Dr. Peter Smallwood

Associate Professor of Biology
Coordinator, Environmental Studies Program


Dr. Smallwood is interested in science policy and science diplomacy.  He has worked as a science advisor to individual US Senators and the Senate Homeland Security/Government Affairs committee.  He was the director of the Wildlife Conservation Society’s program in Afghanistan (2008—2009), and ran the US State Department’s program for Iraqi scientists (2004-05).  He teaches conservation biology, science policy, and ecology.

Grants and Fellowships

USAID Higher Education for Development/Initiative for Conservation in the Amazon and Andes: Building Conservation Capacity for a Changing Amazonia. D. Salisbury, M. Finley-Brook, R. de Sá, P.D. Smallwood, & Peruvian Consortium. Submitted June 28, 2012. Final Agreement signed 12/20/2012. US $750,000 over 2.5 years, to June 30, 2015.

Mellon Foundation, Support Course Development for Cross-School Seminars. Civilians Who Serve: The Privatization of Warfare. Smallwood, P.D. (Biology), Jones, J.P (Law), and Browder, L (American Studies) Funded, 8 December 2011 for Summer 2012: $4500.

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.
Sydney Watts, Ori Belkind, Scott Davis, Douglas 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. PIs: M.A. Steele (Wilkes University), P.D. Smallwood (University of Richmond), J.E. Carlson (Pennsylvania State University), and W.B. Terzaghi (Wilkes University)


Smallwood, P.D. 2013. Current Efforts in Cooperative Scientific Engagement and Mechanisms for Sustainability of the Programs and Relationships. Presentation to invitation-only conference,
“International Biosecurity: Engagement between American and MENA Scientists” Hosted by the American Association of the Advancement of Science and the Morrocan Biosafety Assocation. Casablanca, Morocco. 18-19 May, 2013.

Smallwood, P.D. 2012. Protecting Natural Areas in Afghanistan – Conservation in a Conflict Zone. Plenary Presentation to the Annual meetings of the Natural Areas Association, 9-12 October 2012.

Johnson, MF, N Kanderian, CC Shank, H Rahmani, D Lawson and PD Smallwood. 2012. Setting priorities for protected area planning in a conflict zone: Afghanistan’s National Protected Area System Plan. Contributed paper to the Annual Meetings of the Ecological Society of America. Portland, Oregon, USA. 5-10 August, 2012.

Smallwood, P.D. 2011. Challenges and opportunities for global biosurveillance from conservation programs. Presentation to the 2nd Biothreat Nonproliferation Conference: Global Biosurveillance. 19-21 January 2011, Santa Fe, New Mexico. Sponsored by DoE, DoD,
and DoS.

Smallwood, P.D. Warfare Ecology: Lessons from Afghanistan. Annual meeting of the Ecological Society of America. Austin, TX, USA, 1-6 August, 2011. (Invited symposium

Smallwood, P.D., M. Johnson, D. Salisbury, and S. Nash. Educating Undergraduates and addressing ecological problems at the same time: two case studies. Annual meeting of the Ecological Society of America. Pittsburgh, USA, 1-6 August, 2010 (contributed paper).

Smallwood, P.D., D. Lawson, and P. Zahler. Conserving Wildlife and Wild lands in conflict zones: Afghanistan. Annual meeting of the Ecological Society of America. Pittsburgh, USA, 1-6 August, 2010 (Invited symposium speaker).

Smallwod, P.D., D. Lawson, and P. Zahler. Afghanistan: Conservation in a country in conflict. Contributed paper in session #5: Community-based Conservation. Annual meeting, International Congress for Conservation Biology, Edmonton, Alberta, Canada. 3-7 July 2010.

Participant in AAAS program, International Engagement: Responsible Bioscience for a Safe and Secure Society. 3-6 October 2010, Amman, Jordan

Smallwood, P. D. Strategies for Conservation in high-profile conflict zones. Contributed paper in session “Conservation Strategies.” Annual meetings of the Association for Environmental Studies and Sciences, Madison, Wisconsin, USA. 8-11 October, 2009.

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. 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.



Steele, M., T. Contreras, L. Hadj-Chikh, S. Agosta, P.D. Smallwood, and C. Tomlinson. 2014. Do scatter-hoarders trade off increased predation risks for lower rates of cache pilferage? Behavioral Ecology 25:206-215.

Mitchell, J.C., and P.D. Smallwood. 2013. Field Notes: Lampropeltis getula (Eastern Kingsnake) and Chelydra serpentina (Snapping Turtle). Catesbeiana. 33: 25

Johnson, M.F., Kandarian, N, Shank, C.C., Rahmani, H., Lawson, D., and Smallwood, P.D. 2012. Setting priorities for protected area planning in a conflict zone – Afghanistan’s National Protected Area System Plan. Biological Conservation 148:146-155.

Smallwood, P.D., Shank, C, Dehgan, A. and Zahler, P. 2011. Wildlife Conservation in Afghanistan. BioScience 61:506-511.

Smallwood, P.D. and Liimaitenen, W.T. 2011. Safeguarding WMD Expertise: Lessons Learned From Iraq. Arms Control Today 41: 8-14.

Ostrowski, S., Zahler, P., Dehgan, A., Stevens, K., Karlstetter, M., and Smallwood, P.D. 2009. Asiatic Black Bear still survives in Nuristan, Afghanistan. International Bear News, 18:14-15.

Steele, M.A., Halkin, S.L., Smallwood, P.D., McKenna, T.J., Mitsopoulos, K, and Beam, M. (2007). Cache protection strategies of a scatter-hoarding rodent: do tree squirrels engage in behavioural deception?  Animal Behaviour: 75:705-714.

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.

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.

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.


Smallwood, P.D. 2013. The Risks of Greening in the Red Zone: Creating Afghanistan's First National Park in the Midst of Conflict. Chapter 21 in (Tidball and Krasny, Editors), Greening in
the Red Zone: Disaster, Resilience and Community Greening. Springer Press.

Smallwood, P.D. 2011. The Risks of Greening in the Red Zone: Creating Afghanistan's First National Park in the Midst of Conflict. Chapter 21 in (Tidball and Krasny, Editors), Greening in the Red Zone: Disaster, Resilience and Community Greening. Springer Press (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, 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}.

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).


Ph.D., University of Arizona

M.S., University of North Carolina

B.S., Ohio State University

Contact Information

A-110 Gottwald Center for the Sciences
(804) 289-8803
(804) 289-8233 (Fax)

Areas of Expertise

Science Policy
Conservation Biology
Environmental Studies