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Sexual selection Opposing forces Conflict forms basis for Darwin’s theory of natural selection How then does cooperation evolve? Group selection arguments are weak Conflict even found within “cooperative relationships” such as mating pairs Levels of cooperation Attractive forces must outweigh repulsive and centrifugal forces for cooperation to occur Fitness Attractive and repulsive forces are measured by fitness Inclusive fitness of an allele is its effect on the individual that bears it (direct fitness) and the fitness of other individuals that carry it (indirect fitness) Remember kin selection and Hamilton’s rule? More on that calculating that later Frequency dependent selection Important in conspecific interactions Rare variant is “released” from competition Evolutionarily stable strategies (ESS) A strategy such that, if all the members of a phenotype adopt it, then no mutant strategy could invade under the influence of natural selection Used extensively in behavioral ecology Hawk versus dove strategy Hawk – More likely to win the prize Dove – More likely to escape injury Is Dove an ESS? Is Hawk an ESS? Yes, if benefit is greater than cost Fitness (V) is greater than cost of injury (C) No, if benefit is less than cost Optimal strategy is variable Mixed strategy results based on amount of fitness payoff Hawk versus dove Individual will adopt hawk behavior with a probability of V/C Sexual selection Differences among individuals of a sex in the number of mates they obtain Intersexual selection – driven by interaction between the sexes Intrasexual selection – driven by interaction within the sexes Variation in mating success is generally greater for males than females Intrasexual competition Sperm or pollen competition Multiple paternity in plants, squirrels, bears, birds, lizards, spiders, human twins Large ejaculates increase probability of fertilizing the egg Medflies ejaculate 2.5x more sperm when mate in presence of other males Testes size in primates Sperm scoopers in damselflies Intrasexual competition Combat or other direct competition Selection for armaments, large body size, tactics Combat in staghorn beetle Color signals and combat in blackbirds Intrasexual competition Infanticide Male lions that take over prides of females Female jacanas that destroy nests incubated by males Leads to faster mating opportunities Summary of intrasexual competition Primarily male-male competition over mates Females may compete over mates if access to males is limiting Three main types of intrasexual selection Combat Sperm competition Infanticide Intersexual selection Female choice Human biases in the study of female choice Examples of female choice Theories about why females are choosy Bias in the study of choice Darwin ridiculed for assertion that females have a ‘sense of beauty’ Females thought to choose correct sex and species only Courtship to overcome female ‘reluctance’ or fear of predation Female choice… Details later…much evidence of choice across a variety of taxa With DNA markers, evidence of multiple mating partners in ‘socially monogamous’ species Female choice example: Tungara frogs Males advertise with calls to females Two calls, simple (tun) and complex (tun gara gara) Females prefer complex calls Cost of sexy calls Bats locate males making complex calls more easily Female choice example: barn swallows Barn swallows and tail length The famous peacock’s tail Why be choosy? Good genes: advertisement a true indicator of quality Barn swallows and mites Why be choosy? Good genes: advertisement a true indicator of quality Eyespot area and offspring weight in peacocks Why be choosy? Good genes: advertisement a true indicator of quality Gray tree frogs Why be choosy? Why be choosy? Resource acquisition: nuptial gifts, or advertisement indicates future paternal care Hangingfly nuptial gifts Sensory bias Why be choosy? Sensory bias: females have intrinsic preferences; advertisements are arbitrary Crests in grassfinches Supernormal stimuli Runaway selection for sensory bias Runaway selection for sensory bias Sexy son hypothesis Genetic correlations between choice and trait due to assortative mating Offspring of female will be ‘sexy’, have same advertisement favored by females Original trait is arbitrary Summary of intersexual selection Advertisements may or may not be informative Single display could have multiple meanings and benefits Male parental care Kittiwakes, monogamous birds, male care, similar variation in offspring number When male care exceeds female care Pipefish: female success limited by space in brood pouch of male Males choose females based on body size, skin markings When male care exceeds female care Red phalaropes, showy females court males, which provide all care for eggs and young When male care exceeds female care “Sex-reversed” birds often have precocious young, polyandry, female-biased sex ratios Nest predation is high, only one parent stays at nest to decrease visibility Females selected for multiple mating (replacement clutches) which apparently decreased parental care Why don’t all species have male care? Most species have NEITHER male nor female care Lactating mammals are an extreme Male care more common when paternity more certain For example, internal fertilization Generally, males aren’t very choosy (especially when polygynous) In some frogs & water striders, males have evolved special ‘release signal’ given to other males who have clasped them in error Wild turkeys will court not only stuffed females but models of female heads suspended 40 cm above the ground Sockeye salmon will spawn with any red object Male mating mistakes