1. Proximate and Ultimate Causes of Animal Behaviour

2. The beewolf wasp(Philanthus triangulum)

3. Beewolf wasp burrows

4. Niko Tinbergen: one of the founders of ethology

8. The Scientific Method • Observation and causal question • Hypothesis • Predictions • Test of predictions • Conclusion

9. The Scientific Method • Observation and causal question: Beewolves find the entrance to their own nests. How do they do it? • Hypothesis: They use local landmarks as cues • Predictions: If they use local landmarks then manipulating those landmarks should mislead them • Test of predictions: Experiments • Conclusion: Yes, local landmarks are used by beewolves as cues in finding own nest

10. Black-headed gull … and its chicks

12. The Scientific Method • Observationand causal question - black headed gulls remove eggshells. Why did black headed gulls evolve behaviour of eggshell removal? • Hypothesis - maybe this behaviour helps parent gulls to conceal their offspring from predators • Predictions - if so, broken eggshells should help predators find food • Test of predictions - experiment • Conclusion - yes, eggshell removal appears to help parent gulls conceal offspring from predators

13. Proximate causes Ultimate causes of behavour

14. Oystercatcher Kittiwake Black-headed gull

15. Oystercatcher Solitary nesting Nest predators Immediate eggshell removal Kittiwake Group nesting No nest predators No eggshell removal Black-headed gull Group nesting Nest predation present Delayed eggshell removal

16. Why do babies cry???