Psy 5120, 2012 p. 1
Psy 5120S, 2012: Issues in Comparative Cognition
Instructor: Sara J. Shettleworth
Telephone: 416 978-7819 email: shettle@psych.utoronto.ca website: http://www.psych.utoronto.ca/users/shettle/
Office: Sidney Smith Hall 4054 course web page: linked to homepage of above
Office hours: drop in or by appointment (I am usually around in the early afternoon)
This course covers selected areas of current research, theorizing, and debate in comparative cognition. The focus is on behavioral studies with nonhuman animals, although we will also touch on their relevance for understanding the brain, brain evolution and cognition in humans.
Course requirements
400-500-word reaction papers on required readings (generally 1/week) class participation
Essay, due on Tuesday, April 10, 2012
40%
20%
40%
The purpose of the reaction papers is to encourage you to make reading and thinking about the material a priority so everyone gets the most out of the seminars. Sometimes I will provide the topic; at other times it will be up to you to identify an issue in the readings worthy of comment. A hard copy of each reaction paper must be submitted at the beginning of the relevant class. They will not be accepted late. Zero will be given for each one missed, but the one lowest reaction paper mark will be disregarded.
Occasionally there may be opportunity to give a 10-minute talk to the class on an extra reading such as a current article related to the week’s topic. A talk may be substituted for that week’s reaction paper.
Website and readings: There is a course web page linked to http://www.psych.utoronto.ca/users/shettle/
The assignments for each week will be posted here as well as links to anything on the reading list not readily available in the U of T’s e-journal collection.
Textbooks: Shettleworth, S. J. (2012). Fundamentals of Comparative Cognition. New York: Oxford
University Press, in press (members of the class will receive pdfs of the page proofs). Sections will be assigned as background reading throughout the course. For more depth and detail you may also want to consult (or to buy) Shettleworth, S. J. (2010). Cognition, Evolution, and Behavior , 2 nd edition.
New York:
Oxford University Press. Paperback available in the U of T textbook store. 1 copy is on 3-hour loan in the
Earth Sciences Library.
Essay . Basically you can write on almost any topic related to the course. This includes comparative aspects of your own research. See the last page of this handout for specifications
Email policy. Short email queries will generally be answered within 24 hours on weekdays, within a couple of days on weekends. Longer discussions of issues raised by the readings should be reserved for class or a personal appointment. So your messages will not be treated as spam you should use a U of T e-address that includes your name and head your message with a specific mention of the course.
Printing policy: Please submit anything to be marked in hard copy, double spaced to give me room to write comments, but printed on both sides of the paper to save trees. For tips on double sided printing, see the page of the UTBiology Environment Action Team: http://utbeat.biology.utoronto.ca/
Notwithstanding this paper-saving policy, please print out the readings and bring the hard copies to class.
Laptops (i.e. for reading pdfs) will not be welcome in the seminar (a policy we can discuss).

Psy 5120, 2012 p. 2
Dates, topics, and readings
Readings should be done in the order listed. * before an item means it can be found on the class web page, linked to http://www.psych.utoronto.ca/users/shettle/
1. Jan. 10 INTRODUCTION TO THE COURSE
2. Jan. 17 NO CLASS. Read the little Fundamentals text to get an overview of the material in the course and think about what you might like to do your essay on.
3. Jan. 24 HISTORY, CURRENT TRENDS AND BIG ISSUES.
The “big issues” are the roles of anthropomorphism, Lloyd Morgan’s canon, and appeals to consciousness in studying comparative cognition.
Fundamentals , Chapter 1 (pp. 1-16)
Wynne, C. D. L. (2004). The perils of anthropomorphism. Nature, 428 , 606, and the reply to it by Burghardt,
Nature, 430 , 15.
Shettleworth, S. J. (2010). Clever animals and killjoy explanations in comparative psychology. Trends in
Cognitive Sciences, 14 , 477-481.
* Heyes, C.M. (2008). Beast machines? Questions of animal consciousness. In M. Davies & L. Weiskrantz
(Eds). Frontiers of Consciousness . Oxford University Press. Pdf of the published chapter will be linked to course site or Google Cecilia Heyes and find a preprint under publications, book chapters.
*Sober, E. (2005). Comparative psychology meets evolutionary biology. Morgan's canon and cladistic parsimony. In L. Daston & G. Mitman (Eds.), Thinking with animals: New perspectives on anthropomorphism (pp. 85-99). New York: Columbia University Press. Handout or pdf (tbd).
4. Jan. 31: EVOLUTION AND THE NATURE OF COMPARISON
Biological and theoretical foundations of comparative psychology. What do we compare, how do we compare it, and what do we hope to learn from doing so?
Papini, M. (2002). Pattern and process in the evolution of learning. Psychological Review, 109 , 186-201.
Emery, N. J., & Clayton, N. S. (2004). The mentality of crows: Convergent evolution of intelligence in corvids and apes. Science, 306 , 1903-1907.
MacLean, E. L., Matthews, L. J., Hare, B. A., Nunn, C. L., Anderson, R. C., et al. (2011). How does cognition evolve?: Phylogenetic comparative psychology. Animal Cognition, in press (if not out by
January, use the “Springerlink” option for the online journal and look in “online first”).
Amici, F., Aureli, F., & Call, J. (2008). Fission-fusion dynamics, behavioral flexibility, and inhibitory control in primates. Current Biology, 18 , 1415-1419. Be sure to study the linked Methods that are only available online.

Psy 5120, 2012 p. 3
5. Feb. 7. MEMORY AND CONSCIOUSNESS: EPISODIC-LIKE MEMORY AND
METACOGNITION.
Fundamentals Chapter 2, pp. 28 -34 only.
Clayton, N. S., & Dickinson, A. (1998). Episodic-like memory during cache recovery by scrub jays. Nature,
395 , 272-274.
Fortin, N. J., Wright, S. P., & Eichenbaum, H. (2004). Recollection-like memory retrieval in rats is dependent on the hippocampus. Nature, 431 , 188-191.
Eacott, M. J., & Easton, A. (2010). Episodic memory in animals: Remembering which occasion.
Neuropsychologia, 48 , 2273-2280.
Hampton, R. R. (2001). Rhesus monkeys know when they remember. Proceedings of the National Academy of Sciences (USA), 98 , 5359-5362.
Smith, J. D., & Washburn, D. A. (2005). Uncertainty monitoring and metacognition by animals. Current
Directions in Psychological Science , 14 , 19-24.
Terrace, H. S., & Son, L. K. (2009). Comparative metacognition. Current Opinion in Neurobiology, 19 , 67-
74.
Hampton, R. R. (2009). Multiple demonstrations of metacognition in nonhumans: Converging evidence or multiple mechanisms? Comparative Cognition & Behavior Reviews, 4 , 17-28.
6. Feb. 14 PLANNING AND REINFORCMENT: MENTAL TIME TRAVEL IN ANIMALS?
Fundamentals Chapter 2, pp. 34-41 (Associative Learning) and Chapter 3, pp. 66-72
Roberts, W. A. (2002). Are animals stuck in time? Psychological Bulletin, 128 , 473-489.
Mulcahy, N. J., & Call, J. (2006). Apes save tools for future use. Science, 312 , 1038-1040.
Also read (of course!) the online supplementary methods.
*Dickinson, A. (2011). Goal-directed behaviour and future planning in animals. In R. Menzel & J. Fischer
(Eds.). Animal Thinking: Contemporary Issues in Comparative Cognition (pp. 79-91). Cambridge
MA: MIT Press. Pdf will be linked to the course web page.
Suddendorf, T., & Corballis, M. C. (2010). Behavioural evidence for mental time travel in nonhuman animals. Behavioural Brain Research, 215 , 292-298.
Cheke, L. G., & Clayton, N. S. (2011). Eurasian jays ( Garrulus gladarius ) overcome their current desires to anticipate future needs and plan for them appropriately. Biology Letters, in press (or may be out)
READING WEEK

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7. Feb. 28. TOOL USE AND PHYSICAL UNDERSTANDING
What animals use tools and what cognitive processes are involved?
Fundamentals , pp. 73-80
Seed, A., & Byrne, R. (2010). Animal tool-use. Current Biology, 20 , R1032-R1039.
Seed, A. M., Call, J., Emery, N. J., & Clayton, N. S. (2009). Chimpanzees solve the trap problem when the confound of tool-use is removed. Journal of Experimental
Psychology: Animal Behavior Processes, 35 , 23-34.
Silva, F. J. & Silva, K. M. (2006). Humans’ folk physics is not enough to explain their tool-using behavior. Psychonomic Bulletin & Review, 13, 689-693.
Bluff, L. A., Troscianko, J., Weir, A. A. S., Kacelnik, A., & Rutz, C. (2010). Tool use by wild New
Caledonian crows Corvus moneduloides at natural foraging sites. Proceedings of The Royal Society of London B, 277 , 1377-1385.
Wimpenny, J. H., Weir, A. A. S., Clayton, L., Rutz, C., & Kacelnik, A. (2009). Cognitive processes associated with sequential tool use in New Caledonian crows. PLoS One, 4 , e6471.
8. March 6. SPACE
Spatial cognition is a huge area of research on ants, bees, rats, birds, and humans, among other species. A central issue has been “do animals have cognitive maps?” We will look at a historic paper that addresses it along with some recent research on people and honeybees.
Fundamentals Chapter 3, pp. 49-57 only; Chapter 5, pp. 118-120 (on modularity)
Tolman, E. C. (1948). Cognitive maps in rats and men. Psychological Review, 55 , 189-208.
Full text with figures online at: http://psychclassics.yorku.ca/Tolman/Maps/maps.htm
Wang, R. F., & Spelke, E. S. (2002). Human spatial representation: Insights from animals. Trends in Cognitive Sciences, 6 , 376-382.
Landau, B., & Lakusta, L. (2009). Spatial representation across species: geometry, language, and maps. Current Opinion in Neurobiology, 19 , 12-19.
(added Jan 31): Esch, H. E., S. Zhang, M. V. Srinivasan, and J. Tautz. (2001). “Honeybee dances communicate distances measured by optic flow.”
Nature 411: 581–583.
Menzel, R. et al. (2005). Honey bees navigate according to a map-like spatial memory.
Proceedings of the National Academy of Sciences (USA), 102 , 3040-3045.
Menzel, R., et al. (2011). A common frame of reference for learned and communicated vectors in honeybee navigation. Current Biology, 21 , 645-650.

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9. Mar. 13 NUMBER
What numerical abilities do other species share with humans and which are uniquely human? How are they related to the ability to represent other relative magnitudes in general?
Fundamentals , pp. 41-48 (concepts and categories); pp. 61-69 (number and economic decisions).
Feigenson, L., Dehaene, S., & Spelke, E. (2004). Core systems of number. Trends in Cognitive Sciences, 8 ,
307-314.
Hauser, M. D., Carey, S., & Hauser, L. (2000). Spontaneous number representation in semi free ranging rhesus monkeys. Proceedings of the Royal Society of London, 267 , 829-833.
Pica, P., Lemer, C., Izard, V., & Dehaene, S. (2004). Exact and approximate arithmetic in an Amazonian indigene group. Science, 306 , 499-503.
Cantlon, J. F., Platt, M. L., & Brannon, E. M. (2009). Beyond the number domain. Trends in Cognitive
Sciences, 13 , 83-89.
Terrace, H. S (2005). The simultaneous chain: A new approach to serial learning. Trends in Cognitive
Sciences, 9, 202-210.
Grosenick, L., Clement, T. S., & Fernald, R. D. (2007). Fish can infer social rank by observation alone.
Nature, 445 , 429-432.
Pompilio, L., & Kacelnik, A. (2010). Context-dependent utility overrides absolute memory as a determinant of choice. Proceedings of the National Academy of Sciences, 107 , 508-512.
10. Mar. 20. SOCIAL COGNITION: THEORY OF MIND
Fundamentals , pp. 81-92.
Sheehan, M. J., & Tibbetts, E. A. (2011). Specialized face learning is associated with individual recognition in paper wasps. Science, 334 , 1272-1275.
Call, J., and M. Tomasello. (2008). “Does the chimpanzee have a theory of mind? 30 years later.”
Trends in
Cognitive Science 12: 187–192.
Penn, D. C., & Povinelli, D. J. (2007). On the lack of evidence that non-human animals possess anything remotely resembling a 'theory of mind'. Philosophical Transactions of the Royal Society B, 362 , 731-
744.
Bugnyar, T., & Heinrich, B. (2005). Ravens, Corvus corax , differentiate between knowledgeable and ignorant competitors. Proceedings of the Royal Society of London B, 272, 1641-1646.
* Cheney, D. L. (2011). Cooperation in nonhuman primates: Function and cognition. In R. Menzel & J.
Fischer (Eds.), Animal thinking: Contemporary issues in comparative cognition (pp. 239-265).
Cambridge, MA: MIT Press.
Cheney, D. L., Moscovice, L. R., Heesen, M., Mundry, R., & Seyfarth, R. M. (2010). Contingent

Psy 5120, 2012 p. 6 cooperation between wild female baboons. Proceedings of the national Academy of Sciences, 107 ,
9562-9566.
Bshary, R., and A. S. Grutter. (2006). “Image scoring and cooperation in a cleaner fish mutualism.”
Nature
441: 975–978.
11. Mar. 27. SOCIAL LEARNING AND CULTURE
Fundamentals , pp. 93-101
Catmur, C., Walsh, V., & Heyes, C. (2009). Associative sequence learning: the role of experience in the development of imitation and the mirror system. Philosophical Transactions of the Royal Society B,
364 , 2369-2380.
Thornton, A., & McAuliffe, K. (2006). Teaching in wild meerkats. Science, 313 , 227-229.
Whiten, A. et al. (1999). Cultures in chimpanzees. Nature , 399 , 682-685.
* Galef, B. G. (2009). Culture in animals? In The Question of Animal Culture , ed. K. N. Laland and B. G.
Galef Jr., 222-246. Cambridge, MA: Harvard University Press.
Tennie, C., Call, J., & Tomasello, M. (2009). Ratcheting up the ratchet: on the evolution of cumulative culture. Philosophical Transactions of the Royal Society (London) B, 364 , 2405-2415.
12. April 3. COMMUNICATION
Fundamentals , pp. 101-111.
Seyfarth, R. M., & Cheney, D. L. (2010). Production, usage, and comprehension in animal vocalizations.
Brain & Language , 115 , 92-100.
Zuberbuhler, K., Cheney, D. L., & Seyfarth, R. M. (1999). Conceptual semantics in a nonhuman primate.
Journal of Comparative Psychology, 113 , 33-42.
Evans, C.S. & Evans, L. (2007). Representational signaling in birds. Biology Letters, 3, 8-11.
Hauser, M. D., Chomsky, N., & Fitch, W. T. (2002). The faculty of language: What is it, who has it, and how did it evolve? Science , 298 , 1569-1579. van Heijningen, C. A. A., deVisser, J., Zuidema, W., & ten Cate, C. (2009). Simple rules can explain discrimination of putative syntactic structures by a songbird species. Proceedings of the National
Academy of Sciences, 106 , 20538-20543.
PAPERS DUE ON TUESDAY APRIL 10

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Psy 5120, 2012: The Essay
The required essay can be on any topic related to the course. For example, you can use it to explore comparative aspects of your own research problem. It should be 12 – 15 pages long, plus cover page (if any) and bibliography. It should include a review and critical analysis of literature on the problem you choose and should conclude with an assessment of the most important gap(s) in knowledge and a proposal for further work. This need not be a specific experimental design but could be a well-justified recommendation for which direction(s) research on this problem should go in, for example, more field observations of a certain kind, more comparative studies on certain kinds of species, etc. Your contribution, in terms of choice of problem, organization, and analysis, as well as clarity of presentation, will be an important component of the mark.
Topics may include, but are not restricted to, further exploration of topics covered in the required.
However, although you can take off from the required readings or class discussion, credit obviously cannot be given for extensive repetition of material covered in class.
Please note that because this course is about the comparative study of cognitive processes and only peripherally about the neurobiology of these processes, if you choose to look at connections to neuroscience you should focus on the cognitive/behavioral aspects of the “animal model(s)” you choose. For instance, if the topic were “is the hippocampus a cognitive map?” it would be very important to analyze whether the behavioral tests used and the effects lesioned or genetically modified animals show in them actually are good evidence for cognitive mapping.
A double-spaced hard copy is officially due at 1 PM on Tuesday, April 10, 2012 ( 1 week after the last class).
Please note that it is now Psychology Graduate Committee policy that students who submit assignments after the course submission deadline should not receive a mark of A+.
You are strongly encouraged to submit an information proposal for the essay, consisting of about a page stating the question to be addressed and giving one or two key references. Feedback will be given within a few days. This process may all take place by email and would be appropriate to initiate whenever you are beginning to think about your topic.
Office hours: No formal office hours are scheduled for this course, but you can get in touch at any time to discuss your essay topic or anything else and I will usually be able to meet within a day or 2. Or feel free to drop by – I’m usually to be found in Sid Smith 4054 in the early afternoons.