1. Resource competition Ideal free distribution Dominance –Correlates –Hierarchies –Hormonal effects Territoriality

2. Ideal free distribution Premise: animals sequentially fill available habitat starting with best patches Assumptions –“Ideal” by possessing perfect information about resource quality –“Free” to disperse appropriately Expectation –Input matching rule: animals disperse to equalize energy intake or reproductive success

3. Ideal Free Distribution

4. Ideal free distribution experiments Stickleback fish: 6 fish in a tank consume Daphnia introduced from the ends with an initial probability ratio of 2:1; after the start of feeding (first arrow) the number of fish in patch 2 quickly approaches the number predicted by the input matching rule (dotted line). At the second arrow, the patch probabilities are reversed (Milinski 1979). Mallard ducks: similar experiment in which 33 ducks were fed pieces of bread from two stations with probability ratio of 2:1 around a pond. The number of ducks at the poorer station quickly approaches the input matching rule prediction (Harper 1982).

5. Deviations from IFD 16 of 20 studies show too many in poor habitat and too few in rich habitat (Sutherland 1986) Perceptual error –Expected if animals choose habitat at random Differences in competitive abilities –Dominants exclude subordinates

6. Competitive differences exist

7. Dominance - why? Regulates access to resources without continual fighting Should be tolerated if the potential risk of injury from fighting exceeds the potential gain from resource acquisition

8. Dominance - how? Ability to control access to a resource is termed resource holding potential (RHP) Correlates include –body size –experience –matrilineal relationships –fat reserves –prior success (or failure) Requires individual recognition or status badges

9. Dominance and size

10. Dominance and age

11. Dominance and residence

12. Matrilineal dominance Daughters outrank older sisters in

13. Dominance and fat reserves No effect of weight But, males with more energy reserves win aerial chases to determine ownership of mating territories along streams

14. Dominance hiearchies Linear, transitive Linear hierarchies are difficult to explain in large groups because RHP should be difficult to rank accurately among average individuals. Intransitive Alliances

15. Mechanisms for linear hierarchies Animals recognize each other using signals that correlate with RHP - status badges –Harris sparrows, house sparrows Animals adjust their perceived status on the basis of the outcome of recent contests (winner-loser effects) –fishes Animals adjust their perceived status after observing contests involving others (bystander effects) –chickens

16. Status badges in Harris sparrows Dominance correlates with size of black patch on head and chest

17. Why not fake it? After Before Darkened birds were attacked more often!

18. Hormones and dominance Winners often have elevated testosterone levels Losers have elevated plasma corticosterone levels compared to winners In lobsters, losers can be made aggressive by infusing serotonin. This effect can be blocked by Prozac, a serotonin inhibitor

19. Effects of testosterone on male vertebrates

20. Steroid hormone synthesis Stress hormoneFemale sexual functionMale sexual function

21. Economics of territoriality Resource must be defensible –Renewable, e.g. nectar, emerging insects –not ephemeral or superabundant Benefits > Costs of defense –Energetic costs increase with Density of intruders Territory size –Benefits accrue by Increasing energy intake rate Reducing search costs Reducing starvation risk

22. Territoriality scales with body weight and diet

23. Optimal territory size

24. Territory size should be adjusted to maximize energy gain © 1999 Bob Pelham

25. Territory size should depend on resource availability Eastern ArizonaNW California

26. Territories should occur at intermediate resource densities

27. VariableSalmon Creek Doran Beach Territory length 41 ± 1082 ± 14 Intruder density 11.5 ± 7.40.5 ± 0.4 Prey density 580 ± 160671 ± 130 Prey density -> Intruder pressure -> territory size (Myers) Territory size should respond to competition

28. Testosterone increases competitive ability

29. Testosterone alters territoriality, at a cost, in juncos Males defend larger territoriesMales feed nestlings less