Cipollini Lab Home Page

 

Dr. Don Cipollini

Professor, Department of Biological Sciences

Director, Environmental Sciences PhD Program

208 Biological Sciences Building
Wright State University
3640 Colonel Glenn Hwy.
Dayton, OH 45435

Email: don.cipollini@wright.edu
Phone (937) 775-3805

FAX (937) 775-3320   

General Research Interests

Work in my laboratory generally focuses on the chemical ecology of plant responses to the environment, with a focus on induced plant responses to herbivores and pathogens and the chemical ecology of invasive plants, insects, and microbes.  One important theme is the influence of environmental factors on the expression of phenotypic plasticity and the outcomes of interactions of plants with their biotic environment.  Within this context, my interests range from highly mechanistic explanations of induced plant responses and their impacts on herbivores and plant fitness, to evolution of plant defenses, to community level impacts.  The current project areas described below represent the major thrusts of our research program, although specific directions are often driven by student interests and the availability of funding.  In particular, you will see that we have adopted Arabidopsis thaliana as an ecological model and have extended our understanding of plant resistance to understand invasiveness and impacts of Alliaria petiolata, an important invasive plant.  Our studies have extended to the expression and role of constitutive and inducible defenses in woody plant ecology, including that of the invasive shrub, Lonicera maackii.  We have worked experimentally with bacteria, fungi, insects, and plants, and use a variety of techniques including basic experimental greenhouse and field ecology techniques, microscopy, molecular analyses, and analytical chemistry techniques such as HPLC.

 

Current Projects

Chemical ecology of the invasive plants, garlic mustard
(Alliaria petiolata) and Amur honeysuckle (Lonicera maackii)


Garlic mustard is a Eurasian biennial herb that was first noted in North America in the mid 1800’s.  Since its introduction, this plant has spread throughout North America where it is considered an invasive threat to forest understory flora and fauna.  There are numerous hypotheses about the success of invasive plants in novel environments, including the Natural Enemies hypothesis, the Evolution of Increased Competitive Ability hypothesis, and the Novel Weapons hypothesis.  Garlic mustard provides an interesting model with which to test some of them.  Our work on garlic mustard has been funded by the Wright State University Research Council, the Ohio Board of Regents, and the Ohio Plant Biotechnology Consortium.  On a related topic, we have also extended our interest in chemical ecology of invasive plants to examine allelopathy, herbivore resistance, plant-soil feedbacks and other mechanisms of invasiveness in Amur honeysuckle, Lonicera maackii, an Asian shrub that is invasive throughout eastern North America.

 

Current projects include:

●Experimental tests of purported invasive mechanisms

●Sources of variation and novelty in the expression of chemical defenses

●Growth and defense responses to insect damage, powdery mildew infection, and their interaction

●Identity and impacts of allelochemicals on plants and their associated mycorrhizae

●Assessment of impacts on plant and animal species and genetic diversity of invaded communities

●Chemical defense responses to climate change

Costs and benefits of induced plant responses to insects
and pathogens in Arabidopsis thaliana

Interest in this system lies primarily in costs, benefits, and tradeoffs of plant responses mediated by salicylic acid and jasmonic acid, which we commonly use in conjunction with genetic mutants to manipulate and study plant defense responses at the biochemical, physiological, and ecological level.  Interest also lies in integrated plant responses to herbivores, light, competitors, and other ecological stimuli.  We have also commonly used Arabidopsis as a target plant in allelopathy studies.  Various aspects of our work on Arabidopsis have been funded by USDA, the Ohio Plant Biotechnology Consortium, the Eppley Foundation for Research, and Sigma Xi.

Current projects include:

●Physiological and ecological costs of resistance

●Interactions between plastic responses to herbivores and competitors

●Functional ecology of methyl jasmonate

●Exploring the potential of Plant Growth Promoting Rhizobacteria to enhance growth and insect resistance

Chemical ecology of induced resistance in woody plants

Our interest in inducible chemical defenses has extended to woody plants, including Austrian Pine and Ash species, in collaboration with Dan Herms and Enrico Bonello, Ohio State University.  Focus has been on genetic and environmental influences on local and systemic defenses responses of both leaves and phloem tissues.  Work on the pine and ash projects has been funded by USDA, OARDC, the Ohio Plant Biotechnology Consortium, and the USDA-APHIS. 

 

Current Projects Include:

●Chemical ecology of Austrian Pine responses to insects and pathogens across a nutrient gradient

●Mechanisms and variation in ash resistance to Emerald Ash Borer

Conservation and restoration ecology

My wife, Kendra Cipollini, and I have collaborated on a number of things, including producing our sons, Emmett and Otto, and in her work on the federally endangered wetland plant, Scirpus ancistrochaetus.  Kendra formerly worked as a conservation scientist with the Nature Conservancy and is a now an assistant professor in Ecology at Wilmington College in Wilmington, OH.  Check out  Kendra's homepage .  Currently, we are exploring population status, genetic diversity and environmental influences on the growth of S. ancistrochaetus, work funded by the US Fish and Wildlife Service.  We are also investigating impacts and restoration methods of sites dominated by the invasive shrub, Lonicera maackii, Ranunculus ficaria, and other invaders. Work on the latter system has been supported by the Hamilton County Park District.

Current projects include:

●Impacts of deer browsing and method of control on restoration success in honeysuckle-remediated areas.

●Interactive effects of water level and nutrient availability on competitive associations of Scirpus ancistrochaetus and Scirpus cyperinus

●Population status and genetic diversity of Scirpus ancistrochaetus, a federally endangered wetland sedge
 

Curriculum Vitae and Publications

EDUCATION

Pennsylvania State University, 1997, Ph.D. Ecology, Advisor: Jack C. Schultz
Indiana University of Pennsylvania, 1993, M.S. Biology, Advisor: Sandra J. Newell
Indiana University of Pennsylvania, 1990, B.S. Biology

ADDITIONAL TRAINING

Tropical Field Ecology (OTS 94-1), Organization for Tropical Studies, Costa Rica, 1994

APPOINTMENTS

Professor, (2008-present), Dept. of Biological Sciences, Wright State University

Director (2009-present), Environmental Sciences PhD Program, Wright State University

Interim Director, (2007-2009) Environmental Sciences PhD Program, Wright State University

Associate Professor, (2004-2008), Dept. of Biological Sciences, Wright State University

Assistant Professor, (1999-2004), Dept. of Biological Sciences, Wright State University

Postdoctoral Research Associate, (1997-1999), University of Chicago

Graduate Teaching and Research Assistant, (1992-1997), Pennsylvania State University,

Graduate Teaching and Research Assistant, (1990-1992), Indiana University of Pennsylvania

PUBLICATIONS

           60.  Cipollini, D., Wang, Q., Whitehill, J.G.A., Powell, J., Bonello, P, and Herms, D. 2011.

Distinguishing defensive characteristics in the phloem of ash species resistant and susceptible to the Emerald Ash Borer.  Journal of Chemical Ecology (in press)

 

59.  Enright, S., and Cipollini, D.  2011.  Overlapping defense responses to water limitation and

Pathogen attack and their consequences for resistance to powdery mildew disease in garlic mustard, Alliaria petiolata.  Chemoecology  21: 89-98.  DOI: 10.1007/s00049-011-0072-8

 

58.  Hillstrom, C, and Cipollini, D. 2011.  Variation in phenotypic plasticity in native and invasive

 populations of Alliaria petiolata.  International Journal of Plant Sciences.  (in press)

 

57.  Koch, A.M., Antunes, P.M., Barto, E.K., Cipollini, D., Mummey, D.L., and Klironomos, J.N.

2011.  The effects of arbuscular mycorrhizal (AM) fungal and garlic mustard introductions on native AM fungal diversity.  Biological Invasions DOI: 10.1007/s10530-010-9920-7

 

56.  Barto, E.K., Antunes, P.M., Stinson, K., Koch, A.M., Klironomos, J.N., Cipollini, D. 2011.

Differences in arbuscular mycorrhizal fungal communities associated with sugar maple seedlings in and outside of invaded garlic mustard forest patches.  Biological Invasions (in press)

 

55.  Ciola, V., and Cipollini, D. 2011.  Distribution and host range of a powdery mildew fungus

attacking garlic mustard in southwestern Ohio.  American Midland Naturalist (in press)

 

54.  Cipollini, K. and Cipollini, D.  2011.  Population status and threats to the federally endangered

Scirpus ancistrochaetus: implications for management.  Northeastern Naturalist (in press)

 
               53.  Barto, E.K., Powell, J., and Cipollini, D.  2010.  How novel are the chemical weapons of garlic 
mustard in North American forest understories?  Biological Invasions 12: 3465-3471, doi: 10.1007/s10530-010-9744-5
 
               52.  Barto, E.K., Friese, C., and Cipollini, D.  2010.  Arbuscular mycorrhizal fungi protect a native
plant from allelopathic effects of an invader.  Journal of Chemical Ecology 36: 351-360.
 
               51.  Cipollini, D., and Heil, M.  2010.  Costs and benefits of induced resistance to pathogens and
herbivores in plants.  CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources doi: 10.1079/PAVSNNR20105005
               

            50.  Wallis, C., Eyles, A., Chorbadjian R.A., Riedl K., Schwartz S., Hansen R., Cipollini, D., Herms,

            D.A., and Bonello P.  2010.  Differential effects of nutrient availability on the secondary

metabolism of Austrian pine (Pinus nigra) phloem and resistance to Diplodia pinea.  Forest Pathology doi: 10.1111/j.1439-0329.2009.00636.x

 
49.  Cipollini, D. 2010.  Constitutive expression of methyl jasmonate-inducible responses delays reproduction
               and constrains fitness responses to nutrients in Arabidopsis thaliana.  Evolutionary Ecology 24: 59-68.
 
               48.  Barto, E.K., and D. Cipollini.  2009.  Garlic mustard (Alliaria petiolata) removal method affects 
               native establishment.   Invasive Plant Science and Management 2:230-236.
 
               47.  Barto, E.K., and D. Cipollini.  2009.  Density dependent phytotoxicity of Impatiens pallida
                plants exposed to extracts of Alliaria petiolata.  Journal of Chemical Ecology 35: 495-
               504.
 
               46.  Cipollini, D., and S. Enright.  2009.  A powdery mildew fungus levels the playing field for
           invasive garlic mustard (Alliaria petiolata) and a North American native plant.  Invasive Plant Science and
           Management 2:253-259
 
45.  Barto, E.K., and D. Cipollini.  2009.  Field concentrations and half-lives of Alliaria petiolata 
secondary metabolites. Chemosphere 76: 71-75
 
44.  Cipollini, K.A., E. Ames, and D. Cipollini.  2009.  Amur Honeysuckle (Lonicera maackii) 
management method impacts restoration of understory plants in the presence of White-Tailed Deer 
(Odocoileus virginiana).  Invasive Plant Science and Management 2:45-54

 

43.  Barto, K., S. Enright, A. Eyles, C. Wallis, R. Chorbadjian, R. Hansen, D.A. Herms, P. Bonello, and D. Cipollini.  2008.  Effects of fertilization and fungal and insect attack on systemic protein defenses of Austrian pine. Journal of Chemical Ecology 34: 1392-1400

 

42. Stireman, J.O., and D. Cipollini.  2008.  Stealth tactics of galling parasites and their potential indirect effects.  New Phytologist, 178: 462-465

 

41.  Adams, J.M., Fang, W., Callaway, R.M., Cipollini, D., Newell, E., and Transatlantic Acer platanoides Invasion Network.  2009.  A cross-continental test of the Enemy Release Hypothesis: leaf herbivory on Acer platanoides (L.) is three times higher in North America than its native Europe.  Biological Invasions, in press

 

40.  Cipollini, D. and M. Dorning.  2008.  Direct and indirect effects of conditioned soils and tissue extracts of the invasive shrub, Lonicera maackii, on target plant performance. Castanea,

 

39.  Cipollini, D., R. Stevenson, S. Enright, A. Eyles, P. Bonello.  2008.  Phenolic metabolites in leaves of the invasive shrub, Lonicera maackii, and their potential phytotoxic and anti herbivore effects.  Journal of Chemical Ecology  34: 144-152

 

38. Cipollini, K.A., G.Y. McClain, and D. Cipollini.  2008.  Separating effects of allelopathy and shading by Alliaria petiolata and Lonicera maackii on growth, reproduction and survival of Impatiens capensis.  American Midland Naturalist, 160: 117-128

 

37.  Cipollini, D., R. Stevenson, K. Cipollini.  2008.  Contrasting direct and indirect effects of

allelochemicals from two invasive plants on the performance of a non-mycorrhizal plant.  International Journal of Plant Sciences 169: 371-375

 

36.  Wallis, C., A. Eyles, R. Chorbadjian, B. McSpadden-Gardener, R. Hansen, D. Cipollini, D. A. Herms, and P. Bonello.  2008.  Systemic induction of phloem secondary metabolism and induction of resistance to a canker pathogen in Austrian pine.  New Phytologist, 177: 767-778.

 

35.  Callaway, R.M., D. Cipollini, K. Barto, G. C. Thelen, S. G. Hallett, D. Prati, K. Stinson, J. Klironomos.  2008.  Novel weapons: invasive plant suppresses fungal mutualisms in America but not in its native Europe.  Ecology, 89:1043-1055

 

34.  Eyles, A., D.A. Herms, W. Jones, K. Riedl, D. Cipollini, S. Schwartz, K. Chan, P. Bonello. 2007.  Comparative phloem chemistry of Manchurian (Fraxinus mandshurica) and two North American ash species (F. americana and F. pennsylvanica).  Journal of Chemical Ecology 33: 1430-1448

 

33.  Enright, S. and D. Cipollini.  2007.  Infection by powdery mildew Erysiphe cruciferarum

(Erysiphaceae) strongly affects growth and fitness of garlic mustard Alliaria petiolata (Brassicaceae).  American Journal of Botany, 94: 1813-1820

 

32.  Eyles, A., R. Chorbadjian, C. Wallis, R. Hansen, D. Cipollini, D. Herms, P. Bonello.  2007.  Cross-induction of systemic induced resistance between an insect and a fungal pathogen in Austrian pine over a fertility gradient.  Oecologia, 153: 365-374

 

31.  Cipollini, D. and B. Gruner.  2007.  Cyanide in the chemical arsenal of garlic mustard, Alliaria petiolata.  Journal of Chemical Ecology 33: 85-94 pdf

 

30.  Cipollini, D.  2007.  Consequences of the overproduction of methyl jasmonate on seed production, tolerance to defoliation, and competitive effect and response of Arabidopsis thaliana.  New Phytologist 173: 146-153  pdf

 

29. Dorning, M, and D. Cipollini.  2006.  Leaf and root extracts of the invasive shrub Lonicera maackii inhibit seed germination of three herbs with no autotoxic effects.  Plant Ecology 184: 287-296 pdf

 

28.  Cipollini, D.  2005.  Interactive effects of lateral shading and jasmonic acid on morphology, phenology, seed production, and chemical defense responses in Arabidopsis thaliana.  International Journal of Plant Sciences 166: 955-959 pdf

 

27.  Barto, K, and D.F. Cipollini.  2005.  Testing predictions of the Optimal Defense Theory and Growth Differentiation Balance Hypothesis in Arabidopsis thalianaOecologia  146: 169-178  doi: 10.1007/s00442-005-0207-0 pdf

26.  Cipollini, D., J. Mbagwu, K. Barto, C. Hillstrom, and S. Enright.  2005.  Expression of constitutive and inducible chemical defenses in native and invasive populations of garlic mustard (Alliaria petiolata).  Journal of Chemical Ecology 31: 1243-1255 pdf

25.  R.O. Musser, D. F. Cipollini, S.M. Hum-Musser, S.A. Williams, J.K. Brown, and G.W. Felton.  2005.  Evidence that the caterpillar salivary enzyme glucose oxidase provides herbivore offense in Solanaceous plants.  Archives of Insect Biochemistry and Physiology 58: 128-137.

24.  Cipollini, D.F., S. Enright, B. Traw, and J. Bergelson.  2004.  Salicylic acid inhibits jasmonic acid-induced resistance of Arabidopsis thaliana to Spodoptera exiguaMolecular Ecology 13: 1643-1653 pdf

23.  Cipollini, D.F.  2004.  Stretching the limits of plasticity: can a plant defend itself from both competitors and herbivores?  Ecology 85: 28-37 (Special feature on Phytohormonal Ecology) pdf

22.  Traw, M.B., Kim, J., Enright, S., Cipollini, D.F., and J. Bergelson.  2003.  Negative cross talk between salicylate and jasmonate-mediated pathways in the Wassilewskija ecotype of Arabidopsis thalianaMolecular Ecology 12: 1125-1135 pdf

21.  Cipollini, D.F., J.W. Busch, K. Stowe, E. Simms, and J. Bergelson.  2003.  Genetic variation and relationships of constitutive and herbivore-induced glucosinolates, trypsin inhibitors, and herbivore resistance in Brassica rapa. Journal of Chemical Ecology  29: 285-302. pdf

20. Cipollini, D.F., C. B. Purrington, and J. Bergelson.  2003.  Costs of induced responses.  Basic and Applied Ecology 4: 79-89 pdf

19. Cipollini, D.F.  2002.  Variation in the expression of chemical defenses in Alliaria petiolata in the field and common garden.  American Journal of Botany  89: 1422-1430 pdf

18. Cipollini, M., E. Paulk, and D. Cipollini.  2002.  Effect of nitrogen and water treatment on leaf chemistry in horsenettle (Solanum carolinense) and relationship to herbivory by flea beetles (Epitrix spp.) and tobacco hornworm (Manduca sextaJournal of Chemical Ecology 28: 2377-2398. pdf

17. Cipollini, D.F.  2002.  Does competition magnify the fitness costs of induced responses in Arabidopsis thalianaA manipulative approach.  Oecologia 131: 514-520. pdf

16. Cipollini, D.F., and J. Bergelson.  2002.  Interspecific competition affects growth and herbivore damage of Brassica napus L. in the field.  Plant Ecology  162: 227-231

15. Cipollini, D.F., and M. Sipe. 2001  Jasmonic acid treatment and mammalian herbivory differentially affect chemical defenses and growth of Brassica kaberChemoecology 11: 137-143.

14. Cipollini, D.F., and J. Bergelson.  2001.  Plant density and nutrient availability constrain the constitutive and wound-induced production of trypsin inhibitors in Brassica napus L.  Journal of Chemical Ecology 27: 593-610.  pdf

13. Redman, A, Cipollini, D.F., and J.C. Schultz.  2001.  Fitness costs of jasmonic acid-induced defense in tomato, Lycopersicon esculentumOecologia 126: 380-385. pdf

12. Cipollini, D.F., and J. Bergelson.  2000.  Environmental and developmental regulation of trypsin inhibitor activity in Brassica napus L.  Journal of Chemical Ecology 26: 1411-1422. pdf

11. Cipollini, D.F. and J.C. Schultz.  1999.  Exploring cost constraints on stem elongation in plants using phenotypic manipulation.  American Naturalist  153: 153: 236-242. pdf

10. Cipollini, D.F.  1999.  Costs to flowering of the production of a mechanically hardened phenotype in Brassica napus L.  International Journal of Plant Sciences  160: 735-741. pdf

9. Cipollini, D.F., and A.M. Redman.  1999.  Age-dependent effects of jasmonic acid treatment and wind exposure on foliar oxidase activity and insect resistance in tomato.  Journal of Chemical Ecology  25: 271-281 pdf

8. Lentz, K., and D.F. Cipollini.  1998.  Effect of light and simulated herbivory on growth of
endangered northeastern bulrush, Scirpus ancistrochaetus Schuyler.  Plant Ecology 139: 125-131.  pdf

7. Moran, P.J., and D.F. Cipollini.  1998.  Effect of fungal infection and mechanical stress on peroxidase activity and resistance to pests in cucumber.  Journal of Phytopathology 147: 313-316.

6. Cipollini, D.F.  1998.  The induction of soluble peroxidase activity in leaves of bean plants by wind induced mechanical perturbation.  American Journal of Botany  85: 1586-1591 pdf

5. Cipollini, D.F.  1998.  Induced defenses and phenotypic plasticity.  Trends in Ecology and Evolution  13: 200.

4. Cipollini, D.F.  1997.  Wind-induced mechanical stimulation increases pest resistance in common bean.  Oecologia 111: 84-90. pdf

3. Cipollini, D.F.  1997.  Gibberellic acid treatment reduces the tolerance of field-grown common bean to leaf removal.  Journal of Plant Growth Regulation  16: 123-127.

2. Cipollini, D.F. and J.C. Schultz. 1997.  Wind and trees.  Trends in Ecology and Evolution 12: 276-277.

1. Cipollini, D.F., S.J. Newell, and A.J. Nastase. 1994. The estimation of total carbohydrates in the nectar of the pitcher plant, Sarracenia purpurea L.  American Midland Naturalist 131: 374-377.

 

CURRENT LAB PEOPLE:

Graduate Students

 

Deah Lieurance (E.S. Ph.D. Program 2008-present):  Exploring the enemy release hypothesis in invasive Lonicera species.

 

Samantha (Sam) Davis (E.S. Ph.D. Program, 2010-present):  Direct and indirect impacts of garlic mustard on a rare woodland butterfly.

 

Andy Officer (M.S. Program, 2009-present):  Interactions between garlic mustard, and mutualistic and pathogenic fungi.

 

Kelly Schradin (M.S. Program, 2010-present):  The role of plant-soil feedback in the invasiveness of Lonicera maackii.

 

LAB ALUMNI:

Wang Qin (M.S. Biology 2010):  Variation in resistance of Asian and North American ash species to the Emerald Ash Borer.  Current: MBA program Wright State University

 

Kathryn Barto (M.S. Biology 2003):  Testing the optimal defense theory and growth differentiation balance hypothesis in Arabidopsis thaliana.
(Ph.D. Environmental Sciences 2008):  An assessment of the allelopathic potential of Alliaria petiolata.  Current: Post-Doc at Freie Universitat, Berlin

 

Eusondia Barto  Undergraduate honor’s thesis (2006):  Influence of wounding and emerald ash borer extracts on defense protein expression in white and green ash.  Current: PhD program in Microbiology, Ohio State University

 

Victoria Ciola (M.S. Program 2007-2009):  Distribution and host specificity of a powdery mildew infecting garlic mustard.  Science Editor, McGraw-Hill

 

Dan Romanek (M.S. Program 2008-2009):  Population and biogeographic variation in allelopathic effects of Lonicera maackii.  Chemical Remediation Industry

Stephanie Enright (M.S. Biology 2006)  Direct and indirect impacts of powdery mildew on Alliaria petiolata.  Current: Lab Coordinator, Wittenberg University

Carl-Johan Hillstrom (M.S. Biology 2005)  Plasticity in growth, antioxidant chemistry, and jasmonic acid induced defenses of Alliaria petiolata across nutrient and moisture gradients.  Current:  PhD program in Plant Physiology, University of Kentucky

Sara Weber (M.S. Biology 2005)  Impacts of Alliaria petiolata invasion on the genetic diversity of Impatiens capensis.   Current:  Environmental Specialist, Ashland Chemical

Monica Dorning  (M.S. Biology 2004) Impacts of honeysuckle on neighboring plant performance and community dynamics.  Past: Research Associate, USDA-APHIS, Florida, chasing citrus canker movement, Current: PhD student at UNC-Charlotte

Michelle Sipe (M.S. Biology 2004)  Thesis title:  Specificity in the induction and effects of chemical defenses in Brassica kaber  Current: 

Cherissa Rainey  Undergraduate research technician 2006-2008

 

Bill Gruner was an undergraduate technician in the lab during 2004-2005.  Current: Graduate of MS Program in Biological Sciences, Wright State, Researcher at WPAFB

Jeremiah Busch served as a research technician during the summer of 2000, who went on to receive his PhD at Indiana University in 2005
Current: faculty at Washington State University

Neela Kumar was a M.S. student in the lab for 9 months during 2001. 
Current:  Graduate of PhD program in Botany at Miami University, OH.

Caleb Slemmons was an independent researcher and undergraduate research technician from Fall 1999-Spring 2002, who went on to receive an MS in Environmental Sciences at Miami University, OH in 2005

Current:

 Lori Healey was an undergraduate research technician from Spring 2002 to Spring 2003, who went on to complete her MS in Microbiology and Immunology at Wright State in 2005

If you are interested in learning more about our lab, please contact me by phone or e-mail.

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Courses

Fall Quarter 2011
Bio 444 Plant Physiology

Winter Quarter 2012
ES 712 Genes, Organisms, and Ecosystems

Spring Quarter 2012
Bio 800 Field Ecology Seminar

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Personal Interests

Hiking, camping, live music, sports   ... and good beer

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Hot List

 Wright State Department of Biological Sciences Home Page

 Environmental Sciences PhD Program

  Ecological Society of America


 

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Last Revised: April 2011