1. Behavioral Ecology Photo from Wikimedia Commons

2. Ethological Underpinnings of Behavioral Ecology Photo from http://www.dabase.org/lorenz.htm Konrad Lorenz Instinct, imprinting, etc.

3. Photo of Tinbergen from Wikimedia Commons Niko Tinbergen Four questions subsumed under Proximate vs. Ultimate Causes; questions concerning, respectively, how a behavior is produced and why it evolved (i.e., evolutionary Benefit / Cost Ratio) Ethological Underpinnings of Behavioral Ecology

4. Of his chosen study organism von Frisch said: “The honey bee is like a magic well: the more you draw from it, the more there is to draw.” Karl von Frisch Photo of von Frisch from Wikimedia Commons Waggle Dance

5. Photo from Wikimedia Commons Konrad Lorenz Niko Tinbergen Karl von Frisch Nobel Prize – 1973 Ethological Underpinnings of Behavioral Ecology

6. E.g., artificial selection experiments suggest a genetic basis for “migratory activity” Genes can influence behavior, so behavior can evolve Pulido (2007) BioScience, Fig. 2 Artificial Selection For higher proportion of migrants For lower proportion of migrants

7. Foraging Behavior Photo from Wikimedia Commons E.g., ambush predator and female fly prey (also illustrates another cost of sex)

8. Items with high profitability (P) are generally preferred Optimal Foraging Theory Photo from http://www.rspb.org.uk/community/ourwork/b/biodiversity/archive/2013/02/18/ guest-blog-in-the-still-of-the-night.aspx P = E t E = net energy value, i.e., energy gained minus energy invested t = encounter time & handling time invested in obtaining & processing the food

9. Conceptual model of OFT Optimal Foraging Theory Net energy gained = (Total energy obtained) – (Cumulative energy investment) Drops off as animal cannot carry nor ingest more Cain, Bowman & Hacker (2014), Fig. 8.6

10. Marginal Value Theorem as applied to profitability of foraging patches Cain, Bowman & Hacker (2014), Fig. 8.8 Within a patch, the marginal value for longer time has diminishing returns Slopes of straight, solid lines = Energy gained / time Tangent maximizes profitability (slope) & determines optimal giving up time Optimal Foraging Theory

11. Foraging (and other) decisions can be modified by predators The Ecology of Fear Beckerman et al. (1997) Proceedings of the National Academy of Sciences, Fig. 1 E.g., caged grasshoppers foraging in the presence or absence of the risk of predation, i.e., with or without a spider (mean  s.e.m. shown)

12. Prey sometimes communicate their awareness of predators to those predators E.g., stotting / pronking The Ecology of Fear Photo from Wikimedia Commons

13. Social Behavior Photo of social grooming from Wikimedia Commons

14. E.g., Optimal Group Size Consider the variable Benefit / Cost Ratio Social Behavior Cain, Bowman & Hacker (2014), Fig. 8.22 Should an individual remain alone or join another to form a group of 2? What is the optimum group size? Should an individual join a group of 2 or 5? What are likely benefits and costs?

15. Reproductive Behavior Photo from Wikimedia Commons

16. The Evolution of Competitive Males & Choosy Females (and sometimes the reverse) Parental Investment is “any investment by the parent in an individual offspring that increases the offspring's chance of surviving (and hence reproductive success) at the cost of the parent's ability to invest in other offspring” (Trivers 1972) Photomicrograph of human egg and sperm cells from Cain, Bowman & Hacker (2014), Fig. 7.7; photo of suckling manatee from http://mammalssuck.blogspot.com/2013/11/mega-mammal-milk-analysis.html; photo of “pregnant” seahorses from http://www.scubadiveasia.com/blog/best-dad-award-goes-to-the-seahorse/ Anisogamy Maternal investment = nursing Paternal investment = brood-pouch “pregnancy”

17. Male-Male Competition Photomicrographs from http://prometheuswiki.publish.csiro.au/tiki-index.php?page= Spikelet+sterility+and+in+vivo+pollen+germination+and+tube+growth+under+high-temperature+stress+in+rice E.g., male pollen grains compete to fertilize female ovules

18. Photo of peacock spider (Maratus volans) from Wikimedia Commons Female Choice – Courtship E.g., courtship in the peacock spider (Maratus speciosus)courtship in the peacock spider (Maratus speciosus)

19. Copulatory Courtship & Cryptic Female Choice Photo of Maria Fernanda Cardosa’s sculptures of male damselfly genitalia from http://livingwithinsects.wordpress.com/2012/04/30/insect-reproductive-morphology/ E.g., male damselfly genitalia (aedeagi, plural of aedeagus)

20. Monogamy Polygyny Polyandry Promiscuity Mating Systems Photo of horseshoe crabs from Wikimedia Commons

21. “Polygyny occurs if environmental or behavioral conditions bring about the clumping of females, and males have the capacity to monopolize them.” Emlen & Oring (1977) Mating Systems Cain, Bowman & Hacker (2014), Fig. 8.8; schema from Emlen & Oring (1977) Science, Fig. 1

22. Polygyny Threshold Model Mating Systems Graphic model from Orians (1969) American Naturalist, Fig. 1 Pick a point on the monogamous female curve. The distance to the right to intercept the bigamous female curve is the polygyny threshold, i.e., the habitat quality increase required to make it worthwhile for the female to share a mate.

23. W. D. Hamilton Inclusive Fitness & Kin Selection Photo of Hamilton from Wikimedia Commons Kin selection exposes the selfish nature of altruism; helping kin can increase one’s inclusive fitness (direct plus indirect fitness) Hamilton’s Rule: rB > C Relatedness * (Benefits to recipient) > (Costs to altruist)

24. Relatedness “I would lay down my life for 2 brothers or 8 cousins” J. B. S. Haldane r – introduced by Sewell Wright as a measure of consanguinity Generation 1 Generation 2 Generation 3 Mother- daughter r = 1/2 Sister r = 1/2 Cousin r = 1/8

25. Eusociality in Diploid Organisms Photos from Wikimedia Commons For most individuals in the colony the benefits to helping the queen outweigh the costs of sacrificing their own reproduction rB > C Naked Mole RatTermites

26. Eusociality in Haplodiploid Organisms Photos of Hymenoptera from Wikimedia Commons For most female individuals in the colony the benefits to helping the queen rear sisters outweigh the costs of sacrificing their own reproduction rB > C Generation 1 Generation 2 Mother- daughter r = 1/2 Sister r = 3/4

27. Adoption Aggression Anti-Predator Behavior Begging Breeding Brood Parasitism Cannibalism Communication Cooperation Copulation Dispersal Dominance Hierarchies Family Dynamics Flocking Grooming Habitat Selection Herding Homing Behavioral Ecology Infanticide Kin Recognition Mate Guarding Migration Nepotism Nesting Parasite Avoidance Parental Care Playing Predator-Prey Interactions Roosting Scent-Marking Sex Change Schooling Symbiotic Maintenance Territoriality Thermoregulation Etc…