342 Martinson et al.: Water Stress and Geographic Origin Influence Borer Infestations in Ash stressed or already declining (Liu et al. 2003; Baranchikov et al. 2008). Additionally, Manchurian ash did not suffer increasing mortality when exposed to EAB in a three-year experiment (Rebek et al. 2008) and has persisted in a long-term exper- imental plantation near the epicenter of the EAB outbreak (D.A. Herms, pers. comm.). Together, these findings suggest that cultivated and wild- type Manchurian ash should persist through the EAB invasion in North America where water stress is minimized and where edaphic and climatic conditions are favorable (Drenkhan et al. 2014). Retaining, planting, and breeding Manchurian ash with North American species may be key to sustaining the genus Fraxinus in North America and potentially preserving the multitude of arthro- pod species that use ash trees during all or part of their lifecycle (Gandhi and Herms 2010b). The researchers suggest that further study of the sus- ceptibility of mature Manchurian ash to North American borers would ensure that planting and breeding schemes are as successful as possible in light of both exotic and North American borers. Acknowledgments. We are grateful for the help of Mehdi Allaverd, Sarah Bass, Erik Bergmann, Christina Chavez, Samuel Grimard, and Brian Raupp in the field, and Dick Bean, Charles Pickett, Kim Rice, and the staff at the MDA for expertise and assistance. We thank Bill Lamp for technical assistance. Dan Herms and an anonymous reviewer provided valuable comments that improved the manuscript. We are grateful for the donation of ash trees from Bailey Nursery and for field access from the Fairfax County For- estry Division. All work was conducted under an APHIS PPQ 526 permit. This study was conducted in cooperation with the USDA-APHIS-PPQ Center for Plant Health Science and Tech- nology, which also provided funding for the project (Project No. 08-8100-1330-CA). This work was also supported by the USDA National Institute of Food and Agriculture, McIntire Stennis proj- ects 0213895 and 1003486. LITERATURE CITED Baranchikov, Y., E. Mozolevskaya, G. Yurchenko, and M. Kenis. 2008. Occurrence of the emerald ash borer, Agrilus planipennis, in Russia and its potential impact on European forestry. EPPO Bulletin 38:233–238. Cappaert, D., D.G. McCullough, T.M. Poland, and N.W. Siegert. 2005. Emerald ash borer in North America: A research and regulatory challenge. American Entomologist 51:152–165. Cappuccino, N., and D. Carpenter. 2005. Invasive exotic plants suffer less herbivory than non-invasive exotic plants. Biology Letters 1:435–438. Cipollini, D., Q. Wang, J.G.A. Whitehill, J.R. Powell, P. Bonello, and D.A. Herms. 2011. Distinguishing defensive characteristics in the phloem of ash species resistant and susceptible to emerald ash borer. Journal of Chemical Ecology 37:450–459. Crawley, M.J. 2007. The R Book. Wiley, West Sussex, England. 942 pp. Cregg, B.M., and M.E. Dix. 2001. Tree moisture stress and insect damage in urban areas in relation to heat island effects. Journal of Arboriculture 27:8–17. Cumming, A.B., D.B. Twardus, and W.D. Smith. 2006. Maryland and Massachusetts street tree monitoring pilot projects. National Forest Health Monitoring Program NA-FR-01-06. U.S. De- partment of Agriculture, Northeastern Area State and Private Forestry, Newtown Square, Pennsylvania. Desurmont, G.A., M.J. Donoghue, W.L. Clement, and A.A. Agrawal. 2011. Evolutionary history predicts plant defense against an invasive pest. Proceedings of the National Academy of Sciences of the United States of America 108:7070–7074. Drenkhan, R., H. Sander, and M. Hanso. 2014. Introduction of Manchurian ash (Fraxinus mandshurica Rupr.) to Estonia: Is it related to the current epidemic on European ash (F. excel- sior L.)? European Journal of Forest Research. DOI: 10.1007/ s10342-014-0811-9. Duan, J.J., M.D. Ulyshen, L.S. Bauer, J. Gould, and R. Van Driesche. 2010. Measuring the impact of biotic factors on populations of immature emerald ash borers (Coleoptera: Buprestidae). Envi- ronmental Entomology 39:1513–1522. Eyles, A., W. Jones, K. Riedl, D. Cipollini, S. Schwartz, K. Chan, D.A. Herms, and P. Bonello. 2007. Comparative phloem chemistry of Manchurian (Fraxinus mandshurica) and two North American ash species (Fraxinus americana and Fraxinus pennsylvanica). Journal of Chemical Ecology 33:1430–1448. Gandhi, K.J.K., and D.A. Herms. 2010a. Direct and indirect effects of alien insect herbivores on ecological processes and interac- tions in forests of eastern North America. Biological Invasions 12:389–405. Gandhi, K.J.K., and D.A. Herms. 2010b. North American arthro- pods at risk due to widespread Fraxinus mortality caused by the alien emerald ash borer. Biological Invasions 12:1839–1846. Haack, R.A. 2006. Exotic bark- and wood-boring Coleoptera in the United States: Recent establishments and interceptions. Canadian Journal of Forest Research-Revue Canadienne De Recherche Forestiere 36:269–288. Hanks, L.M., T.D. Paine, J.G. Millar, C.D. Campbell, and U.K. Schuch. 1999. Water relations of host trees and resistance to the phloem-boring beetle Phoracantha semipunctata F. (Coleoptera: Cerambycidae). Oecologia 119:400–407. Herms, D.A. 2002. Strategies for deployment of insect resistant ornamental plants, pp. 217–237. In: Wagner, M.R., K.M. Clancy, F. Lieutier, and T.D. Paine (Eds.). Mechanisms and Deployment of Resistance in Trees to Insects. Kluwer Academic, Dordrecht, The Netherlands. Herms, D.A. 2007. Distinguishing Emerald Ash Borer from Native Borers. Ohio State University OARDC Extension Publication Fact Sheet. Wooster, Ohio. Herms, D.A., and D.G. McCullough. 2014. Emerald ash borer inva- sion of North America: History, biology, ecology, impacts, and management. Annual Review of Entomology 59:13–30. Huberty, A.F., and R.F. Denno. 2004. Plant water stress and its con- sequences for herbivorous insects: A new synthesis. Ecology 85:1383–1398. Johnson, W.T., and H.H. Lyon. 1994. Insects that Feed on Trees and Shrubs, 2nd edition, revised. Cornell University Press, Ithaca, New York, U.S. ©2014 International Society of Arboriculture
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