Core Concepts in Animal Behavior – Winter 2013                                                                               11 March 2013


Michael Beecher

Professor of Psychology & Biology

Guthrie 327, 543-6545


PSYCH 502, 3 credits (graded)                   Tues Thurs 11:00-12:20 – Guthrie 315          



We will discuss the major concepts and research approaches in the contemporary study of animal behavior, focusing on topics that lie at the interface of animal behavior, evolutionary biology, neurobiology and psychology. Each week will focus on one topic. We will read both classic and ‘hot new’ papers. Because there are more core concepts than there are weeks in the course, I have left some out and have left some play in the schedule. In particular one week is open for a topic that can any one of you can suggest. In addition, appropriate papers for the other weeks can be suggested.


Course requirements and student responsibilities: Because this is a graduate seminar course, there will be no exams; there will be no term paper either. But the other side of the coin is that the student’s presence and active participation are de rigueur. Your grade will depend entirely on participation in the actual class sessions. Each week’s papers will be divided up among the class participants so that each student will have one specific assignment each week besides being responsible for reading the rest of the material. Any student participating fully in the course will get a 4.0. You need to let me know if there is a class you cannot attend (e.g., because you are going to a scientific conference). Unexcused absences are pretty much the only route to a less-than-4.0 grade.


Usually it will make sense to bring a ppt presentation to class to illustrate the paper (or part of a paper) on which you are the ‘point person’. Therefore I will bring a projector (and also a laptop if you request) to each class session.











Jan 8, 10

History of the field

  to ~1960

  to present

Wynne-Edwards (1963)

Wynne-Edwards; Maynard Smith; Perrins (1964) 

Williams (1966)

Krebs & Davies extract


Jan 15, 17

Altruism & Kin Selection

  Slides for Sherman, Emlen, Krakauer

  Kin Recognition

Sherman (1977)

Emlen et al (1995)

Krakauer (2005)

Manning et al (1992)


Jan 22, 24

Altruism: Reciprocity, Limited Control & Concession Models

Wilkinson (1992)

Carter & Wilkinson (2012)

Fischer (1988)

Clutton-Brock (1998); Clutton-Brock et al (2001)

Snyder-Mackler et al 2012


Jan 29, 31

Eusociality & the Return of Group Selection      Caglar lecture notes

Wilson & Wilson (2007)

Queller & Strassman (1998)

West-Eberhard (1987)

Ratnieks & Wenseleers (2008)


Feb 5, 7

Sexual Selection

Krebs & Davies, Chapt 8

Andersson, M. (1982)

Wilkinson & Reillo (1994)

Gonzalez-Voyer et al (2008)


Feb 12, 17


Searcy & Nowicki (2005)

Laidre (2009)

Logue et al (2010)

Backwell et al (2000)


Feb 19, 21

Animal Cognition

Gould (2004)

Herrmann et al (2007)

Dean et al (2012)

de Waal (2012)


Feb 26

Feb 28

Animal Cognition

Evolutionary Psychology

de Waal (2008)

Haidt & Kesebir (2010)


Mar 01

Mar 03

Evolutionary Psychology

Integrative Approaches

Tooby & Cosmides: EP Primer  (see also optionals below)

Dioniak et al (2006)

Kotrschal et al (2013)

Zanette et al (2011)


Mar 08, 10

Integrative Approaches

Fernald & Maruska (2012)

Van Oers et al (2004)

Glocker et al (2009)

Musso et al (2003)


Readings for Winter 2013               *optional reading           **really optional, usually added after class

Altruism & Kin Selection

Williams, G. C. (1966).  Adaptation and Natural Selection, Princeton University Press (chapt 1).

Wynne-Edwards, V. C. (1963). Intergroup selection in the evolution of social systems. Nature 200: 623-626.

Wynne-Edwards, V., Maynard Smith, J., & Perrins, C. (1964).  Group selection and kin selection; Survival of young swifts in relation to brood-size. Nature  201: 1145-1149.

Krebs, J. R. & Davies, N. B. (1981). pp 14-18 in An Introduction to Behavioural Ecology, Blackwell (3rd edition 1993).

Classic Empirical studies on Altruism & Kin Selection

Sherman, P.W. (1977). Nepotism and the evolution of alarm calls. Science 197: 1246-1253.

Emlen, S.T., Wrege, P.W., and Demong, N.J. (1995). Making decisions in the family: an evolutionary perspective. American Scientist 83: 148-157.   bigger better pdf

Krakauer, A. H. (2005). Kin selection and cooperative courtship in wild turkeys. Nature 434: 69-72.

Manning, C. J., Wakeland, E. K. & Potts, W. K. (1992). Communal nesting patterns in mice implicate MHC genes in kin recognition. Nature 360: 581-583.

* Holmes, W. G. (2004). The early history of Hamiltonian-based research on kin recognition. Annales Zoologici Fennici 41: 691-711.

* Ruff, J. S., Nelson, A. C., Kubinak, J. L. & Potts, W. K. (2012). MHC signaling during social communication. In Self and Nonself,  Carlos LópezLarrea, C. (Ed.). Landes Bioscience.

Classic Empirical studies on Reciprocal Altrusim, Limited-Control & Concession Models

Wilkinson, G. S. (1984). Reciprocal food sharing in the vampire bat. Nature 308: 181-184.

Carter GG, Wilkinson GS. (2013). Food sharing in vampire bats: reciprocal help predicts donations more than relatedness or harassment. Proceedings of the Royal Society B  280: 2573.

* Wilkinson, G. S. (1990). Food sharing in vampire bats. Scientific American.

Fischer, E. A. (1988). Simultaneous hermaphroditism, tit-for-tat, and the evolutionary stability of social systems. Ethology & Sociobiology 9: 119-136.

Clutton-Brock, T. (1998). Reproductive skew, concessions and limited control. Trends in Ecology & Evolution 13: 288-292.

Clutton-Brock, T. et al. (2001). Cooperation, control, and concession in meerkat groups. Science 291: 478-481.

Snyder-Mackler, N., Alberts, S. C. & Bergman, T. J. (2012) Concessions of an an alpha male? Cooperative defence and shared reproduction in multi-male primate groups. Proceedings of the Royal Society B  279: 3788-3795.

Eusociality & the Return of Group Selection

Queller, D. C. & Strassman, J. E. (1998). Kin selection and social insects. Bioscience 48: 165-175.

Wilson, D. S. & Wilson, E. O. (2007). Rethinking the theoretical foundation of sociobiology. 82: 327-348.

West Eberhard, M. J. (1987). Flexible strategy and social evolution. pp 35-51 in Ito, Y., Brown, J. L. & Kikkawa, J. (eds.) Animal Societies: Theories and Facts.

Ratnieks, F. L. W. & Wenseleers, T. (2006). Altruism in insect colonies and beyond: voluntary or enforced. Trends in Ecology & Evolution 23: 45-52.

* Strassman, J. E. & Queller, D. C. (2007). Insect societies as divided organisms: the complexities of purpose and cross-purpose. Proceedings of the National Academy of Sciences 104: 8619-8626.

* Nowak, M. A., Tarnita, C. E. & Wilson, E. O. (2010). The evolution of eusociality. Nature 466: 1057-1062. [Note that I’ve attached the supplementary material to the article itself. Consider that to have a **!]

* Many mad-as-hornets authors (2011). Inclusive fitness theory and eusociality. Nature 471: E1-10.

** Alexander, R. D. (1974).  The evolution of social behavior. Annual Review of Ecology & Systematics 5: 325-383.

Sexual Selection

Andersson, M. (1982). Female choice selects for extreme tail length in a widowbird. Nature 299: 818-820.

Wilkinson, G. S. & Reillo, P. R. (1994). Female choice response to artificial selection on an exaggerated male trait in a stalk-eye fly. Proceedings of the Royal Society B 255: 1-6.

Gonzalez-Voyer, A., Fitzpatrick, J. L. & Kolm, N. (2008). Sexual selection determines parental care patterns in cichlid fishes. Evolution 62: 2015-2026.

 [see also Communication below]


Searcy, W. A. & Nowicki, S. (2005). “Introduction” from The Evolution of Animal Communication: Reliability and Deception in Signaling Systems.  Princeton.

Laidre, M. E. (2009). How often do animals lie about their intentions? An experimental test. American Naturalist 173, 337-346.

Logue, D. M. et al (2010). Does signalling mitigate the cost of agonistic intentions? A test in a cricket that has lost its song. Proceedings of the Royal Society B 277: 2571-2575.

Backwell et al (2000). Dishonest signalling in a fiddler crab. Proceedings of the Royal Society B 267: 719-724.

* Rendall, D., Owren, M. J. & Ryan, M. J. (2009). What do animal signals mean? Animal Behaviour 78: 233-240.

* Seyfarth, R. L. et al. (2010). The central importance of information in studies of animal communication. Animal Behaviour 80: 3-8.

* Ryan, M. J. (1991). Sexual selection and communication in frogs. Trends in Ecology & Evolution 6: 351-356.

* Dawkins, R. & Krebs, J. R. (1978). Animal signals: Information or manipulation. In J. R. Krebs & N. B. Davies (eds.), Behavioural Ecology : An Evolutionary Approach.  Sinauer.

** Kimball, R. T., Braun, E. L., Ligon, J. D., Lucchini, V. & Randi, E. (2001). A molecular phylogeny of the peacock-pheasants (Galliformes: Polyplectron spp.) indicates loss and reduction of ornamental traits and display behaviours. Biological Journal of the Linnean Society 73: 187-198.

** Ron, S. R. (2008). The evolution of female mate choice for complex calls in tungara frogs. Animal Behaviour 76: 1783-1794.

** Meyer, A., Morrissey, J. M. & Schartl, M. (1994). Recurrent origin of a sexual selected trait in Xiphophorus fishes inferred from a molecular phylogeny. Nature 368: 539-542.

** Alan Grafen’s ppt on the handicap model:

** See also Carl Bergstrom’s web module on honest signalling:

Animal Cognition (Cognitive Ethology)

Gould, J. L. (2004). Thinking about thinking: how Donald R. Griffin (1915–2003) remade animal behavior. Animal Cognition 7: 1–4.

Herrmann, E., Call, J., Hernandez-Lloreda, M. V., Hare, B. & Tomasello, M.  (2007).  Humans have evolved specialized skills of social cognition:  The cultural intelligence Hypothesis.  Science 317: 1360-1366.

Dean, L. G., Kendal, R. L., Schapiro, S. J. & Laland, K. N.  (2012).  Identification of the social and cognitive processes underlying human cumulative culture. Science 335: 1114-1118.

de Waal, F. B. M. (2012).  The antiquity of empathy. Science 336: 874-876.

de Waal, F. B. M. (2008).  Putting the altruism back into altruism: The evolution of empathy. Annual Review of Psychology 59: 279-300.

Evolutionary Psychology

Haidt, J. & Kesebir, S. (2010). Morality.  In S. Fiske, D. Gilber & G. Lindzey (eds.), Handbook of Social Psychology, pp 797-832.

Cosmides, L. & Toobey, J. (1997). Evolutionary Psychology: A Primer. [online]

* Daly, M. & Wilson, M. (1999). Human evolutionary psychology and animal behaviour.  Animal Behaviour 57: 509–519.

* Smith, E. A., Borgerhoff Mulder, M. & Hill, K. (2000). Evolutionary analyses of human behaviour: A commentary on Daly & Wilson. Animal Behaviour 60: F21-26.

Integrative Approaches

Dioniak, S. M., French, J. A. & Holekamp, K. E. (2006). Rank-related maternal effects of androgens on behaviour in wild spotted hyaenas. Nature 440: 1190-1193.

Kotrschal, A. et al (2013). Artificial selection on relative brain size in the guppy reveals costs and benefits of evolving a larger brain. Current Biology 23: 1-4.

Zanette, L. Y., White, A. F., Allen, M. C. & Clinchy, M. (2011). Perceived predation risk reduces the number of offspring songbirds produce per year. Science 334: 1398-1401.

Fernald, R. D. and Karen P. Maruska. (2012). Social information changes the brain.  Proceedings of the National Academy of Sciences 109: 17194–17199.

van Oers, K., Piet J. Drent, P. J., de Goede, P. & van Noordwijk, A. J. (2004). Realized heritability and repeatability of risk-taking behaviour in relation to avian personalities. Proceedings of the Royal Society B 271: 65-73.

Glocker, M. L. et al. (2009).  Baby schema modulates the brain reward system in nulliparous women. Proceedings of the National Academy of Sciences 106: 9115-9119.

Musso, M. et al. (2003). Broca’s area and the language instinct. Nature Neuroscience 6: 774-781.