Assignment 4
Assignment 4: Constructing an Ethogram and Hypotheses about Animal Behavior
1. The goal of this assignment is to have you apply your knowledge of behavioral ecology to behaviors that you personally observe. a. First, you will quantify behavioral observations using an ethogram. You will use your ethogram to construct a “behavior budget” that describes how often the animal performs each behavior. b. Second, you will generate a hypothesis about your behavioral observations and propose a way to experimentally test your hypothesis. Assignment 4
Example Ethogram from MacNulty 2007 Assignment 4
Assignment 4: Constructing an Ethogram and Hypotheses about Animal Behavior
5. Develop a hypothesis or set of hypotheses about one or more of the behaviors that you observed. The hypothesis should deal with a reason why the animal that you observed exhibited the behavior or behavioral budget that you documented. You should have both a null and alternative hypothesis.
6. Develop a hypothetical experiment or set of experiments that could be used to test your hypothesis. A good source of inspiration of possible experiments are the studies discussed in class. Note that your experiments should be feasible for one person to do with a budget of up to $10,000 (the budget of many graduate student projects) in the time frame of one year. Proposing to do a breeding experiment with elephant seals is not feasible because they are not easily manipulated. Love Darts in Snails and Slugs
1. During copulation, partners place large spermatophores in the spermatophore-receiving (SRO) organ of their partner.
2. The spermatophore is then digested in the SRO. Only ~0.1% of the spermatozoa escapes to where fertilization occurs.
3. Individuals shoot each other with love darts. The love dart is laced with hormones and increases the number of escaping spermatozoa from the SRO
4. By darting its partner, a snail can increase its chances of paternity.
5. The Japanese Euhadra subnimbosa uses its darts to stab its partner >3000 times during 22 min of ‘foreplay’ before copulation. Diversity in Love Darts
Lesson: Sexual conflict can lead to diversification. Love Darts in Snails and Slugs
1. Darts have evolved multiple times, and have been lost multiple times.
2. Darting species may be locked in sexually antagonistic selection which leads to an arms race of defense/offense.
a. SRO complexity correlates positively with dart shape (e.g. the presence of flanges).
Hermaphroditic flatworm, Pseudobiceros bedfordi. Have two penises, which they fence with. Each tries to smear the other with sperm, but tries to avoid getting smeared. Hermaphroditic flatworm, Pseudobiceros bedfordi.
St = areas with sperm Hermaphroditic flatworm, Pseudobiceros bedfordi.
H = holes caused by sperm, Sc = Scars Hermaphroditic flatworm, Pseudobiceros bedfordi.
* = Large hole where large amount of sperm hit Body then tore in two! Lessons
1. Sexual conflict can be costly. Can impose a large selective force. Can lead to partner manipulation.
2. Even hermaphrodites have different interests when mating. Each is trying to maximize its individual fitness.
3. Sexual antagonism can select for diversity in weapons and mating behaviors. Mediating the Conflict
1. Many hermaphrodites inseminate reciprocally. This “sperm trading” ensures reciprocity during mating.
2. Test: Create cheaters by vasectomizing one partner of hermaphroditic sea slugs (Study by Anthes, 2005)
3. Result: Partners of the cheaters were less likely to trade intromissions/inseminations, and were more likely to terminate a mating encounter. Further Reading on Wacky Mating Systems: Dr. Tatiana’s Sex Advice to all Creation
Lecture Outline: Mating Systems
Diversity among species Basic Categories of Mating Systems and How they are determined by ecology.
Diversity within species Alternative mating strategies and their evolution.
Reminder: Assignment 2 is due on Wednesday Basic Categories of Mating Systems
1. Monogamy: A male and female form a pair bond (short or long term). Often both parents care for eggs/young.
2. Polygyny: A male mates with several females. Females mate with just one male. Usually female provides parental care.
3. Polyandry: A female mates with several males. Often the male provides parental care.
4. Promiscuity: Both males and females mate several times with different individuals. Either sex may care for the young.
5. Polygamy: a general term for when an individual of either sex has more than one mate. Ecological Theory of Mating System Evolution Emlen and Oring, 1977
1. Mating systems depend on the ability of a portion of the population to control the access of others to potential mates.
2. Multiple mates must be energetically defendable by individuals a. Even distribution of mates = little polygamy potential. b. Patchy distribution = high polygamy potential. Ecological Theory of Mating System Evolution Emlen and Oring, 1977
3. Benefits of defense must outweigh the costs. a. Energy spent defending or seeking mates comes at the cost of parental care.
4. Environmental factors determine the costs and benefits. For instance, how females are spaced.
5. Ecology constrains the degree to which sexual selection can operate. Monogamy
1. Sexual selection theory suggests that a male’s reproductive success is usually related to the number of females he inseminates.
2. Why monogamy? Hypotheses for Monogamy
1. Mate guarding hypothesis: Monogamy may be adaptive if a female left by the male would probably acquire another mate who would fertilize her eggs.
2. Mate guarding likely to occur if females are receptive after mating, widely scattered, and difficult to locate.
3. Example: Clown shrimp, females are widely dispersed, males spend weeks with a female. Hypotheses for Monogamy
1. Mate assistance hypothesis: Males stay with a single female to help rear their mutual offspring.
2. The additional offspring that survive due to paternal care may more than compensate the male for the chance to reproduce with other females.
3. Example: Seahorse Hippocampus whitei, pairs ignore any others of the opposite sex. The male’s pouch can only hold 1 clutch of eggs, so as long as his pouch is full he is maximizing his reproduction.
4. Example: Monogamous California mice, males help rear offspring. ~2 offspring born. With male: 1.5 on average survive. Without male: .6 on average survive. Hypotheses for Monogamy
1. Female enforced monogamy: Females attempt to block their partners’ polygynous moves in order to monopolize their parental assistance.
2. Example: Razorbills (a seabird), females attack their partner if they show interest in a neighbor. Monogamy
2. Example: Burying beetles. Male and female bury a mouse to feed offspring. Once buried, the male may release a pheromone to attract other females. His mate pushes him from his perch, reducing his ability to signal. (from Eggert and Skaluk, 1995). Social vs Genetic Monogamy
1. About 90% of all birds are socially monogamous: form long-term partnerships during a breeding system. a. Male birds (unlike many mammals) can increase fitness substantially by helping with eggs/nestlings. Mate assistance hypothesis.
2. In some birds social monogamy = genetic monogamy. Ex. Common loons and Florida scrub jays.
3. Many socially monogamous birds have extra pair copulations (EPCs). Not genetically monogamous.
4. EPCs benefit males by allowing him to sire offspring. Tradeoff is while he is gone, he gives up mate guarding. Females and EPCs
1. What do females gain from EPCs?
2. Good genes hypothesis: Gain good genes for offspring. a. Female tits seek EPCs if mated to an unattractive partner (ie the partner gets few EPCs).
3. Fertility insurance hypothesis: EPCs reduce risk of having infertile parter. Test = look at fertilization success. a. Gunnison’s praire dogs: Polyandrous females: pregnant 100% of time, monogamous females: pregnant 92% of the time.
3. More resources hypothesis: Mating with multiple males allows access to more resources. a. Dunnocks: Females seek out a subordinate mate. Will copulate 100s of times for a clutch of eggs. Both males help rear the offspring. Polyandry
1. Galapagos hawks: As many as 8 males may pair with a female, helping her rear a single offspring per breeding episode.
a. Suitable breeding territories are scarce, males may cooperate to keep other males out, all males have an equal chance of fertilization.
Polyandry
1. Spotted sandpipers a. Females fight other females for territories. b. A territory may attract more than one male who incubate a single female’s clutches. c. This “sex role reversal” could be due to female only being able to lay 4 eggs at a time. If resources are abundant, they need another mate to take care of another clutch. d. Food is often abundant.
Female Defense Polygyny
1. Female defense polygyny: Females occur in defensible clusters, males compete for control of the clusters.
2. Examples: a. Male Greater Spear-Nosed Bats defend roosting females. Successful defenders may sire up to 50 offspring. Female Defense Polygyny b. A marine amphipod: Forms a house out of shells and pebbles. Males capture females and glue their houses to his! Resource Defense Polygyny
1. Resource defense polygyny: a. Females do not live together permanently. b. Male may become polygynous if he controls a rich patch of resources that the female’s visit
2. Examples: a. Male black-winged damselflies defend floating vegetation. Females mate with the male and lay eggs on the vegetation. Resource Defense Polygyny
Example 2 a. African cichlid fish, Lamprologus callipterus. Need safe location for eggs. b. Female lays eggs in an empty snail shell, and guards them. c. Males collect shells, sometimes stealing them from rival males. Observed to collect up to 86 shells. Up to 14 females nesting on one male’s territory Testing Resource Defense Polygyny
1. Predicts that female distribution is controlled by key resources
2. Test = Manipulate the resources, see if females alter their locations.
3. Example: Dunnocks (a songbird), Davies and Lundberg a. Normally females search for food widely. This means that female go on multiple males territories. They are polyandrous. b. Experiment: Supplemented some home ranges with food. Testing Resource Defense Polygyny
Result: With supplemented food, females decreased their range size. This decreased the number of mates that they had. Males were able to monopolize females. Scramble Competition Polygyny
1. Scramble competition polygyny: Females are widely dispersed, which reduces the benefits of territorial behavior. Males try to outrace rivals to receptive females.
2. Examples: a. Male thirteen-lined squirrels search widely for females that are receptive for 4-5 hours during the breeding season. Males bias their searching at places where a female was about to go into estrous. Scramble Competition Polygyny
Example 2: Explosive breeding assemblages
a. Horseshoe crabs: Females lay eggs during just a few nights. Males scramble to get a mate.
b. Wood frogs: Females lay eggs in just one or a few nights. The high density of females and rival males does not make territoriality feasible. Lek Polygyny
1. Lek Polygyny: Sometimes males advertise to females with elaborate visual, acoustic, or olfactory displays. Males do not hunt for mates. Females watch males display at territories that do not contain food, nesting sites, or anything useful.
2. Sometimes males aggregate into groups and each male defends a small territory that contains no resources at all-sometimes just a bare patch of ground. a. When territories are clumped in a display area = a lek.
2. Male mating success is highly skewed on lecks a. Manakins: males jump between perches, snapping. feathers. At a lek of 10, there were 438 copulations. One male = 75%, second male = 13%, six others = 10%. Leks
Mating success if usually strongly skewed on leks with the majority of matings going to a small proportion of males.
36 Leks
1. Leks have been reported in only 7 species of mammals and 35 species of birds
2. Thought to occur when males are unable to defend economically either the females themselves or the resources they require a. In both antelope and grouse, the lekking species are those with the largest female home ranges b. In Uganda kob, topi and fallow deer, males lek at high population densities but defend territories or harems at low densities Why Lek?
1. Why do males all congregate to display? Lots of competition.
2. Hotspot hypothesis: females tend to travel along certain routes and males congregate where routes intersect.
3. Hotshot hypothesis: subordinate males cluster around highly attractive males hoping to interact with females attracted to the hotshots.
4. Test: Remove males that were successful in attracting females. a. Hotspot prediction = other males will move into best site. b. Hotshot prediction = subordinates will leave site.
5. Example: Great snipe. Removal of dominant male caused subordinates to leave. Removal of a subordinate resulted in another subordinate coming in. Why Lek?
6. Female preference hypothesis: females prefer groups of large males where they can more quickly compare the quality of males.
7. Test: Look at whether reproductive success varies with number of males at lek.
8. Example: Ruffs (a sandpiper). More males attract more females up to a point. After 6 or more males, the dominant’s reproductive success goes down. Why Lek?
9. Reducing male predation hypothesis: As lek size increases likelihood of any one male being predated decreases (dilution effect) a. Tungara frogs are safer from bat predation when calling in large choruses (panel a).
10. Increased female attraction hypothesis: By displaying together males provide greater attraction for females and draw them in from a greater distance a. Tungara frog payoffs per individual male increased with lek size (panel b).
11. These hypotheses are not exclusive and many be important depending on species and spatial scale. Alternative Mating Systems
1. Monopolization of females by males, leaves many males without a mate.
2. This can favor the evolution of sneaking strategies. Often get “female mimics.”
3. Traits favored for sneaking may differ from those of territorial males. Often different morphs become markedly distinct.
4. Selection is often frequency dependent selection. The rare types have a fitness advantage. This leads to the stable maintenance of multiple morphs. a. Prediction: over time the fitnesses of the strategies should be about equal.
5. Can be polymorphic, polyphenic, or plastically determined. Polymorphic side-blotched lizard strategies
Orange Morph: Territory Usurper Large size Large territories Tradeoff: Low survival to next year
Yellow Morph: Sneaker Blue Morph: Mate Guarder Mimics females Small territories Not territorial
Sinervo & Lively 1996, Zamudio & Sinervo 2000, Sinervo et al 2000, Sinervo & Svensson 2002, Sinervo & Clobert, 2003, Sinervo et. al. 2006 Selection on males is frequency dependent: rock-paper-scissors Alternative Mating Strategies
Marine Isopod Morphs
Shuster and Wade
1. Male types are a polymorphism. a. Alpha males: Have pincers and defend sponges. b. Beta males: Female mimics c. Gamma males: Invest in sperm, dive into matings.
2. Mating success approximately equal: Alpha = 1.51 mates, Beta = 1.35 mates, gamma = 1.37 mates. Dung Beetle Mating Strategies
Emlen & Nijhout 1. Male types are a polyphenism. Morphology (horns/no horns) is plastically determined by the amount of dung the larva was raised on.
2. Mating behavior is variable: defend or sneak.
3. Developmental tradeoff: Big horns can mean small eyes! Plastic Natterjack Toad Mating Strategies
1. Male types are plastic. Males make the “best of a bad job” = makes best of poor circumstances by adopting alternative strategy.
2. Large males call loudly and attract the most females. 60% were callers, but got 80% of matings.
3. Small males are “satellites” and attempt to intercept females. Will switch behaviors when they get bigger. Also, if remove the large males, then they will call.