Exam 1 Results: Mean: 75.7 Std. Dev: 10.3 Histogram 12 Frequency Distribution for Column 1 To (<) Count 43.500 49.111 1 49.111 54.722 0 54.722 60.333 2 60.333 65.944 4 65.944 71.556 7 71.556 77.167 10 77.167 82.778 11 82.778 88.389 7 88.389 94.000 4 Total 46 10 8 Count From (!) 6 4 2 0 40 50 60 70 80 Column 1 90 100 Recall: Sexual selection… • Viability: probability of survival to reproductive age • Mating Success Sexual selection: differential reproductive success due to differential mating success • Fecundity: viable offspring per female Differences in energetic costs of reproduction lead to competition by one sex for the other. usually, males compete for females So far, we’ve seen intrasexual selection: females mate with which ever male gets access to them Selects for male traits that allow them to dominate, intimidate or sexually outcompete rival males… 1 Intrasexual selection: competition between males 1. Direct competition for mates Forms of intrasexual selection… 2. Sperm competition: e.g., barbed penises e.g., large testes 3. Infanticide: 2 Intersexual selection: mate choice Females often mate preferentially with showy males Enlarged, ornamented traits; coloration; vocalization; other behavior. Showy male Drab female Like intrasexual selection, leads to evolution of sexual dimorphism Mate choice: female widowbirds prefer longer-tailed males Futuyma, Fig. 11.9 3 Intersexual selection and speciation Leads to trait divergence among isolated populations: role in reproductive isolation, ultimately speciation Traits favored by mate choice my compromise a male’s viability e.g., guppy coloration with predators e.g., Vocalizations in túngara frogs: Physalaemus pustulosis 4 Rapid spread of silent phenotype following parasite invasion (2006) http://www.newsroom.ucr.edu/cgi-bin/display.cgi?flash=1418 On Kauai, > 90% of male crickets now have mutant wing (“flatwing”), less than 20 generations after introduction of parasitoid fly Toothy comb (normal male) >30% parasitized Also see shift in male behavior: flatwings move near calling males to pick up females Mutant (flatwing) male: <1% parasitized 5 Mate choice: How and why do female preferences arise? Explanation 1: Direct benefits. Females choose traits with direct benefits for her or her offspring Direct fitness advantage to choosiness, so the preference is favored by natural selection. e.g., female birds choose males with best territory for resources Mate choice: How and why do female preferences arise? Explanation 1: Direct benefits. Females choose traits with direct benefits for her or her offspring Direct fitness advantage to choosiness, so the preference is favored by natural selection. e.g., female birds choose males with best territory for resources e.g., female hangingflies prefer males who provide larger prey during mating. 6 Explanation 2: Sensory bias. Intrinsic preference for particular sensory stimuli. Female’s preference precedes the evolution of the preferred male trait. e.g., Prey orientation: lizard courtship movements Explanation 2: Sensory bias. Intrinsic preference for particular sensory stimuli. Female’s preference precedes the evolution of the preferred male trait. e.g., Prey orientation: lizard courtship movements e.g., Females of swordfish fish species 1 prefer long tails even though no males possess them: preference predates the species. Species 1 Species 2 Female preference for long tails arose before Species 1 and 2 diverged from their common ancestor P+: females prefer long tails T+: males possess long tails 7 Sensory bias and female preference… Zebra finches females prefer males with an artificial white crest Explanation 3: Indirect benefits. Females prefer males with traits that indirectly benefit the female or her offspring. a) “Sexy son” hypothesis (= Runaway sexual selection) (=“Fisherian model”) Indirect fitness benefit to female who chooses attractive male: her sons will be attractive and so she’ll have more grandchildren 8 Explanation 3: Indirect benefits. Females prefer males with traits that indirectly benefit the female or her offspring. a) “Sexy son” hypothesis (= Runaway sexual selection) (=“Fisherian model”) Indirect fitness benefit to female who chooses attractive male: her sons will be attractive and so she’ll have more grandchildren Future generations will pass on the alleles for the trait (expressed in males) and the alleles for the trait preference (expressed in females). (Establishes genetic correlation between the trait (T) and preference for the trait (P)) Can lead to fixation of the trait & preference through positive frequency-dependent selection Sexy son hypothesis illustrated: sandfly Offspring of most vs least attractive males do not differ in viability or fecundity (=no direct benefits). Lutzomyia longipalpis But sons of attractive fathers have greatest mating success. 9 Indirect benefits… b) Indicators of genetic quality: ‘Good genes’ Females evolve preference for male traits associated with high fitness (viability). e.g., Female stickleback prefer males with red bellies. Red-bellies= Males are well nourished and have good immune systems (from carotenoids). Offspring are resistant to parasites: Good genes… Females prefer energetic courtship displays 10 Good genes… Male gray tree frogs with longer calls sire fitter offspring: Hyla versicolor http://video.google.com/videoplay?docid=-1630917301467247878&q=gray+tree+frog&hl=en Evolutionary Conflict between Sexes For unrelated mates: the fitness of a mate only matters to the extent that it ensures the fitness of the offspring… Intersexual selection can lead to antagonistic coevolution: selection favors traits in one sex that reduce the fitness of the other sex Evolutionary “arms race” between the sexes 11 Antagonistic coevolution… At the level of gametes: 1. Abalones (e.g., Haliotis) • Selection on sperm: increased speed at penetrating eggs (lysin protein activity) • Selection on eggs: increased resistance, to prevent polyspermy Haliotis sp. Molecular evolution of lysin gene: rapid change at nonsynonymous sites; selection to alter protein to overcome egg barrier lysin 2. Proteins that make Drosophila sperm resistant to dislodging are toxic to females. Antagonistic coevolution… At the level of individuals: Copulation leads to injury or death e.g., traumatic insemination (bedbugs) e.g., drowning of female ducks 12 Antagonistic coevolution… At the level of individuals: Copulation leads to injury or death e.g., traumatic insemination (bedbugs) e.g., drowning of female ducks May select for female traits/preferences that confer resistance to males’ inducements to mate: “Chase-away sexual selection” e.g., water striders: Males have evolve flattened abdomens that facilitate forced copulation; females have evolved spines to obstruct males’ grip Futuyma, Fig. 14.14 Chase-away sexual selection may explain why female preference for a male trait is sometimes greatest in species where the trait hasn’t yet evolved (e.g., swordtails, zebra finches) Antagonistic coevolution at the level of individuals… Sexual cannibalism >60% of the diet of female praying mantises may be males. 13 2. 1. 4. 3. Male praying mantises avoid being eaten when they can. Males are wary around hungry females: jump on her from far away to avoid opportunties of being eaten 14 But not all sexual cannibalism is coevolutionary antagonism: Australian redback spider: males encourage cannibalism Males that are cannibalized: • mate >2X longer • produce 2X as many offspring • insert genital plug. Direct benefits http://www.scar.utoronto.ca/~mandrade/video/somersault_clip_large.mov 15