Sexual Conflict and Sperm Competition Sperm Competition • Sperm compete to fertilize an egg • Occurs in animals with external as well as internal fertilization • Probably the first type of sexual competition to evolve due to anisogamy • Can be present in relatively simple and sessile animals Sperm of territorial and sneaker males differs in quality in male bluegill sunfish (Lepomis macrochirus). Sneaker male sperm is higher quality and fertilizes more eggs Sperm Competition • With internal fertilization, sexual selection can continue after copulation if females mate multiply • Sperm from rival males compete for fertilizations… • … OR cryptic female choice Sperm Competition • Males clearly benefit from extra copulations • Females can get all the sperm needed to fertilize their eggs from a single copulation • So, why do females copulate with multiple males? Number of partners Female Multiple Mating Hypotheses 1. Costs of resistance exceed the costs of acquiescence – No reproductive benefit, but may be harassed or harmed if she resists Female Multiple Mating Hypotheses 2. Direct (material) benefits – Ensure all eggs are fertilized – Increase amount of parental care (males may co-operate in care if they have a chance of paternity) – Reduce male harassment of herself and her young by giving all males a chance of paternity (e.g. primates) Concealed ovulation coupled with promiscuous mating may help to prevent infanticide in primates Female Multiple Mating Hypotheses 3. Indirect (genetic benefits) – Increase the genetic quality of her offspring – E.g. Socially monogamous birds frequently engage in extra-pair matings • Colourful plumage and displays are viability indicators that function to attract other males’ females for extrapair matings • Females paired to attractive males are more faithful – Different sires are better for sons versus daughters Sexual Conflict • Occurs when the optimal outcome is different for males and females • Male and female adaptations and counteradaptations Enforced copulation in Cimex lectularius Waterfowl • Length of male phallus is correlated with frequency of forced extra-pair copulations – Length varies from 1.5 to 40 cm • In species where phallus is long and elaborate, females have more elaborate vaginal morphology – Reduce chances of male intromission without co-operation Vagina Mallard duck (Anas platyrhynchos) 2 cm Phallus In species where males have a longer phallus, females have a more elaborate vagina Forced Copulations – Water Striders • Male pounces on female and tries to secure a mating by grasping her • Across species there is correlated evolution of male morphology to increase grasping (elongation of grasping genitalia) and female morphology to resist (elongation of abdominal spines to thwart the male) MALE Gerris incognitus FEMALE MALE Gerris thoracicus FEMALE Male grasping genital segment and female abdominal spine correlate across species Hemipteran beetle – Xylocoris maculipennis • Male simply pierces the body wall of the female to inject sperm – Swim around until they meet an egg • Does this to other males too – Sperm make their way to victim’s testes where they can be passed on to females Mate Guarding by Males • Ensure paternity by guarding mates or by copulating frequently to swamp the sperm of rivals • E.g. Socially monogamous birds – Males follow females closely during the time she is fertile – Reduce efforts in feeding the chicks if female is absent during this time Bottlenose Dolphins • 2 – 3 males form alliances to herd females and defend them from other males • Males sometimes use force to ensure that females stay with them Arthropods • Prolonged copulation in arthropods is a form of mate guarding Male Adaptations • Post-copulatory adaptations of males include: – Sperm removal • Remove sperm of rival males before depositing own – Sperm displacement • Insemination flushes out previous inseminations Uninflated penis Inflated penis Flagellum caries barbs to remove sperm Sperm removal in Odonata: Penis of Crocethemis erythraea (top) and Orthetrum cancellatum (bottom) Male Adaptations • Post-copulatory adaptations of males include: – Copulatory plugs • Male “cements” up female’s genital opening to prevent other males from mating with her • E.g. Moniliformis dubius (parasitic acanthocephalan worm in rats) seals up both females and other males – Anti-aphrodisiacs • Males deposit a scent after mating that discourages other males Male Adaptations • Post-copulatory adaptations of males include: – Sterile sperm • Males produce two types of sperm – eusperm (fertile) and parasperm (sterile) • Parasperm is a “cheap filler” of female sperm storage structures and also helps protect eusperm from spermicide Male Adaptations • Post-copulatory adaptations of males include: – Accessory gland proteins (Acps) • Ejaculate contains a cocktail of proteins that influence female behaviour and physiology • Functions include incapacitating rival sperm, protecting sperm, increasing egg laying rate in female, and decreasing female’s propensity to re-mate • Reduce female longevity Acps decrease longevity in female Drosophila melanogaster (dark blue line) relative to controls Male Adaptations • Post-copulatory adaptations of males include: – Strategic allocation of sperm • Male fowl increase their sperm investment when more male competitors are around, and when females have larger combs (produce larger eggs) • Males may use less sperm or reject females with whom they have already mated Gallus gallus Female Adaptations • Extra-pair copulations (EPC) – Females of many seemingly monogamous or polygynous species engage in infidelity – Common in birds Frequency (+SE) of extrapair young in relation to the social mating system (M = monogamy, P = polygyny) of 40 species of passerine birds Female Adaptations • Cryptic female choice – Females may have some control over which male’s sperm fertilizes her eggs • E.g. Feral fowl (Gallus gallus): – Subordinate males may force copulations – Females contract cloaca to eject sperm Field Crickets (Gryllus bimaculatus) • Mated females with two males: – Both siblings – One sibling and one non-sibling – Both non-siblings • Hatching success decreased only if both males were siblings • DNA markers showed that females preferentially stored sperm of unrelated males Sexual Cannibalism • Females eat the males they copulate with • Common in mantids and spiders • Males may or may not deliberately sacrifice themselves Redback Spiders • Males offer themselves as a meal after copulation • Males that get eaten fertilise more eggs than uneaten males, and thus benefit from sexual suicide • Cost to male is low – Little chance of mating again
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