Ecology Lecture 10 Life History Patterns 1 Topics covered (both Life History Lectures) Sexual selection What criteria do individuals use to choose mates? Mating systems How many mates does an individual have What factors determine this? Parental care/parental investment in offspring Allocation of resources Quick reproduction vs. growth & long life (r and K strategists, etc…) Relevance to Population Ecology Life-history characteristics determine Organization of individuals in space and time How the population perpetuates itself Examples… discretely vs. continuously; rapidly vs. slowly, etc… Which characteristics will be selected for and persist within the population (due to sexual selection) Conversely, selective pressures (abiotic factors, distribution of resources, etc..) will affect the life-history patterns observed Sexual selection Defined: Selection for characteristics/behaviors that maximize chances of mating and producing the most, and highest quality, offspring. A category within “natural selection” General pattern: Male-male competition and female choice Common pattern with many exceptions! Example: Satin bowerbird mating behavior Each male build an elaborate bower where he conducts his courtship display Bowers located near each other Each female visits several times, finally chooses a mate Bowerbird males: # of mates Some males much more successful than others Bowerbird females: # of mates Only ~1/3 of females have >1 mate Why do males usually compete, while females choose? Hypothesis 1, A.J. Bateman: “Eggs are expensive, sperm is cheap!” Amount of energy invested in a single gamete is much greater for females (eggs) than for males (sperm) Female bird may invest up to 30% of body weight in eggs. “Eggs are expensive, sperm is cheap.” Difference in investment per gamete Example: Fairy wrens Males have 8 billion sperm in testes at once Females lay six eggs maximum per clutch Female limited by egg production, Male limited by number of mates only (presumably unlimited sperm) Operational sex ratio skewed toward males Is sperm really cheap? It often takes a large number of sperm to fertilize a single egg, due to Hostile environment within female Acid Attacks by the immune system Is sperm really cheap? Sperm competition among males Occurs when females have multiple mates Possibly the predominant situation Fitness advantages for female (will explore in next lecture) Some males may actually run out of sperm… Garter snakes, zebra finch, blue crabs, rams… What if there is no sperm competition? In sea horses, eggs are deposited into pouches, and there is not sperm competition. Why not? Male sea horses have relatively low sperm counts! Is sperm really cheap? Drosophila bifurca: one sperm with long tail Sperm tail is 20x length of his body His testes make up 11% of his body mass. Why do males usually compete, while females choose? Hypothesis 2 (R. Trivers): Competition vs. choice is based on individual with the most total parental investment Often the female (example: mammals) But in some species, male makes a greater total investment Gulf pipefish While male cares for a single brood, a female can produce two clutches of eggs male has greater total parental investment Operational sex ratio skewed toward females. Males choose large, ( ornamented females over small, drab ones. Female Male Types of sexual selection Intrasexual selection: maintenance of traits that assist in competition within the gender successful mating Intersexual selection: maintenance of traits that are attractive to the opposite gender Intrasexual selection 1: Adaptations to gain access to females Dominance behavior & characteristics Example 1: sexual dimorphism in elephant seals Male-male competition and sexual dimorphism (seals) Male dung beetle, Phanaeus vindex (Rattlebox photography) Intrasexual selection 1: Adaptations to gain access to females Weaponry for fighting with other males. Example: dung beetle! Intrasexual selection 1: Adaptations to gain access to females Sneaker strategies Example 1: Plainfin midshipmen Dominant male features and behavior Nest building, singing, guarding Sneaker male features and behavior No nest, no singing, just sex… Small fish, big balls! Dung beetles: two morphs Behavioral and morphologial differences similar to midshipmen Large, dominant males with horns defend burrows Small, hornless males with “big balls” sneak Midshipmen Genetically-based differences Dung beetles Nutritionally-based differences Intrasexual selection 2: Adaptations favoring the use of one’s sperm Displacing or inactivating rival sperm Damselfly “scooper” penis Intrasexual selection 2: Adaptations favoring the use of one’s sperm Displacing or inactivating rival sperm Example: Chemical sperm inactivation in fruit flies Photo: San Francisco Exploratorium Intrasexual selection 2: Adaptations favoring the use of one’s sperm Mechanisms to avoid sperm displacement Mate guarding (Example: many crab species) Intrasexual selection 2: Adaptations favoring the use of one’s sperm Mechanisms to avoid sperm displacement Prolonged mating and cannibalism (example: redback spider) Female less likely to mate with another if she eats him He has low likelihood of finding a new mate (high predation) Andrade, 1996 Intrasexual selection 2: Adaptations favoring the use of one’s sperm Mechanisms to avoid sperm displacement Anti-aphrodisiac (Example: Heliconius erato) Intersexual selection Focus on female choice Material benefits: Nutrition Ex: hangflies) Length of mating time depends on quality and size of “courtship gift” Intersexual selection Focus on female choice Material benefits: Anti-predator substances Defensive compounds in arctiid moths Photo: butterfly-conservation.org Intersexual selection Focus on female choice Ability of males to provide sufficient sperm Female fruit flies (some species) choose virgin males Photo: San Francisco Exploratorium Intersexual selection Focus on female choice Parental ability Cannot assess directly May be correlated with other features of the male Example 1: Redwing blackbird “Epaulettes” correlated with nest defense Courtship effort correlated with feeding effort Photo: Vancouverislandbirds.com Intersexual selection Focus on female choice Parental ability (Ex: sedge warbler) Size of song repertoire correlates with chick weight at fledging Sedge warbler (cont.) Female sedge warblers choose on the basis of repertoire size. Thus they choose the most fit males Intersexual selection Focus on female choice Health/Genetic quality Example: song repertoire in great reed warbler Females chose males with larger repertoires This was correlated with greater offspring survival (unrelated to parental care) Intersexual selection Focus on female choice Health/Genetic quality Example: Bright coloration of sticklebacks negatively correlated with low parasite loads Intersexual selection Focus on female choice Health/Genetic quality (bright color negatively correlated with parasite load) Advantages to females choosing these males Avoid getting parasites while mating Avoid transferring parasites to young Are choosing healthier males; their health status may be related to “genetic quality” Satin Bowerbirds: multiple signals of health and fitness (and good genes?) Origin/maintenance of mate choice for “exaggerated” characteristics In some species, why do males develop what appear to be extreme traits that actually can hamper their survival? Example: Peacock’s tail Origin/maintenance of mate choice for “exaggerated” characteristics Hypothesis 1: Runaway selection (R.A. Fisher) Directional that takes on a life of its own Starts as an “honest signal” more extreme. Mechanism: Females choose males with large tails, multiple eyespots. the next generation has a higher proportion of these males. Will work even if his traits are not honest signals of quality. Why? Evidence of arbitrary choices by females (bird band example) Origin/maintenance of mate choice for “exaggerated” characteristics Hypothesis 2: Handicap or “good genes” hypothesis (R.A. Fisher) Exaggerated trait might decrease chance of survival, only males with superior genes can survive despite the handicap Example: peacock tail as a handicap. In this case, a female choosing a male with these traits would be improving her fitness. (His signal is an “honest signal” of fitness.) Distinguishing between Runaway Selection and “Good Genes” (Petrie) Methods Males of different ornamentation/tail length and randomly bred them with females Young raised under identical conditions and then released Results Why random? Offspring of the “attractive” males weighed more at day 84. Offspring of the attractive males were more likely to be alive after two years Which hypothesis is supported by this data? Petrie’s peacock data Origin/maintenance of mate choice for “exaggerated” characteristics How extreme can a characteristic become? Under what conditions will directional selection stop? (Think about costs vs. benefits…)