Biological Invasions

, Volume 16, Issue 4, pp 859–873 | Cite as

Ash (Fraxinus spp.) mortality, regeneration, and seed bank dynamics in mixed hardwood forests following invasion by emerald ash borer (Agrilus planipennis)

  • Wendy S. KloosterEmail author
  • Daniel A. Herms
  • Kathleen S. Knight
  • Catherine P. Herms
  • Deborah G. McCullough
  • Annemarie Smith
  • Kamal J. K. Gandhi
  • John Cardina
Original Paper


Emerald ash borer (EAB; Agrilus planipennis) has killed millions of ash trees and threatens ash throughout North America, and long-term persistence of ash will depend on the potential for regeneration. We quantified ash demography, including mortality and regeneration, of Fraxinus americana (white ash), Fraxinus pennsylvanica (green ash), and Fraxinus nigra (black ash) in mixed hardwood forests near the epicenter of the EAB invasion in southeastern Michigan and throughout Ohio. Plots were established across a gradient of ash densities. Ash was the most important species in hydric sites, and ranked second among all species in mesic and xeric sites. In sites nearest the epicenter in Michigan, ash mortality exceeded 99 % by 2009, and few or no newly germinated ash seedlings were observed, leaving only an “orphaned cohort” of established ash seedlings and saplings. As ash mortality increased, the number of viable ash seeds in soil samples decreased sharply, and no viable seeds were collected in 2007 or 2008. In Ohio sites farther from the epicenter, densities of new ash seedlings were much higher in plots with healthy ash trees compared to plots where trees had died. EAB was still present in low densities in Michigan and Ohio stands in 2012 where average mortality of ash was nearly 100 %. The future of ash at these sites will depend on the outcome of the dynamic interaction between the orphaned cohort of previously established ash seedlings and saplings and low density EAB populations.


Ash regeneration Demography Invasive forest pest Seedlings 



For their extensive assistance with field and laboratory work we thank Limbania Aliega, Lourdes Arrueta, Stephanie Blumer, Alejandra Claure, Christian Colindres, Kyle Costilow, Tara Dell, Mary Douglas, Eileen Duarte, Charles Flower, Stefanie Fluke, Tim Fox, Joan Jolliffas, Lawrence Long, Rina Mejía, Rachel Nieswander, Sarahí Nuñez, Delmy Sánchez, Sebastián Sáenz, Stephanie Smith, and Nathan Yaussy, as well as other members of the Herms, Knight and McCullough Labs. We are especially grateful to Diane Hartzler who organized and coordinated much of the field work in Michigan since the inception of the study. We also thank the two anonymous reviewers for their constructive suggestions that helped us improve this manuscript. This research was funded by grants from the USDA Forest Service Northeastern Research Station’s Research on Biological Invasions of Northeastern Forests program; USDA National Research Initiative Biology of Weedy and Invasive Species in Agroecosystems competitive grants program; the National Institute of Food and Agriculture; Cooperative Agreements with the USDA Forest Service Northern Research Station, Delaware, OH; and state and federal funds appropriated to the Ohio Agricultural Research and Development Center and The Ohio State University.


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Copyright information

© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  • Wendy S. Klooster
    • 1
    Email author
  • Daniel A. Herms
    • 2
  • Kathleen S. Knight
    • 3
  • Catherine P. Herms
    • 1
  • Deborah G. McCullough
    • 4
  • Annemarie Smith
    • 2
  • Kamal J. K. Gandhi
    • 2
    • 5
  • John Cardina
    • 1
  1. 1.Department of Horticulture and Crop Science, Ohio Agricultural Research and Development CenterThe Ohio State UniversityWoosterUSA
  2. 2.Department of Entomology, Ohio Agricultural Research and Development CenterThe Ohio State UniversityWoosterUSA
  3. 3.Forestry Sciences Laboratory, Northern Research StationUSDA Forest ServiceDelawareUSA
  4. 4.Departments of Entomology and ForestryMichigan State UniversityEast LansingUSA
  5. 5.Daniel B. Warnell School of Forestry and Natural ResourcesThe University of GeorgiaAthensUSA

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