University of Richmond

Dr. Peter Smallwood

Associate Professor of Biology
A-110 Gottwald Science Center
Office: (804) 289-8803
Fax: (804) 289-8233

Dr. Smallwood is on leave for spring and fall semester of 2008.  He is in Kabul, serving as country director for the Wildlife Conservation Society's project in Afghanistan. He was recently featured on National Public Radio and in The Collegian for his work abroad.

Research:
I am a behavioral ecologist by training. My work extends from studies of the behavior of individual organisms to plant-animal interactions and their community-wide effects. I have studied the foraging strategies of long jawed spiders, as well as their breeding behaviors. I recently published a study of the breeding behavior of Kestrels (a small falcon). Currently, most of the work in my lab is focused on the interactions between squirrels, mice, and other small animals and the oaks they depend upon for their winter food supply. The work is usually field-based, and often involves experiments and observations of free-ranging squirrels, mice, and other small vertebrates.

Education:
Ph.D., University of Arizona

Selected Publications:
Steele, M.A., Halkin, S.L., Smallwood, P.D., McKenna, T.J., Mitsopoulos, K, and Beam, M. (2007 in press). Cache protection strategies of a scatter-hoarding rodent: do tree squirrels engage in behavioural deception?  Animal Behaviour: in press.

Michael A. Steele, John E. Carlson, Peter D. Smallwood, Amy B. McEuen, Thomas A. Contreras, and William B. Terzaghi (2007) Linking seed and seedling shadows:  A case study in the oaks (Quercus) in: Seed dispersal: Theory and its application in a changing world. (Dennis, A. J., Schupp, E. W., Green, R. and Westcott, D. W., eds)  CAB International, Wallinford, U.K.{invited chapter, peer reviewed}.  

Steele, M.A., Manierre, S., Genna, T., Contreras, T.A., Smallwood, P.D., & Pereira, M.E. (2006) The innate basis of food-hoarding decisions in grey squirrels: evidence for behavioural adaptations to the oaks. Animal Behaviour 71: 155-160.

Steele, Micheal A., and Peter D. Smallwood. 2002.  Acorn dispersal by birds and mammals. Chapt 12  in Ecology and Management of Oaks for Wildlife. (W. McShea and W. M. Healy, eds.)  Johns Hopkins University Press. {invited chapter, peer reviewed}.  
 
Danielson-Francois, A.,   C. Fetterer, and P.D. Smallwood. (2001)  Body condition and mate choice in Tetragnatha elongata.  Journal of Arachnology 30:20-30.

Smallwood, P.D.,  Steele, M.A., and Faeth, S. H. (2001) The ultimate basis of the caching preferences of rodents, and the oak-dispersal syndrome: tannins, insects, and seed germination.  American Zoologist 41: 840-851.  

Steele, M.A.,  P.D. Smallwood, A. Spunar, and E. Nelsen.  (2001) Proximate cues influencing caching decisions: Can grey squirrels detect seed dormancy?  American Zoologist 41: 852-864.

Smallwood, P. D. (2001) An integrative approach to the study of plant-animal interactions.  
American Zoologist 41:807-809. (Invited overview)

Steele, M. A., G. Turner, P.D. Smallwood, J.O. Wolff, and J. Radillo.  2001. Cache management by small mammals: Experimental evidence for the significance of acorn embryo excision.   Journal of Mammalogy 82:35-42

Smallwood, P.D., and J.A. Smallwood. (1998) Seasonal shifts in the sex ratios of fledgling American kestrels: The early bird hypothesis. Evolutionary Ecology  12:839-853.

Smallwood, P.D., M.A. Steele, E. Ribbens, and William F. McShea (1998). The effects of seed hoarders on the distribution of tree species: Grey squirrels and oaks as a model system. pp. 211-222 in (M.A. Steele, D. Zegers, and J. Merritt, eds), Ecology and Evolutionary Biology of Tree Squirrels. Virginia Museum of Natural History, special publication #6, 320pp. (invited chapter, peer reviewed).

Hadj-Chikh, L.Z.,  M.A. Steele, and P.D. Smallwood (1996)  Caching decisions by Grey Squirrels: A test of the handling time and perishability hypotheses.  Animal Behavior 52:941-8.
 
Smallwood, P.D. (1996)   An introduction to risk sensitivity: The use of Jensen's inequality to clarify evolutionary arguments of adaptation and constraint. American Zoologist 36:392-401.
    
Smallwood, P.D., and R.V. Cartar (1996) Risk sensitivity in behavior: Where are we now? American Zoologist 36:389-391 (Invited overview).

Steele, M.A., and P.D.Smallwood (1994)  What are squirrels hiding?  
Natural History 10/94: 40-45.

Smallwood, P.D. (1993)  Web-site tenure in the Long-Jawed Spider: Is it risk-sensitive foraging, or conspecific interactions?  Ecology 74: 1826-1835.

Smallwood, P.D., and W.D. Peters  (1986)  Grey squirrel food preferences: The effects of tannin and fat concentration. Ecology 67:168 174.

Presentations:
M. Shafqat, J. Cao, J. Carlson, T. Genna, K. Hicks, D. Layer, H. Marking, A. McBrearty, A. McEuen, P. Smallwood, M. Steele, D. Terzaghi, W. Terzaghi Determining parent-offspring distances in oak forests by DNA fingerprinting Presented at American Society of Plant Biologists, meeting in Seattle, WA July 16-20, 2005

Steele, M. A. J.E. Carlson, P.D. Smallwood, A.B. McEuen, T.A. Contreras, W. B. Terzaghi. M. Steele. (Invited plenary presentation) Linking seed and seedling shadows in temperate forests: A test of the differential dispersal hypothesis in oaks.  Fourth International Conference on Frugivory and Seed Dispersal, July, 2005, Brisbane, Australia (NSF sponsored)

P.D. Smallwood,  W. Terzaghi,  J.E. Carlson,  J.E. Mank,  E. Ribbens,  T.A. Contreras,  L. Caudill, and M. A. Steele.  Searching for effects of animal dispersal on the distribution of oak seedlings.   To be presented at the Annual meetings of the Ecological Society of America, 4 August 2002, Tucson, Arizona, USA.

P.D. Smallwood, W. B. Terzaghi, M.A. Steele, J.E. Carlson, and J.E. Mank.   Further investigations into the interactions between small vertebrates and oaks: A multidisciplinary approach.   Presented at the annual meetings of the Society for Integrative and Comparative Biology, 4 January 2001.  Chicago, Illinois, USA.

P.D. Smallwood, Danielson-Francois, A., and Fetterer, C.A.  Mate Choice in an orb-weaving spider: Body condition matters.  Presented at the meetings of the Animal Behavior Society, Corvalis, OR, USA.  15 July 2001. 

Ngumezi, C., T. A. Contreras, P. D. Smallwood, and M. A. Steele. Effects of acorn size and habitat structure on the location of caches by eastern Gray Squirrels (Sciurus carolinensis).   Poster, presented at the Annual Meetings of the American Society of Mammalogists,  17 June 2001,  Missoula, Montana, USA.

Smallwood, P.D., M.A. Steele, J. Brown, and J.F. Merritt.  Title: Common caching behaviors of several tree squirrel species, and their consequences.  2nd International Colloquy on the Ecology of Tree Squirrels.  7-11 May 2000, Salem, Oregon, USA.

Smallwood, P. D.,    M. A. Steele, and J. Brown.From behavioral decisions to forest structure: Caching behaviors of animals and their effects on the dispersal of oaks.    Society for Integrative and Comparative Biology, Atlanta, GA,  7 January 2000.  Society-Wide Symposium on Terrestrial Plant-Animal Interactions

Smallwood, P.D. M.A. Steele, and J. Radillo. Several small mammal species discriminate between red and white oak acorns: A general phenomenon for caching foragers. Annual meetings of the Animal Behaviour Society.  Bucknell University, Lewisburg, PA.  26 June 1999.

Smallwood, P.D.  M.A. Steele, and Juan Radillo. Discriminating between red and white oak acorns: A general phenomenon for caching foragers?   Annual Meetings of the Society for Integrative and Comparative Biology.  Denver, CO,  6 January 1999.

Smallwood, P.D., M.A. Steele.  Squirrel foraging behavior: ultimate causes and consequences.  Semi-annual meetings of the  International Society for Behavioral Ecology.  Asilomar, CA, 31 July 1998.

Smallwood, P. D.  The misuse of the concept of heritability by people who know better: US!   Oral presentation to the Division of Comparative Issues, Society for Integrative and Comparative Biology.  Boston, MA, 5 January 1998.

Smallwood, P. D.,  E. Ribbens, and M.A. Steele. Title: The consequences of squirrel caching decisions: The spatial distribution of oaks and the structure of oak forests. International Theriological Conference VII,  Accapulco, Mexico.  9 September 1997.   (invited poster in the Symposium in honor of the 25th anniversary of the publication of R. H. MacArthur’s Geographical Ecology.

Awards:

National Science Foundation, Major Instrumentation Program. $273,951.
Craig Kinsley, Massimo Bardi, Cindy Bkack, John Warrick, Gary Radice, Peter Smallwood, Kelly Lambert, and Luciana Felicio. Title:  "MRI:  Acquisition of Apparatus to Support Multi-User Basic Research and Instruction in the Neurosciences."

Teagle Foundation Grants in Higher Education; Big Questions Working Groups.
Sidney Watts, Ori Belkind, Scott Davis, Douglass Hicks, Ray Hillard, David Leary, and Peter Smallwood. Title: “The Problem of Belief and Doubt in Liberal Education.”

NSF -special interdisiplinary competition for Collaborative Research at Undergraduate Institutions (C-RUI). Title: A multi-disciplinary approach to the ecological and evolutionary interactions between food-hoarding animals and the oaks. PI's M. A. Steele (Wilkes
University), P. D. Smallwood (University of Richmond), J.E. Carlson (Pennsylvania State University), and W.B. Terzaghi (Wilkes University)