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1 January 2005 William Brewster Memorial Award, 2004: Russell P. Balda and Alan C. Kamil

Russell P. Balda and Alan C. Kamil combined their training in animal behavior, ecology, evolution, and comparative psychology to pioneer the study of avian cognition and contribute significantly to our understanding of corvid biology. More than two decades ago, they saw the potential in the fact that seed-caching birds are self-motivated to return to inconspicuous points in the landscape to retrieve hidden food, and they began to employ the cache-recovery paradigm as a model system for studying learning, memory, and “spatial information processing.” Their unusually fruitful collaboration has led to enormous advances in the understanding of spatial learning in birds and, in the previous decade, to the publication of approximately 50 scientific papers.

The Balda—Kamil partnership's careful consideration of the ecological context and evolutionary history of bird behavior was enhanced by their use of a wider battery of methodologies and perspectives than either an ecologist or a comparative psychologist working alone might employ. By asking their birds to cache in arrays of sand-filled cups, for example, they were able to control for spacing or site preferences of caches and thus assess the importance of fixed patterns of movement and mnemonic devices in enabling cache retrieval (Kamil and Balda 1990).

They demonstrated that spatial cognition of related corvid species is correlated with differences in natural history and the degree to which species depend on stored food for winter survival and breeding: Clark's Nutcrackers (Nucifraga columbiana) and Pinyon Jays (Gymnorhinus cyanocephalus) outperform the less challenged Western Scrub-Jays (Aphelocoma californica) and Mexican Jays (A. ultramarina) (Balda et al. 1996, Balda and Kamil 1998). They found that the exceptional spatial memory of Clark's Nutcracker is coupled with a disproportionately large hippocampal volume (Basil et al. 1996). They also developed an analog of the radial-arm maze, which allowed them to compare the memory of various seed-caching and non-seed-caching species (e.g. Kamil et al. 1994).

Balda and Kamil also found that spatial memory of cache sites in Clark's Nutcracker is highly resistant to interference from other spatial memory tasks (Bednekoff et al. 1997). Together with Peter Bednekoff, they linked corvid sociality to observational spatial memory, demonstrating that the highly social Pinyon Jay may be more adept at learning cache sites by observing other birds than the more solitary nutcracker and Western Scrub-Jay (Bednekoff and Balda. 1996a, b6). Those studies not only demonstrated that the spatial memory of corvids is extraordinary in both capacity and duration, but they helped develop a general understanding of the adaptive nature of avian cognition.

In addition to their work together, Balda and Kamil have made many contributions independently and with other collaborators. For example, Alan Kamil has conducted important studies on the effects of visual predators (Blue Jays [Cyanocitta cristata]) on prey crypticity and how predator choice can function to maintain prey polymorphism (Bond and Kamil 1998, 2002). Russ Balda, on the other hand, has made important contributions to the ecology, breeding biology, and sociality of Pinyon Jays and other birds in a long series of studies in the field (e.g. Balda 2002). Their joint research has been funded by the National Science Foundation throughout the past two decades, and they continue today to make important new contributions to the study of avian behavior.

For their ground-breaking work on North American corvid biology, especially cognition, memory, learning, and seed-caching, and associated aspects of social behavior, The American Ornithologists' Union is pleased to present the 2004 Brewster Memorial Award to Russell P. Balda and Alan C. Kamil.

Award criteria

The William Brewster Memorial Award consists of a medal and an honorarium provided through the endowed William Brewster Memorial Fund of the American Ornithologists' Union. It is given annually to the author or coauthors (not previously so honored) of the most meritorious body of work on birds of the Western Hemisphere published during the 10 calendar years preceding a given AOU meeting.

Literature Cited


R. P. Balda 2002. Pinyon Jay (Gymnorhinus cyanocephalus). In The Birds of North America, no. 605 (A. Poole and F. Gill, Eds.). Birds of North America, Philadelphia. Google Scholar


R. P. Balda and A. C. Kamil . 1998. The ecology and evolution of spatial memory in corvids of the southwestern USA: The perplexing Pinyon Jay. Pages 29–64 in Animal Cognition in Nature (R. P. Balda, I. M. Pepperberg, and A. C. Kamil, Eds.). Academic Press, London. Google Scholar


R. P. Balda, A. C. Kamil, and P. A. Bednekoff . 1996. Predicting cognitive capacity from natural history: Examples from four species of corvids. Pages 33–66 in Current Ornithology, vol. 13 (V. Nolan, Jr., and E. D. Ketterson, Eds.). Plenum Press, New York. Google Scholar


J. A. Basil, A. C. Kamil, R. P. Balda, and K. V. Fite . 1996. Differences in hippocampus volume among food storing corvids. Brain, Behavior and Evolution 47:156–164. Google Scholar


P. A. Bednekoff and R. P. Balda . 1996a. Observational spatial memory in Clark's Nutcrackers and Mexican Jays. Animal Behaviour 52:833–839. Google Scholar


P. A. Bednekoff and R. P. Balda . 1996b. Social caching and observational spatial memory in Pinyon Jays. Behaviour 133:807–826. Google Scholar


P. A. Bednekoff, A. C. Kamil, and R. P. Balda . 1997. Clark's Nutcracker (Aves: Corvidae) spatial memory: Interference effects on cache recovery performance? Ethology 103:554–565. Google Scholar


A. B. Bond and A. C. Kamil . 1998. Apostatic selection by Blue Jays produces balanced polymorphism in virtual prey. Nature 395:594–596. Google Scholar


A. B. Bond and A. C. Kamil . 2002. Visual predators select for crypticity and polymorphism in virtual prey. Nature 415:609–613. Google Scholar


A. C. Kamil and R. P. Balda . 1990. Spatial memory in seed-caching corvids. Pages 1–25. in The Psychology of Learning and Motivation, vol. 26 (G. H. Bower, Ed.). Academic Press, New York. Google Scholar


A. C. Kamil, R. P. Balda, and D. J. Olson . 1994. Performance of four seed-caching corvid species in the radial-arm maze analog. Journal of Comparative Psychology 108:385–393. Google Scholar


Russell P. Balda (right) and Alan C. Kamil: March, 2004. Photo by Judith Balda.

"William Brewster Memorial Award, 2004: Russell P. Balda and Alan C. Kamil," The Auk 122(1), 359-361, (1 January 2005).[0359:WBMARP]2.0.CO;2
Published: 1 January 2005
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