David Ward

David Ward

Biological Sciences
Professor and Herrick Chair
Contact Information
(330) 672-9394


David Ward

Art and Margaret Herrick Endowed Professor of Plant Biology

Research Interests

My research interests lie in the field of the ecology of plant species redistributions. This includes studying both invasive and encroaching plant species. I am also interested in studying the natural process of succession. Most of my research involves trees. I also study the effects of herbivory by large mammals (such as elephants) on the population biology, community ecology and conservation of plant populations. I firmly believe in the value of field experiments to allow us to gain a mechanistic understanding of the factors that create large-scale vegetation patterns.


CURRENTLY ACCEPTING STUDENTS: PhD student to study the ecology and ecophysiology of encroaching and resident eastern red cedars


Selected Recent Publications

Link to Google Scholar Citations: https://scholar.google.com/citations?user=pTy-CwcAAAAJ&hl=en

Link to ISI list of publications: http://www.researcherid.com/rid/E-9814-2010



Ward, D. 2016. The biology of deserts. Oxford University Press, Oxford, UK. 2nd edition.

Griffiths, M.E., Davis, M.A. & Ward, D. 2020. Problem Plants of Ohio. Kent State University Press, Kent, U.S.A.


Journal Articles:

  1. Grellier, S., Janeau, J.-L., Richard, P., Florsch, N., Ward, D., Bariac, T. & Lorentz, S. 2020. Water uptake plasticity of Acacia in encroached grassland: small trees follow the adults. African Journal of Range and Forage Science [doi: 10.2989/10220119.2020.1834453].
  2. McMahon, K. & Ward, D. 2020. The effects of tree canopies on invasive Lantana camara: a follow-up study 18 years later. African Journal of Range and Forage Science [doi: 10.2989/10220119.2020.1850522].
  3. Pillay, T.P. & Ward, D. 2020. Grass competition is more important than fire for suppressing encroachment of Acacia sieberiana saplings. Plant Ecology [doi: 10.1007/s11258-020-01094-1].
  4. Ward, D. 2020. Shade is the most important factor limiting growth of a woody range expander. PLoS ONE 15: e0242003.
  5. Schmitt, M.H., Shrader, A.M. & Ward, D. 2020. Megaherbivore browsers vs. tannins: is being big enough? Oecologia 194: 383–390. [doi: 10.1007/s00442-020-04784-9].
  6. Valencia, E. et al. (incl. Ward, D.) 2020. Synchrony matters more than species richness in plant community stability at a global scale. Proceedings of the National Academy of Sciences, U.S.A. 117: 24345–24351 [doi: 10.1073/pnas.1920405117].
  7. Baldarelli, L., Throop, H. L., Collins, S. L. & Ward, D. 2020. Nutrient additions have direct and indirect effects on biocrust biomass in a long-term Chihuahuan Desert grassland experiment. Journal of Arid Environments 184: e104317. [doi: 10.1016/j.jaridenv.2020.104317].
  8. Ward, D., Kirkman, K.P., Tsvuura, Z., Morris, C.D. & Fynn, R.W.S. 2020. Are there common assembly rules for different grasslands? Comparisons of long-term data from a subtropical grassland with temperate grasslands. Journal of Vegetation Science 31:780–791. [doi: 10.1111/jvs.12906].
  9. De Bello, F., Valencia, E., Ward, D. & Hallett, L. 2020. Why we still need to invest in permanent plots for vegetation science. Journal of Vegetation Science 31:679–685. [doi: 10.1111/jvs.12928].
  10. Ward, D., Schmitt, M.H., & Shrader, A.M. 2020. Are there phylogenetic differences in salivary tannin-binding proteins between browsers and grazers, and ruminants and hind-gut fermenters? Ecology and Evolution 10:10426–10439. [doi: 10.1002/ece3.6698].
  11. Perkovich, C. & Ward, D. 2020. Protein: carbohydrate ratios in the diet of gypsy moth Lymantria dispar affect its ability to digest tannins. Journal of Chemical Ecology 46: 299–307. [doi: 10.1007/s10886-020-01161-x].
  12. Schmitt, M.H., Shrader, A.M. & Ward, D. 2020. Salivary tannin-binding proteins: a foraging advantage for goats? Livestock Science 234: e103974. [doi: 10.1016/j.livsci.2020.103974].
  13. Schmitt, M.H., Shuttleworth, A., Shrader, A.M. & Ward, D. 2020. The role of volatile plant secondary metabolites as pre-ingestive cues and potential toxins dictating diet selection by African elephants. Oikos 129: 24–34. [doi: 10.1111/oik.06665].
  14. Swemmer, A. & Ward, D. 2020. Patterns and determinants of woody plant growth in savannas. Pages 331–438 in P. F. Scogings and M. Sankaran, editors. Savanna Woody Plants and Large Herbivores. Cambridge University Press, Cambridge, U.K. [doi: 10.1002/9781119081111.ch12].
  15. Mureva, A., Ward, D., Pillay, T.P., Chivenge, P. & Cramer, M.D. 2018. Soil organic carbon increases in semi-arid regions while it decreases in humid regions due to woody-plant encroachment of grasslands of South Africa. Scientific Reports 8: art15506. [doi: 10.1038/s41598-018-33701-7].
  16. Koerner, S.E., Smith, M.D., Burkepile, D.E., Hanan, N., Avolio, M.L., Collins, S.L., Knapp, A.K., Lemoine, N.P., Forrestel, E.J., Eby, S., Thompson, D.I., Aguado-Santacruz, G., Anderson, J.P., Anderson, M., Angassa, A., Bagchi, S., Bakker, E.S., Bastin, G., Baur, L.E., Beard, K.H., Beever, E.A., Bohlen, P.J., Boughton, E.H., Canestro, D., Cesa, A., Chaneton, E., Cheng, J., D'Antonio, C.M., Deleglise, C., Dembélé, F., Dorrough, J., Eldridge, D., Fernandez-Going, B., Fernández-Lugo, S., Fraser, L.H., Freedman, B., Garcia-Salgado, G., Goheen, J.R., Guo, L., Husheer, S., Karembé, M., Knops, J.M.H., Kraaij, T., Kulmatiski, A., Kytöviita, M., Lezama, F., Loucougaray, G., Loydi, A., Milchunas, D.G., Milton, S., Morgan, J.W., Moxham, C., Nehring, K.C., Olff, H., Palmer, T.M., Rebollo, S., Riginos, C., Risch, A.C., Rueda, M., Sankaran, M., Sasaki, T., Schoenecker, K., Schultz, N.L., Schütz, M., Schwabe, A., Siebert, F., Smit, C., Stahlheber, K.A., Storm, C., Strong, D.J., Su, J., Tiruvaimozhi, Y.V., Tyler, C., Val, J., Vandegehuchte, M.L., Veblen, K.E., Vermeire, L.T., Ward, D., Wu, J., Young, T.P., Yu, Q. & Zelikova, T.J. 2018. Change in dominance determines herbivore effects on plant biodiversity. Nature Ecology and Evolution 2: 1925-1932. [doi: 10.1038/s41559-018-0696-y].
  17. Tomiolo, S. & Ward, D. 2018. Species migrations and invasions: A global synthesis of causes and consequences of range expansions. Perspectives in Plant Ecology, Evolution and Systematics 33: 62-77. [doi: 10.1016/j.ppees.2018.06.001].
  18. Tomiolo, S. & Ward, D. 2018. Soil properties and climate mediate the effects of biotic interactions on the performance of a woody range-expander. Ecosphere 9: e02186. [doi: 10.1002/ecs2.2186].
  19. Schmitt, M.H., Shuttleworth, A., Ward, D. & Shrader, A.M. 2018. African elephants use plant odours to make foraging decisions across multiple spatial scales. Animal Behaviour 141: 17-27. [doi: 10.1016/j.anbehav.2018.04.016].
  20. Ward, D., Trinogga, J., Wiegand, K., Du Toit, J., Okubamichael, D., Reinsch, S. & Schleicher, J. 2018. Large shrubs increase soil nutrients in a semi-arid savanna. Geoderma 310: 153-162. [doi: 10.1016/j.geoderma.2017.09.023].
  21. Ward, D., Kirkman, K., Hagenah, N. & Tsvuura, Z. 2017. Soil respiration declines with increasing nitrogen fertilization and is not related to productivity in long-term grassland experiments. Soil Biology & Biochemistry 115: 415–422. [doi: 10.1016/j.soilbio.2017.08.035].
  22. Mureva, A. & Ward, D. 2017. Soil microbial biomass and functional diversity in shrub-encroached grasslands along a precipitation gradient. Pedobiologia 63: 37-45. [doi: 10.1016/j.pedobi.2017.06.006].
  23. Ward, D., Kirkman, K. & Tsvuura, Z. 2017. An African grassland responds similarly to long-term fertilization to the Park Grass experiment. PLoS ONE 12: e0177208. [doi: 10.1371/journal.pone.0177208].
  24. Ward, D., Muller, K. & Shrader, A.M. 2017. Soil fertility on granite and sedimentary soils is associated with seasonal differences in foraging by elephants. Plant and Soil 413: 73-81. [doi: 10.1007/s11104-016-3067-y].
  25. Accatino, F., Wiegand, K., Ward, D. & De Michele, C. 2017. Carrying capacity in arid rangelands during droughts: the role of temporal and spatial thresholds. Animal 11: 309-317. [doi: 10.1017/S1751731116001531].
  26. Griffiths, M.E., Ruiz, N., Ward, D., 2017. Mistletoe species richness patterns are influenced more by host geographic range than nitrogen content. African Journal of Ecology 55, 101–110.
  27. Okubamichael, D.Y., Griffiths, M.E. & Ward, D. 2016. Host specificity in parasitic plants - perspectives from mistletoes. AoB Plants plw069. [doi: 10.1093/aobpla/plw069].
  28. Russell, J.M. & Ward, D. 2016. Historical land use and vegetation change in northern KwaZulu-Natal, South Africa. Land Degradation & Development 27: 1691-1699. [doi: 10.1002/ldr.2476].
  29. Accatino, F., Wiegand, K., Ward, D. & De Michele, C. 2016. Trees, grass, and fire in humid savannas - the importance of life history traits and spatial processes. Ecological Modelling 320: 135-144. F1000Prime Recommended.
  30. Schmitt, M.H., Ward, D. & Shrader, A.M. 2016. Incorporating secondary metabolites, tannin-binding proteins, and diet breadth into carrying-capacity models for African Elephants. Ecological Modelling 332: 8–18.
  31. Ward, D. 2016. Clipping frequency but not nutrients affect the architecture and non-structural carbohydrates of a browsing lawn. Plant Ecology 217: 21-29.
  32. Tjelele, T.J., Ward, D. & Dziba, L.E. 2015. The effects of passage through the gut of goats and cattle, and the application of dung as a fertilizer on seedling establishment of Dichrostachys cinerea and Acacia nilotica. The Rangeland Journal 37: 147–156.
  33. Tjelele, T.J., Ward, D. & Dziba, L.E. 2015. The effects of seed ingestion by livestock, dung fertilization, trampling, grass competition and fire on seedling establishment of two woody plant species. PLoS One 10(2): e0117788. [doi:10.1371/journal.pone.0117788].
  34. Ford, A.T., Goheen, J.R., Otieno, T.O., Binder, L., Isbell, L., Palmer, T.M., Ward, D., Woodroffe, R. & Pringle, R.M. 2014. Large carnivores make savanna tree communities less thorny. Science 346: 346-349.



Fall Semester Spring Semester
Odd Years Even Years Odd Years Even Years

Plant Ecology (3 Cr.), U/G

Climate Change Biology (3Cr.), U/G

Invasion Biology (3 Cr.), U/G

Climate Change Biology (3 Cr.), U/G

Field Course to South Africa (5 Cr.) U/G Evolutionary Biology Directed Reading (2 Cr.), G only

U = undergraduate students     G = graduate students




Plant Ecology, Invasion Biology, Biological Sciences, Faculty Advisor, environmental knowledge of poverty-stricken resource users in Africa, land use and degradation, desertification, plant conservation biology, plant population ecology, plant community ecology

Research Institutes and Initiatives

Environmental Science and Design Research Initiative, Global Understanding Research Initiative