolution; sexual conflict and selection Parental care, sexual conflict; Sex-biased accidental sperm and seminal fluids, 126 gene expression sexually transmitted diseases in wound Blarina brevicauda, 389 infliction, 126 Bombyx mori, 182, 186, 19 traumatic penetration from accidental Brevipalpus phoenicis, 312 copulatory wounding, 125–126 Bryobia, 313 variations in incidence, 129–131

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Copulatory wounding (CW) (Continued) EPY. See Extra-pair young hermaphrodites, 123 ESD. See Environmental sex determination humans ESS. See Evolutionarily stable strategy epidemiology of genital wounds, 132 Eurema hecabe, 312 male injuries, 132 Evolutionarily stable strategy (ESS), 3, 9, 15 physiology, 132 Extrapair paternity (EPP), birds rape and nonconsensual sex, 132–133 benefits, 146–147 immune response, 124 conflicts of interest, 141–144 infection prevention, 124 costs, 147 life-history consequences, 124–125 episodes of conflict, 147–148 males, 123 history of study, 141–143 mechanical origins mating pattern variation as sexual conflict driver, extragenital wounding, 122–123 148–150 intragenital wounding, 122 prospects for study, 150–151 overview, 115–116 Extra-pair young (EPY), 250 phylogenetic distribution of traumatic mating, 117 physiological costs, 123–124 prospects for study, 133–135 F self-infliction, 123 Feeding, seminal fluid protein studies, 56–57 terminology, 116 Female resistance. See Sexually antagonistic traits types, 116 Female-targeted sperm release, evolution, 14–18 Cordylochernes scorpoides, 313 Feminization, reproductive parasitism, 311–312 Coridromius bulbopella, 87, 93 FEPC. See Forced extrapair copulation Cornu aspersum, 281 Fiddler crab, genital coevolution, 386 Cryptodiopsis dalmani, 173 Forced extrapair copulation (FEPC), 395 Culex pipiens, 166, 168–169 Franklinothrips vespiformis, 313 CW. See Copulatory wounding Cytoplasmic incompatibility (CI), 316–317, 321–322 Cytoplasmic male sterility (CMS), 314–315, 319 G Gallus gallus, 108, 183–184 Gambusia, 388, 392 D Gamete competition theory, 8 Danio rerio, 182–183 Gamete limitation theory, 8 Daphnia pulex, 187 Gamete size, dimorphism evolution, 7–10 Darwin–Bateman paradigm (DBP), 2, 13, 19 Gammarus duebeni, 312, 318, 320 DBP. See Darwin–Bateman paradigm Genetic sex determination (GSD) Dermestes maculatus, 108 evolution patterns in sex-determination cascades Deroceras, 281 neutral evolution, 293–295 Dicrostonyx torquatus, 167 overview, 292–293 Diptera, 173 genetic and developmental architecture, 302–304 Drosophila. See Copulatory wounding; Genital selection sources on novel sex-determining genes coevolution; Reproductive parasitism; direct selection Seminal fluid proteins; Sex-biased gene pleiotropic benefits, 296–297 expression; Sex chromosome drive; Sex sex-ratio selection, 295–296 chromosomes; Speciation indirect selection Dytiscus latissmus, 87, 94 deleterious mutations on heterogametic sex chromosome, 301–302 intralocus sexual conflict, 298–301 E Genital coevolution Echinotheridion, 374 examples, evidence, and mechanisms Egg production, seminal fluid protein studies, 55–56 birds, 400 Elymus virginicus, 315 fish, 399 Environmental sex determination (ESD), transition to insects, 398 genetic sex determination, 295–297 mammals, 401 Ephestia cautella, 320 snakes, 399–400 EPP. See Extrapair paternity spiders, 398–399

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table, 386–389 fraternal birth order effect, 234 female mate choice role, 393–394 genetic model limitations, 235–237 mechanism distinction, 396–398 heritability estimates, 234–235 natural selection pedigree asymmetries, 233–234 lock-and-key mechanism to prevent population genetic models hybridization, 392–393 candidate mechanisms for balancing selection mechanical feasibility of copulation, 391–392 in males, 237 overview, 385, 390–391 female candidate selection mechanisms, prospects for study, 401–402 239–240 sexually antagonistic coevolution, 394–396 male genetic models, 237–239 Genomic conflict. See Sexually antagonistic zygotic drive prevalence and features, 231–232 Gerris, 87, 92–93, 387, 395 prospects for study, 240–241 Gerris gillettei, 362 stability over time, 231 Glossina pallidipes, 387, 394 twin studies, 234–235 Gonad expenditure, 18 Horse Gonado-somatic index (GSI), 10–13 genital coevolution, 389 Graphoderus zonatus verrucifer,94 infanticide, 205, 212 Gryllodes sigillatus,90 HP1, 169 Gryllodes sigillatus, 106 Hypolimnas bolina, 309, 318, 320 GSD. See Genetic sex determination GSI. See Gonado-somatic index I Igf2r, parental conflict, 39–40 H Infanticide Hemachromatosis protein (HFE), 39 female counterstrategies Hermaphrodites individual counterstrategies, 211–213 copulatory wounding, 123 sexual and reproductive counterstrategies, fitness routes, 268 208–211 gonochorist sexual conflict comparison, 268–270 social counterstrategies, 213–218 interlocus sexual conflict male strategies overview, 267–269 adaptive and nonadaptive hypotheses, sequential hermaphrodites, 271–273 202–208 intralocus sexual conflict sexual selection hypothesis, 200–202 sequential hermaphrodites, 270–271 overview, 199–200 simultaneous hermaphrodites, 273–274 prospects for study, 218–219 overview of sexual conflict, 266–268 Interlocus sexual conflict postmating conflicts definition, 267 overview, 279 hermaphrodites sperm donor interests, 279–281 overview, 267–269 sperm recipient interests, 281–282 sequential hermaphrodites, 271–273 premating conflicts longevity effects overview, 274–275 environmentally mediated changes in degree resolution routes, 277–279 of sexual conflict, 103 sex-role preferences, 275–277 evidence, 105–106 reproductive parasitism in plants, 314–316 evolutionary consequences, 103 sequential hermaphrodite features, 270 potential for sexual conflict over longevity simultaneous hermaphrodite features, 273 and aging, 105 HFE. See Hemachromatosis protein prospects for study, 109–111 HIRA, 161 proximate effects, 101–103 Homonamagna, 317 sex-specific selection, 104–105 Homosexuality, human susceptible systems, 107 evolutionary investigations, 229–231 speciation role, 357–361 family clustering, 232 Intralocus sexual conflict fecundity definition, 267 females, 232–233 genetic sex determination evolution, 298–301 males, 232 hermaphrodites

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Intralocus sexual conflict (Continued) Micaria sociabilis, 370 sequential hermaphrodites, 270–271 Microbiome–Mother’s Curse paradox. See Mother’s simultaneous hermaphrodites, 273–274 Curse sex chromosome change, 343–344 MicroRNA, sex-biased gene expression role, 189, 192 speciation role, 356–358 Microspora, 312 Microtus oregoni, 173 Mitochondria. See Mother’s Curse K Mollusks, genital coevolution, 386 Kentish plover. See Parental care, sexual conflict Mother’s Curse (MC) competitive coexistence of symbionts and pathogens, 78–79 L fertility fitness, 75–76 Lacerta vivipara, 108 inbreeding Lactation/gestation ratio, 200–201, 206 fertility fitness, 76 Lag load, 26 viability fitness, 76 Lamprologus callipterus,33 kin selection, 77–78 Latrodectus, 374–375 microbiome paradox, 74–75, 78–80 LD. See Linkage disequilibrium nuclear compensatory mutations, 77 Leptopilina clavipes, 318 overview, 73–75 Life span. See also Aging prospects for study, 80 interlocus sexual conflict effects symbiont transmission mode evolution, 79 environmentally mediated changes in degree viability fitness, 75 of sexual conflict, 103 MSCI. See Meiotic sex chromosome inactivation evidence, 105–106 Muroidea, 167 evolutionary consequences, 103 Mus minutoides, 167 potential for sexual conflict over longevity and Musca domestica, 175, 295 aging, 105 Muscidifurax uniraptor, 313 prospects for study, 109–111 Myopus schisticolor, 167 proximate effects, 101–103 sex-specific selection, 104–105 susceptible systems, 107 N seminal fluid protein studies, 58–59, 102 Nasonia vitripennis, 314 sexual dimorphism, 106 Neochrysocharis formosa, 313 variation effects on sexual conflict, 109 Neococcoidea, 175 Linkage disequilibrium (LD), 42 Nephila fenestrata, 373, 375 Lion, infanticide, 210, 212, 214 Nephila pilipes, 373 Littorina saxatilis, 182, 187 Nephilengys malabarensis, 376 Longevity. See Aging; Life span Nerodia, 393 Lymnaea stagnalis, 280 Nesidiocoris tenuis, 332 Lysin, 62 Nicrophorus defodiens, 251, 255 Niophalaria glabrata, 275 Nosema granulosis, 312 M Notiomystis cincta, 334 Macrolophus pygmacus, 332 Macrostomum lignano, 280, 282 Major histocompatibility complex (MHC), male O competition, 32 Offspring desertion. See Parental care, Male adaptations to sperm competition. See Sperm sexual conflict competition Onthophagus taurus, 387, 392 Manduca sexta, 186 Operational sex ratio (OSR), 2–3 Mating conflict, definition, 267 OSR. See Operational sex ratio MC. See Mother’s Curse Ostrinia scapulalis, 314, 321 Meiotic drive. See Sex chromosome drive Ovd, 175 Meiotic sex chromosome inactivation (MSCI), Overdominance, homosexuality genetic model in 175, 347 males, 238 MHC. See Major histocompatibility complex Ovulin, 51–52

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P Reproductive parasitism Parafontaria tonominea, 388 cytoplasmic incompatibility, 316–317, Parasitism. See Reproductive parasitism 321–322 Parental antagonism host responses to sex-ratio distorters, overview, 40–41, 43 319–321 resolution, 44–45 male lethality and virulence, 317 Parental care, sexual conflict mating system/reproductive ecology diversity of care strategies, 247–249 evolution, 318 ecology of conflict and cooperation, 256–258 mechanisms of action, 322–323 evidence, 249–250 multiphenotype model, 323–324 fitness consequences, 250 obligate asexuality evolution, 317–318 macroevolution considerations, 256 overview, 309–310 manipulation and parental tactics, 251–252 population biology overview, 245–247 sex ratio/sex allocation distortions prospects for study, 258–259 embryonic male killing, 312–314 sexual conflict detection over care feminization, 311–312 comparative analysis, 255–256 hermaphroditic plants, 314–316 experiments, 254–255 parthenogenesis induction, 312–313 field observations, 252–254 symbiont–host conflict, 310–311 Kentish plover, 253–254, 258 prospects for study, 322–324 Parental conflict, 39–40 Rheumatobates rileyi, 87, 92 Parental investment (PI), 2–3 Rhizoglyphus robini, 362 Parthenogenesis, induction in reproductive parasitism, Rickettsia, 312–314 312–313 Penile fracture, human copulatory wounding, 132 Phyllostomus hastatus,41 S Phyrnus marginemaculatus, 249 SA-ZD. See Sexually antagonistic zygotic drive Physa acuta, 275 Schistosoma mansoni, 187 PI. See Parental investment Scira coprophila, 175 Pig, genital coevolution, 389, 394 Seal, infanticide, 205 Pisaura mirabilis, 372 Selfish genetic elements. See Sex chromosomes Plodia interpunctella, 106 Seminal fluid proteins (SFPs) PMSC. See Postmeiotic sex chromatin chromosomal distribution of loci, 63 Pnigalio soemius, 313 coevolution with receptors in females, Poecilia reticulata, 388, 399 61–62 Postcopulatory sexual selection. historical perspective, complexity in Drosophila,50–52 3–4 expression, 50 Postmeiotic sex chromatin (PMSC), 174 functions, 50–51 Potential rates of reproduction (PRR), 3, 13 male adaptations to sperm competition, 333 Power, sexual conflict, 26–27 overview, 49–50 Prairie dog, infanticide, 206 postmating sexual conflict studies Precopulatory sexual selection, historical perspective, egg production, 55–56 2–3 food intake and processing, 56–57 Pre-ejaculatory sexual selection, evolution, 7, 13–14 immune activation, 57–58 Premating competition, 18 life span, 58–59 Primates overview, 52–53 genital coevolution, 389, 401 remating, 52, 54 infanticide, 202–204, 206–207, 209–210, 214–215, sperm transfer, storage, retention, and usage, 217 54–55 Primordial sexual conflict, definition, 266 practical applications, 65 PRR. See Potential rates of reproduction processing, 51–52 prospects for study, 64–66 rapid evolution, 60–61 R sexual conflict in evolution Rape. See Copulatory wounding conflict resolution evidence, 63–64 Remating, seminal fluid protein studies, 52, 54 molecular signatures, 59–63

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Sepsis cynipsea, 106 Sex determination, see Environmental sex Sex-biased gene expression determination; Genetic sex Anopheles gambiae, 185–186, 191 determination birds, 183–184, 189 Sex peptide (SP), 52, 54, 62, 102 Bombyx mori, 182, 186, 191 Sex peptide receptor (SPR), 54, 62 Danio rerio, 182–183 Sex ratio Daphnia pulex, 187 direct selection sources on novel sex-determining development studies, 191–194 genes, 295–296 Drosophila, 182, 184–185, 188–193 distortion in reproductive parasites. See evolution, 188–190 Reproductive parasitism gene identification, 190–191 operational sex ratio, 2–3 Littorina saxatilis, 182, 187 unity sex ratio evolution, 10 Manduca sexta, 186 Sexual antagonism nematodes, 186–187 homosexuality selection in males, 238 overview, 181–182 overview, 40 prospects for study, 194 resolution, 44–45 quantification, 182, 188 sexual conflict coevolution, 84–86, 91–95 rodents, 183 Sexual cannibalism. See Cannibalism, sexual Schistosoma mansoni, 187 Sexual cascade Solenopsis, 186 evolution of sexual strategies, 6–7 Triboleum castaneum, 186 female-targeted sperm release evolution, Xenopus, 184 14–18 Sex chromosome drive transitions Diptera, 166 events leading to ancestral isogamnous Drosophila eukaryote, 7 overview, 166, 168–169 gamete size dimorphism evolution, 7–10 polymorphism maintenance, 171–172 mobility/behavioral complexity, 11–15 population extinction, 169–171 overview, 7 suppressor evolution, 172–173 pre-ejaculatory sexual selection, sex roles, and evolutionary consequences sexual conflict, 13–14, 18–19 mating system evolution, 173 sperm competition, 10–11 sex determination, sex chromosome, and unity sex ratio evolution, 10 genome evolution, 173–174 Sexual conflict speciation, 174–175 cannibalism. See Cannibalism, sexual heterochromatin and small RNA role, 169 copulatory wounding. See Copulatory wounding mosquitoes, 166, 168–169 definitions, 24–26, 38, 266 overview, 165–166 genetic benefits, 28–32 prospects for study, 175–176 hermaphrodites. See Hermaphrodites rodents, 166–168 historical perspective, 4–6 X versus Y chromosome, 166 infanticide. See Infanticide Sex chromosomes longevity studies. See Aging; Life span conflict role in turnover and change parental care. See Parental care, sexual conflict overview, 342–343 population-level consequences, 33–34 intralocus sexual conflict and change, seminal fluid proteins. See Seminal fluid 343–344 proteins population size, linkage, and turnover, sex chromosomes. See Sex chromosomes 344–346 sexual antagonism coevolution, 84–86, dosage compensation studies, 346 91–95 selfish genetic element sexual cascade, 13–14 attraction, 349 sexual selection relationship, 26–34 sexually antagonistic allele interplay, speciation. See Speciation, sexual conflict role 349–350 sperm competition. See Sperm competition turnover role, 341–342, 346–347 Sexually antagonistic traits sexualization of X and Z chromosomes, coevolution 347–349 female resistance, 85–86 transmission, 341–342 mating economy, 86

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predictable versus stochastic models, 86 overview, 355–356 case studies reinforcement and sexual conflict, 357, 362 diving beetles, 94–95 Sperm competition seed beetles, 94 male adaptations to sperm competition traumatic insemination, 93–95 copulation extension or repeating, 331 water striders, 91–93 copulatory plugs, 333–334 conditional economics of conflict, 90–91 ejaculate variation females seminal fluid proteins, 333 genetic variation for resistance to harm, 90 sperm number, quality, and morphology, genital coevolution, 394–397 331–333 overview, 89 mate guarding, 330–331 resistance types and forms, 89–90 overview, 329–330 males, 86–89 strategic allocation of mating effort, 334 overview, 83–85 sexual cascade, 5, 10–11 prospects for study, 95 sexual conflict Sexually antagonistic zygotic drive (SA-ZD) costs and benefits evolution of zygotic driver, 157 females, 335 hybrid infertility role, 161 males, 334–335 logic of genetic conflict, 157 mitigation and benefits, 335–336 mechanisms, 158–159 Sperm release. See Female-targeted sperm release molecular pathway, 161 Spiroplasma ixodetis, 314 prior evidence lack, 159–160 Spiroplasma poulsoni, 314 prospects for study, 161–162 SPR. See Sex peptide receptor sex-linked segregation distortion mechanism, SR, 165, 169–173 160–161 Sry, 167 zygotic drive overview, 156 Symbiosis. See Mother’s Curse; Reproductive Sexual selection parasitism definitions, 24–26 evolution. See Sexual cascade historical perspective T postcopulatory sexual selection, 3–4 Teleogryllus commudus, 106 precopulatory sexual selection, 2–3 Telostylinus angusticollis, 102 sexual conflict relationship, 26–34 Tenrec caudatus, 389 SFPs. See Seminal fluid proteins Thamnophis sirtalis, 29–32, 388 Siphopteron, 118, 120, 281 TI. See Traumatic insemination Slx1, 168, 174–175, 293, 297, 350 Tidarren, 374 Sly, 168, 174–175, 350 Tmy, 174 Snakes, genital coevolution, 399–400 Traumatic insemination (TI). See also Copulatory Solenopsis, 186 wounding SP. See Sex peptide Drosophila, 118–119 Speciation, sexual conflict role selection from accidental sperm and seminal ecological co-adaptation, 357–358 fluids, 126 experimental studies sexually antagonistic trait, 93–95 evolution of reproductive isolation, Traumatic mating. See Copulatory wounding 361–362 Tribolium castaneum, 55, 186 female polymorphism, 363–353 Trichogramma, 313 fertilization traits and proteins, 362–363 species richness, 363 interlocus sexual conflict, 357–361 U intralocus sexual conflict, 356–358 Unity sex ratio, evolution, 10 modeling evolutionary dynamics of between-locus conflict, 358–361 V game-theoretic models, 358 VERL, 62–63 population genetic models of within-locus Vertically transmitted symbiont. conflict, 358 See Mother’s Curse

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W Y W chromosome. See Sex chromosomes Y chromosome. See Sex chromosome drive; Sex Wolbachia, 161, 311–314, 316–318, 320, 321–322 chromosomes Y-linked genes, 43 X X chromosome. See Sex chromosome drive; Sex Z chromosomes Z chromosome. See Sex chromosomes Xenopus, 184 Zyginidia pullula, 312 Xiphophorus maculatus, 295 Zygotic drive. See Sexually antagonistic X-linked genes, parental antagonism, 43–44 zygotic drive

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