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Territoriality • Territoriality is where individuals organisms maintain exclusive use of an area that contains some limiting resource • Individuals actively defend their territory, which is costly • Benefits gained from maintaining territory, must outweigh costs Competition for Space Sexual Selection • Sexual selection is a type of natural selection, driven by male/female relationships Sexual Selection • Secondary sexual characteristics: horns, long feathers, coloration Reproductive competition • Sexual selection occurs when individuals compete for mating opportunities involves both intrasexual and intersexual selection leads to evolution of secondary sexual characteristics Intersexual selection • Intersexual selection Benefits of mate choice Males may help rear young, gather food, defend nest, etc.. • Indirect benefits Females may choose healthiest or oldest males overall genetic or physiological health more vigorous offspring Reproductive strategies • Animals use different reproductive strategies to increase reproductive success Mate choice Parental investment Mating systems Parental investment and mate choice • Mate choice occurs when individuals do not mate at random, but appear to make decisions on mates base on quality common in females • Parental investment refers to contributions each sex makes in producing and rearing offspring usually higher in females different selective pressures gametes Parenting behavior • Enhancing the survival of offspring can increase parents’ reproductive success • Parental behavior comes at a cost; drains time and effort that could be spent producing additional offspring Male reproductive strategy • Males produce energetically inexpensive sperm and often provide no parental care • Males maximize reproductive success by mating with as many females as possible (quantity) Female reproductive strategy • Females produce large, energetically expensive eggs and often provide parental care • Females increase reproductive success by increasing the quality of their mates Choosy females • Female choice can dictate rules of male competition and shape male behavior • Selects for traits in males that appeal to females, and relate to their “vigor” • Male hangingflies offer nuptial gifts (better chances with a large gift!) • Some controversy as to why females are choosy, e.g handicap theory Sexual selection leads to sexual dimorphism • Secondary sexual characteristics evolve to attract members of the opposite sex Intersexual Selection Handicap hypothesis Only genetically-superior males can survive with a handicap Sensory exploitation involves evolution in males of an attractive signal that exploits preexisting biases Intrasexual selection • Intrasexual selection Individuals of one sex compete for the opportunity to mate with individuals of the other sex Selection will strongly favor sexual dimorphism. Sperm competition Male Contests • Females of some species (e.g. elk) cluster in groups • Males of such species may fight one another for access to harems • Selects for large males that can defeat other males in contests • Lekking is form of male contest Mating systems • Three types Monogamy Polygyny Polyandry • Behavioral ecologists are using DNA fingerprinting to find the benefits of different mating system strategies Are finding that extra-pair copulations are important! Mating systems • Extra-pair copulations Researchers found that in one study, 20% of red-winged blackbird offspring were a result of extra-pair copulations. May be very pervasive Males benefit by increased mating success. Females may benefit by increased rearing assistance. Mating systems • Number of mates monogamy - one male and one female polygyny - one male and many females polyandry - one female and several males • Needs of offspring altricial - require extensive, prolonged care precocial - require little parental care Sociality and altruism Benefits of group living • Improved detection and repulsion of predators Sawflies feed gregariously Produce a noxious regurgitant when irritated Selfish herd • A group held together by self-interest • Other members of the group form a living shield against predators • Individuals may compete for the safest spots Dominance hierarchy • Some individuals accept subordinate status to others • Dominant members have higher reproductive success than subordinates • Subordinates wait their turn, or become “sneaky” Costs of group living • Increased competition for food, mates, and other limiting resources • Increased vulnerability to disease and parasitism • Risk of exploitation by other group members Selection for altruistic behavior • Nonbreeding helpers are found in mammals, birds, and insects • Altruists apparently sacrifice their reproductive success to help others • How are genes for altruism perpetuated? Factors favoring altruism • Altruism - performance of an action that benefits another individual at a cost to the actor (nest helpers) Natural selection would seem to argue against altruism Such acts may not be truly altruistic, and may be benefiting the actor Nest helpers may gain parenting experience or inherit territory Altruism • Reciprocity - Individuals may form partnerships in which mutual exchanges of altruistic acts occur. • Kin selection - By directing aid toward close genetic relatives, an altruist may increase reproductive success of its relatives enough to compensate for the reduction in its own fitness. The more closely related the individuals, the more likely the potential genetic gain. Theory of kin or indirect selection • Proposed by William Hamilton • Genes associated with caring for relatives may be favored by selection • Altruists pass on genes indirectly, by helping relatives who share close genetic similarity to survive and reproduce Kin selection (or, why some organisms are nice to their relatives) • Premise: altruism can increase in frequency if altruistic individual is related to recipient • Kin selection is natural selection that favors the spread of alleles that increase the indirect component of fitness Inclusive fitness • Includes direct fitness and indirect fitness • Fitness can accrue two ways: Descendant kin (own offspring) Collateral kin (cousins, nieces, nephews) • Helping either descendant or collateral kin will lead to copies of own alleles surviving Inclusive fitness • Inclusive fitness = fitness via descendant kin + fitness via collateral kin = own offspring + offspring sum effects on r = relatedness production to others of others • Organisms should maximize inclusive fitness Coefficient of relatedness • The probability that homologous alleles in two individuals are identical by descent Quantifying relatedness rparent-offspring = 1/2 rsiblings = 1/2 raunt-nephew = 1/4 General case: (1/2)n, where n = number of links Hamilton’s rule • An allele for altruistic behavior will spread if rB > C or Br-C > 0 B is benefit to recipient C is cost to the actor B and C are measured in units of surviving offspring Evolution of social systems • Society - group of organisms of the same species organized in a cooperative manner • Insects All ants, some bees, some wasps, and all termites are eusocial. Social insect colonies are composed of different castes of workers that differ in size and morphology and have different tasks to perform. Social insects • Workers in colonies of social insects are sterile • These colonies are extended families • Workers pass on their genes indirectly by helping relatives reproduce Termites • Workers and soldiers are sterile • A single queen and one or more kings are the parents of the entire colony Eusocial hymenoptera • Haplo-diploid sex determination • Females diploid, males haploid • r queen—worker = ½ • r worker—worker = ¾ (share all genes from dad) Eusocial hymenoptera Mother Sister Father Brother • Raising sisters increases inclusive fitness more than reproducing yourself, or raising brothers Naked Mole Rats • Only mammals known to have a sterile worker caste • Single queen mates with one to three males • DNA evidence shows clan members are all closely related Naked mole rats also exhibit altruism