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Education, Research And Technical Assistance For Managing Our Natural Resources
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Dr. W. David Walter

Research Publications

  • Miller, W.L., J. Edson, P. Pietrandrea, C. Miller-Butterworth, and W. David Walter. Identification and evaluation of a core microsatellite panel for use in white-tailed deer (Odocoileus virginianus). BMC Genetics 20:49.
  • Schneider, A.L., A.T. Gilbert, W.D. Walter, G.S. Vandeberg, J.R. Boulanger. Spatial Ecology of Urban Striped Skunks (Mephitis mephitis) in the Northern Great Plains: A Framework for Future Oral Rabies Vaccination Programs. Urban Ecosystems 22:539-552. Abstract | 
  • Miller, W.L. and W. D. Walter. Spatial heterogeneity of prion gene polymorphisms in an area recently infected by chronic wasting disease. Prion 13: 65-76. Abstract | 
  • Amor, J.M., R. Newman, W.F. Jensen, B.C. Rundquist, W.D. Walter, and J.R. Boulanger. Seasonal home ranges and habitat selection of three elk (Cervus elaphus) herds in North Dakota. PlosONE 14(2): e0211650.
  • Walter, W.D., T.S. Evans, D. Stainbrook, B.D. Wallingford, C.S. Rosenberry, and D.R. Diefenbach. 2018. Heterogeneity of a landscape influences size of home range in a North American cervid. Scientific Reports 8:14667. DOI:10.1038/s41598-018-32937-7. Abstract |  Download  |  Publisher Website | 
  • Williamson, L. T., W. David Walter, S. R. Klinger, and D. R. Diefenbach. 2018. Incorporating detection probability to estimate pheasant density. Journal of Wildlife Management. 82(8):1680–1688; DOI: 10.1002/jwmg.21545 Download  | 
  • Carrollo, E.M., H.E. Johnson, J.W. Fischer, M. Hammond, P.D. Dorsey, C.W. Anderson, K.C. VerCauteren, and W. David Walter. Influence of precipitation and crop germination on resource selection by mule deer (Odocoileus hemionus) in southwest Colorado. Scientific Reports Abstract | 
  • Miller, W.L. and W.D. Walter. 2017. CWDPRNP: a tool for cervid prion sequence analysis in program R. Bioinformatics. 33(19):3096–3097. Abstract | 
  • King, D.T., J.W. Fischer, B. Strickland, W.D. Walter, F.L. Cunningham, and G. Wang. 2016. Winter and summer home ranges of American white pelicans (Pelecanus erythrorhynchos) captured at loafing sites in the southeastern United States. Waterbirds 39(3):287–294.
  • Fischer, J.W., C.R. Blass, W.D. Walter , C.W. Anderson, M.J. Lavelle, W.H. Hall, and K.C. VerCauteren. 2016. Evaluating a strategy to deliver vaccine to white-tailed deer at a landscape level. Wildlife Society Bulletin 40(2):394–399. Abstract | 
  • Fischer, J.W., D. McMurtry, C.R. Blass, W.D. Walter , J. Beringer, and K.C. VerCauteren. 2016. Effects of simulated removal activities on movements and space use of feral swine. European Journal of Wildlife Research 62(3):285–292. Abstract | 
  • Haley, N.J., C. Siepker, L.L. Hoon-Hanks, G. Mitchell, W.D. Walter, M. Manca, R.J. Monello, J.G. Powers, M.A. Wild, E.A. Hoover, B. Caughey, and J. Richt. 2016. Seeded amplification of chronic wasting disease prions in nasal brushings and recto-anal mucosal associated lymphoid tissues from elk by real time quaking-induced conversion. Journal of Clinical Microbiology 54(4): 1117-1126.
  • Haley, N.J., C. Siepker, W.D. Walter, B.V. Thomsen, J.J. Greenlee, A.D. Lehmkuhl, and J.A. Richt. 2016. Antemortem detection of chronic wasting disease prions in nasal brush collections and rectal biopsies from white-tailed deer by real time quaking-induced conversion. Journal of Clinical Microbiology 54(4): 1108-1116.
  • Evans, T.S., M. Kirchgessner, B. Eyler, C.W. Ryan, and W.D. Walter. 2016. Habitat influences distribution of of chronic wasting disease in white-tailed deer. Journal of Wildlife Management Abstract | 
  • Walter, W.D., D.P. Onorato, and J.W. Fischer. 2015. Is there a single best estimator? Selection of home range estimators using area- under- the-curve. Movement Ecology Abstract |  Download  | 
  • Evans, T.S., K.L. Schuler, W.D. Walter. 2014 Surveillance and monitoring of white-tailed deer for chronic wasting disease in the northeastern United States. Journal of Fish and Wildlife Management 5(2):387-393.
  • Walter, W.D. and J.W. Fischer. Manual of Applied Spatial Ecology. Online: <> Abstract |  Publisher Website | 
  • Walter, WD,, Kurle, CM, Hopkins III, JB. 2014. Applications of stable isotope analysis in mammalian ecology. Isotopes in Environmental and Health Studies 50(3):287-290.
    Abstract | 
  • Walter, W.D. 2014. Use of stable isotopes to identify dietary differences and subpopulations of a free-ranging generalist herbivore. Isotopes in Environmental and Health Studies 50(3):399-413.
  • Johnson, H.E., J.W. Fischer, M Hammond, P. Dorsey, W. D. Walter, C.W. Anderson, and K.C. VerCauteren. 2014. Evaluation of techniques to reduce deer and elk damage to agricultural crops. Wildlife Society Bulletin 38(2):358–365. Abstract | 
  • Walter, W.D., R. Smith, M. Vanderklok, and K.C. VerCauteren. 2014. Linking bovine tuberculosis on cattle farms to white-tailed deer and environmental variables using Bayesian hierarchical analysis. Plos ONE 9(3):e90925. Abstract | 
  • Berentsen, A.R., M.R. Dunbar, C.E. Fitzpatrick, W.D. Walter. 2013. Spatial ecology of urban raccoons in Northeastern Ohio: implications for oral rabies vaccination. Prairie Naturalist 45(1): 39-45. Abstract | 
  • Fischer, J.W., W.D. Walter, M.A. Avery. 2013. Brownian bridge movement models to characterize home ranges of avian species. The Condor 115(2): 298-305. Abstract | 
  • Walter, W.D., J.W. Fischer, T.J. Zimmerman, S.E. Hygnstrom, J.A. Jenks, K.C. VerCauteren. 2013. Topographic home range of large mammals: is planimetric home range still a viable method? Prairie Naturalist 45(1): 21-27. Abstract | 
  • Walter,W.D. ,Fischer,J.W. Anderson,C.W., Marks,D.R., Deliberto,T., Robbe-Austerman,S., VerCauteren,K.C. 2013. Surveillance and movements of Virginia opossum (Didelphis virginiana) in the bovine tuberculosis region of Michigan. Epidemiology and Infection, Special Issue on TB, 141(7): 1498-1508. Abstract |  Publisher Website | 
  • Walter, W.D., C. W. Anderson, R. Smith, M. Vanderklok, J. Averill, and K.C. VerCauteren. 2012. On-farm mitigation of transmission of tuberculosis from white-tailed deer to cattle: literature review and recommendations. Veterinary Medicine International Article ID 616318. Download  |  Publisher Website | 
  • Walter, W.D., C.W. Anderson, and K.C. VerCauteren. 2012. Evaluation of remote delivery of passive infrared transponder (PIT) technology to mark large mammals. Plos ONE 7(9): e44838 Download  |  Publisher Website | 
  • Henningsen, J.C., A.L. Williams, C.M. Tate, S.A. Kilpatrick, and W.D. Walter. 2012. Distribution and prevalence of Elaeophora schneideri in moose in Wyoming. Alces 48:35-44
  • Walter, W.D., J.W. Fischer, M.A. Avery J.S. Humphrey, T.S. Daughtery, M.P. Milleson, and E.A. Tillman. 2012. Using three-dimensional flight patterns at airfields to identify hotspots for avian-aircraft collisions. Applied Geography 35:53-59. Download  |  Publisher Website | 
  • Walter, W.D., J.W. Fischer, S. Baruch-Mordo, and K.C. VerCauteren. 2011. What is the proper method to delineate home range of an animal using today’s advanced GPS telemetry systems: the initial step. In: Modern telemetry, Intech, Open Access Publisher. Download  |  Publisher Website | 
  • VerCauteren, K.C., C.W. Anderson, T.R. Van Deelen, D. Drake, W.D. Walter, S.M. Vantassel, and S.E. Hygnstrom. 2011. Regulated commercial harvest to manage overabundant white-tailed deer: an idea to consider? Wildlife Society Bulletin 35(3):185–194. Publisher Website | 
  • Avery, M.L., J.S. Humphrey, T.S. Daughtery, J.W. Fischer, M.P. Milleson, E.A. Tillman, W.E. Bruce, and W. D. Walter. 2011. Vulture flight behavior and implications to aircraft safety. Journal of Wildlife Management 75(7): 1581–1587. Publisher Website | 
  • Walter, W.D., J. Beringer, L.P. Hansen, J.W. Fischer, J.J. Millspaugh, and K.C. VerCauteren. 2011. Factors affecting space use overlap by white-tailed deer in an urban landscape. International Journal of Geographical Information Science 25(3):379–392. Publisher Website | 
  • Walter, W.D., D.M. Baasch, S.E. Hygnstrom, B.D. Trindle, A.J. Tyre, J.J. Millspaugh, C.J. Frost, J.R. Boner, and K.C. VerCauteren. 2011. Space use of sympatric deer in a riparian ecosystem in an area chronic wasting disease is endemic. Wildlife Biology 17(2):191–209. Publisher Website | 
  • Walter, W.D., D.P. Walsh, M.L. Farnsworth, D.L. Winkelman, and M. W. Miller. 2011. Soil clay content underlies prion infection odds. Nature Communications 2:200.
  • Walter, W.D., M.J. Lavelle, J.W. Fischer, T..J. Johnson, S.E. Hygnstrom, and K.C. VerCauteren. 2010. Management of damage by elk (Cervus elaphus) in North America: a review. Wildlife Research 37(8):630–646. Publisher Website | 
  • Davis, C. A., D. M. Leslie, Jr., W. D. Walter, and A. E. Graber. 2010. Mountain biking trail use affects reproductive success of nesting gold-cheeked warblers. Wilson Journal of Ornithology 122:465-474.
  • Walter, W. D., D. M. Leslie, Jr., E. C. Hellgren, and D. M. Engle. 2010. Identification of subpopulations of North American elk (Cervus elaphus) using multiple lines of evidence: habitat use, dietary choice, and fecal stable isotopes. Ecological Research 25:789-800.
  • Walter, W. D., T. J. Zimmerman, D. M. Leslie, Jr., and J. A. Jenks. 2009. Dietary response of sympatric deer to fire using stable isotope analysis of liver tissue. Wildlife Biology in Practice 5:128-135.
  • Spraker, T. R., K. C. VerCauteren, T. L. Gidlewski, R. D. Munger, W. D. Walter, A. Balachandran, and K. I. O’Rourke. 2009. Impact of age and sex of Rocky Mountain elk (Cervus elaphus nelsoni) on follicle counts from rectal mucosal biopsies for preclinical detection of chronic wasting disease. Journal of Veterinary Diagnostic Investigation. 21(6): 868−870.
  • Walter, W.D., and D.M. Leslie, Jr. 2009. Stable isotope ratio analysis to differentiate temporal diets of a free-ranging herbivore. Rapid Communications in Mass Spectrometry. 23(14): 2190−2194
  • Walter, W.D., K.C. VerCauteren, H. Campa, III, W.R. Clark, J.W. Fischer, S.E. Hygnstrom, N.E. Mathews, C.K. Nielsen, E.M. Schauber, T.R. Van Deelen, and S.R. Winterstein. 2009. Regional assessment on influence of landscape configuration and connectivity on range size of white-tailed deer. Landscape Ecology. 24(10): 1405−1420
  • Ward, A.I., K.C. VerCauteren, W.D. Walter, E. Gilot-Fromont, S. Rossi, G. Edwards-Jones, M. Lambert, M.R. Hutchings, and R.J. Delahay. 2009. Options for the control of disease 3: targeting the environment. Pages 147−168 in R.J. Delahay, M.R. Hutchings, and G.C. Smith (eds) Management of disease in wild mammals. Springer. Publisher Website | 
  • Walter, W. D., R. L. Bryant, and D. M. Leslie, Jr. 2005. Unusual documentation of elk behaviors using automated cameras. Proceedings of the Oklahoma Academy of Science 85:81-83.
  • Walter, W. D., D. M. Leslie, Jr., J. H. Herner-Thogmartin, K. G. Smith, and M. E. Cartwright. 2005. Efficacy of immobilizing free-ranging elk with Telazol and xylazine hydrochloride using tranmitter-equipped darts. Journal of Wildlife Diseases 41:395-400.
  • Walter, W. D., H. J. Kilpatrick, and M. A. Gregonis. 2003. Does immunocontraception improve condition of free-ranging white-tailed deer? Journal of Wildlife Management 67:762-766.
  • Walter, W. D., and D. M. Leslie, Jr. 2002. Harvest strategies and numbers of elk (Cervus elaphus) in Oklahoma, 1987-2201. Proceedings of the Oklahoma Academy of Science 82:89-94.
  • Walter, W.D., P.J. Pekins, A.T. Rutberg, and H.J. Kilpatrick. Evaluation of immunocontraceptive adjuvants, titers, and fecal pregnancy indicators in free-ranging white-tailed deer. Wildlife Society Bulletin 30:908-914.
  • Walter, W. D., P. J. Pekins, A. T. Rutberg, and H. J. Kilpatrick. 2002. Evaluation of immunocontraception in a free-ranging suburban white-tailed deer herd. Wildlife Society Bulletin 30:186-192.
  • Kilpatrick, H.J., and W.D. Walter. A controlled archery deer hunt in a residential community: cost, effectiveness, and deer recovery rates. Wildlife Society Bulletin 27:115–123.
  • Kilpatrick, H.J., and W.D. Walter. 1997. Urban deer management: a community vote. Wildlife Society Bulletin 25:388–390.

Current Staff

Federal Staff: 3

Masters Students: 5

Phd Students: 2

Post Docs: 2

University Staff: 3

5 Year Summary

Students graduated: 20

Scientific Publications: 82

Presentations: 116


Contact Us

Pennsylvania Cooperative Fish and Wildlife Research Unit University Park, PA 16802 Phone: (814) 865 - 4511 Fax: (814) 863 - 4710

Unit Leader

Duane Diefenbach
Duane R. Diefenbach

Upon completion of my Ph.D. I worked for the Pennsylvania Game Commission as their biometrician for 7 years. There I began my current research involving grassland sparrows and black bears. Since 1999 I have been a research scientist with the Pennsylvania Unit and enjoy the challenges in integrating research to improve management decisions.


  1. Penn State University Campus Maps

Pennsylvania Cooperative Fish and Wildlife Research Unit Cooperators

  1. Pennsylvania Fish and Boat Commission
  2. Pennsylvania Game Commission
  3. Pennsylvania State University
  4. U.S. Fish and Wildlife Service
  5. U.S. Geological Survey
  6. Wildlife Management Institute