, Volume 19, Issue 2, pp 248–270 | Cite as

Potential Species Replacements for Black Ash (Fraxinus nigra) at the Confluence of Two Threats: Emerald Ash Borer and a Changing Climate

  • Louis Iverson
  • Kathleen S. Knight
  • Anantha Prasad
  • Daniel A. Herms
  • Stephen Matthews
  • Matthew Peters
  • Annemarie Smith
  • Diane M. Hartzler
  • Robert Long
  • John Almendinger


The emerald ash borer (Agrilus planipennis; EAB) is causing widespread mortality of ash (Fraxinus spp.) and climate change is altering habitats of tree species throughout large portions of North America. Black ash (F. nigra), a moist-soil species common in the Northwoods of Minnesota, Wisconsin, and Michigan, USA, is under a double threat of losing habitat from climate change and near annihilation from EAB. Because black ash often occurs in nearly pure stands, planting non-ash species is a management strategy already underway or being planned for thousands of acres. Tools are needed to assist managers in prioritizing sites for early treatment and to select potential species to replace black ash. This study explores the implications of threats to black ash ecosystems using analyses of field data and models to assess both the threats to, and potential replacement species for, black ash in Minnesota. For our analysis we (1) assessed the status of ashes and co-occurring species in forest inventory plots throughout Minnesota; (2) modeled the risk of EAB attack for multiple years in Minnesota; (3) modeled potential impacts of climate change on tree species with current or potential future habitat in Minnesota; (4) evaluated species co-occurring with black ash in plots in Ohio and Michigan, southeast of Minnesota; and (5) synthesized these results to provide a classification for candidate replacement species, both from within Minnesota and from points farther south. Though this process is demonstrated for black ash in Minnesota, the elements to be considered and modeled would be similar for any other location with a pest or pathogen threat for a species which simultaneously faces a changing climate.


assisted range expansion invasive insect spread model climate change emerald ash borer multiple forest threats restoration species distribution models 



The work was sponsored by the authors’ primary research institutions: the Northern Research Station of the US Forest Service, The Ohio State University, and the Minnesota Department of Natural Resources. The authors are indebted to many, many organizations and individuals for the data represented here—the people who sampled the thousands of FIA plots and the others who provided the analysis and maps of ash and other species across the region; those who provided data on EAB abundance, major roads and traffic density, campground size and usage, the wood products industry, and human population density; those who sampled all the mosquito-laden plots in Michigan and Ohio; and those carrying out the planting experiment on the Chippewa National Forest. We are truly indebted to these people, for without their field-based and mapping efforts none of these studies would have been possible. Thanks to Rich McCullough, US Forest Service, for assistance with FIA queries. Thanks also to Tony D’Amato, Stephen Handler, Herman Shugart, and anonymous reviewers for helpful comments on the manuscript.


  1. Anderson MK, Nelson G. 2003. Black ash, Fraxinus nigra Marsh. NRCS plant guide. Natural Resources Conservation Service.
  2. Beyers B. 2013. Rising from the ashes: What happens when the trees disappear from the forest? St. Paul, MN: University of Minnesota, College of Food, Agricultural and Natural Resource Sciences.
  3. Cowardin LM, Carter V, Golet FC, LaRoe ET. 1979. Classification of wetlands and deepwater habitats of the United States. FWS/OBS 79/31. Washington, DC: US Department of the Interior, Fish and Wildlife Service.Google Scholar
  4. Crosthwaite JC, Sobek S, Lyons DB, Bernards MA, Sinclair BJ. 2011. The overwintering physiology of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). J Insect Physiol 57:166–73.CrossRefPubMedGoogle Scholar
  5. Delworth T, Broccoli A, Rosati A, Stouffer RJ et al. 2006. GFDL’s CM2 global coupled climate models—Part 1—formulation and simulation characteristics. J Climate 19:643–74.CrossRefGoogle Scholar
  6. DeSantis RD, Moser WK, Gormanson DD, Bartlett MG, Vermunt B. 2013a. Effects of climate on emerald ash borer mortality and the potential for ash survival in North America. Agric For Meteorol 178–179:120–8.CrossRefGoogle Scholar
  7. DeSantis RD, Moser WK, Huggett RJ, Li R, Wear DN, Miles PD. 2013b. Modeling the effects of emerald ash borer on forest composition in the Midwest and Northeast United States. Gen. Tech. Rep. NRS-112. Newtown Square, PA: US Department of Agriculture, Forest Service, Northern Research Station.Google Scholar
  8. Flower CE, Knight KS, Gonzalez-Meler MA. 2013. Impacts of the emerald ash borer (Agrilus planipennis) induced ash (Fraxinus spp.) mortality on forest carbon cycling and successional dynamics in the eastern United States. Biol Invasions 15:931–44.CrossRefGoogle Scholar
  9. Gandhi KK, Herms D. 2010. Direct and indirect effects of alien insect herbivores on ecological processes and interactions in forests of eastern North America. Biol Invasions 12:389–405.CrossRefGoogle Scholar
  10. Hanberry BB, Palik BJ, He HS. 2013. Winning and losing tree species of reassembly in Minnesota’s mixed and broadleaf forests. Plos One 8:e61709.CrossRefPubMedPubMedCentralGoogle Scholar
  11. Handler S, Duveneck M, Iverson L, Peters E, Scheller R, Wythers K, Brandt L, Butler P, Janowiak M, Swanston C, Kolka R, McQuiston C, Palik B, Turner C, White M, Adams C, Barrett K, D’Amato A, Hagell S, Johnson R, Johnson P, Larson M, Matthews S, Montgomery R, Olsen S, Peters M, Prasad A, Rajala J, Reich P, Shannon PD, Daley J, Davenport M, Emery M, Fehringer D, Hoving C, Johnson G, Johnson L, Neitzel D, Rissman A, Rittenhouse C, Ziel R. 2014. Minnesota forest ecosystem vulnerability assessment and synthesis: A report from the Northwoods Climate Change Response Framework. Gen. Tech. Rep. NRS-133. Newtown Square, PA: US Department of Agriculture, Forest Service, Northern Research Station.Google Scholar
  12. Herms DA, McCullough DG. 2014. Emerald ash borer invasion of North America: history, biology, ecology, impact and management. Ann Rev Entomol 59:13–30.CrossRefGoogle Scholar
  13. Intergovernmental Panel on Climate Change (IPCC). 2014. Climate Change 2014: Impacts, Adaptation, and Vulnerability. Working Group II Contribution to the IPCC 5th Assessment Report Intergovernmental Panel on Climate Change, Stanford, CA.Google Scholar
  14. Iverson L, Prasad AM, Matthews S, Peters M. 2011. Lessons learned while integrating habitat, dispersal, disturbance, and life-history traits into species habitat models under climate change. Ecosystems 14:1005–20.CrossRefGoogle Scholar
  15. Iverson LR, Prasad A, Bossenbroek J, Sydnor D, Schwartz MW. 2010. Modeling potential movements of the emerald ash borer: the model framework. In: Pye J, Raucher M, Sands Y, Lee D, Beatty J, Eds. Advances in threat assessment and their application to forest and rangeland management. Portland, OR: US Department of Agriculture, Forest Service, Pacific Northwest and Southern Research Stations. p 581–97.Google Scholar
  16. Iverson LR, Prasad AM, Matthews SN, Peters M. 2008. Estimating potential habitat for 134 eastern US tree species under six climate scenarios. For Ecol Manag 254:390–406.CrossRefGoogle Scholar
  17. Kashian DM, Witter JA. 2011. Assessing the potential for ash canopy tree replacement via current regeneration following emerald ash borer-caused mortality on southeastern Michigan landscapes. For Ecol Manag 261:480–8.CrossRefGoogle Scholar
  18. Klooster W, Herms D, Knight K, Herms C, McCullough D, Smith A, Gandhi KK, Cardina J. 2014. Ash (Fraxinus spp.) mortality, regeneration, and seed bank dynamics in mixed hardwood forests following invasion by emerald ash borer (Agrilus planipennis). Biol Invasions 16:859–73.CrossRefGoogle Scholar
  19. Knight KS, Slavicek JM, Kappler R, Pisarczyk E, Wiggin B, and Menard K. 2012a. Using Dutch elm disease- tolerant elm to restore floodplains impacted by emerald ash borer. In: Sniezko RA, Yanchuk AD, Kliejunas JT, Palmieri KM, Alexander JM, and Frankel S.J. Tech. Coords. Proceedings of the 4th International Workshop on the Genetics of Host-Parasite Interactions in Forestry: Disease and Insect Resistance in Forest Trees. July 31–August 5 2011, Euguene, OR. Gen. Tech. Rep. PSW-GTR-240. Albany, CA: U.S.Google Scholar
  20. Knight KS, Herms D, Plumb R, Sawyer E, Spalink D, Pisarczyk E, Wiggin B, Kappler R, Ziegler E, Menard K. 2012b. Dynamics of surviving ash (Fraxinus spp.) populations in areas long infested by emerald ash borer (Agrilus planipennis). Sniezko R, Yanchuk A, Kliejunas J, Palmieri K, Alexander J, Frankel S editors. Proceedings of the 4th international workshop on the genetics of host-parasite interactions in forestry: disease and insect resistance in forest trees. Gen. Tech. Rep. PSW-GTR-240. Albany, CA: US Department of Agriculture, Forest Service, Pacific Southwest Research Station, p143-152.Google Scholar
  21. Knight KS, Brown JP, Long RP. 2013. Factors affecting the survival of ash trees (Fraxinus spp.) infested by emerald ash borer (Agrilus planipennis). Biolog Invasions 15:371–83.CrossRefGoogle Scholar
  22. Knight KS, Flash BP, Kappler RH, Throckmorton JA, Grafton B, Flower CD. 2014. Monitoring ash (Fraxinus spp) decline and emerald ash borer (Agrilus planipennis) symptoms in infested areas. Gen. Tech. Rep. NRS-139. Newtown Square: U.S. Department of Agriculture, Forest Service, Northern Research Station.Google Scholar
  23. Koch F, Yemshanov D, Colunga-Garcia M, Magarey R, Smith W. 2011. Potential establishment of alien-invasive forest insect species in the United States: where and how many? Biol Invasions 13:969–85.CrossRefGoogle Scholar
  24. Koch J, Carey D, Knight K, Poland T, Herms D, Mason M. 2012. Breeding strategies for the development of emerald ash borer resistant—North American ash. Sniezko R, Yanchuk A, Kliejunas J, Palmieri K, Alexander J, Frankel S editors. Proceedings of the 4th international workshop on the genetics of host-parasite interactions in forestry: disease and insect resistance in forest trees. Gen. Tech. Rep. PSW-GTR-240. Albany, CA: US Department of Agriculture, Forest Service, Pacific Southwest Research Station. pp 143–152.Google Scholar
  25. Koch JL, Carey DW, Mason ME, Poland TM, Knight KS. 2015. Interspecific variation in Fraxinus pennsylvanica responses to emerald ash borer (Agrilus planipennis). New Forests 46:995–1011.CrossRefGoogle Scholar
  26. Landscape Change Research Group. 2014. Climate change atlas. Delaware, OH: US Department of Agriculture, Forest Service, Northern Research Station.
  27. Lodge DM, Williams S, MacIsaac HJ, Hayes KR, Leung B, Reichard S, Mack RN, Moyle PB, Smith M, Andow DA, Carlton JT, McMichael A. 2006. Biological invasions: recommendations for U.S. policy and management. Ecol Appl 16:2035–54.CrossRefPubMedGoogle Scholar
  28. Looney C, Damato A, Palik B, Slesak RA. 2015. Overstory treatment and planting season affect survival of replacement tree species in emerald ash borer-threatened Fraxinus nigra forests in Minnesota, USA. Can J For Res 45:1728–38.Google Scholar
  29. Lunt ID, Byrne M, Hellmann JJ, Mitchell NJ, Garnett ST, Hayward MW, Martin TG, McDonald-Maddden E, Williams SE, Zander KK. 2013. Using assisted colonisation to conserve biodiversity and restore ecosystem function under climate change. Biol Conserv 157:172–7.CrossRefGoogle Scholar
  30. Matthews SN, Iverson LR, Prasad AM, Peters MP, Rodewald PG. 2011. Modifying climate change habitat models using tree species-specific assessments of model uncertainty and life history factors. For Ecol Manag 262:1460–72.CrossRefGoogle Scholar
  31. Minnesota Department of Natural Resources. 2003. Field guide to the Native Plant Communities of Minnesota: the Laurentian Mixed Forest Province. Ecological Land Classification Program, Minnesota County Biological Survey, and Natural Heritage and Nongame Research Program. St. Paul: MNDNR.Google Scholar
  32. Miles PD, Brand GJ. 2007. Minnesota’s forest resources in 2005 Resource Bulletin NRS-6. Newtown Square: US Department of Agriculture, Forest Service, Northern Research Station.Google Scholar
  33. Miles PD, Jacobson K, Brand GJ, Jepsen E, Meneguzzo D, Mielke ME, Olson C, Perry CHH, Piva R, Wilson BT, Woodall C. 2007. Minnesota’s forests 1999-2003 (Part A). Resource Bulletin NRS-12A. Newtown Square: US Department of Agriculture, Forest Service, Northern Research Station.Google Scholar
  34. Montoya D, Rogers L, Memmott J. 2012. Emerging perspectives in the restoration of biodiversity-based ecosystem services. Trends Ecol Evol 27:666–72.CrossRefPubMedGoogle Scholar
  35. Nakicenovic N, Alcamo J, Davis G, de Vries B, Fenhann J, Gaffin S, Gregory K, Grübler A, Jung TY, Kram T, Lebre La Rovere E, Michaelis L, Mori S, Morita T, Pepper W, Pitcher H, Price L, Riahi K, Roehrl A, Rogner H, Sankovski A, Schlesinger M, Shukla P, Smith S, Swart R, van Rooijen S, Victor N, Dadi Z. 2000. IPCC special report on emissions scenarios. Cambridge: Cambridge University Press.Google Scholar
  36. Nowacki GJ, Abrams MD. 2015. Is climate an important driver of post-European vegetation change in the Eastern United States? Global Change Biol 21:314–34.CrossRefGoogle Scholar
  37. Palik BJ, Ostry ME, Venette RC, Abdela E. 2011. Fraxinus nigra (black ash) dieback in Minnesota: regional variation and potential contributing factors. For Ecol Manag 261:128–35.CrossRefGoogle Scholar
  38. Palik BJ, Ostry ME, Venette RC, Abdela E. 2012. Tree regeneration in black ash (Fraxinus nigra) stands exhibiting crown dieback in Minnesota. For Ecol Manag 269:26–30.CrossRefGoogle Scholar
  39. Pederson N, D’Amato AW, Dyer JM, Foster DR, Goldblum D, Hart JL, Hessl AE, Iverson LR, Jackson ST, Martin-Benito D, McCarthy BC, McEwan RW, Mladenoff DJ, Parker AJ, Shuman B, Williams JW. 2015. Climate remains an important driver of post-European vegetation change in the eastern United States. Global Change Biol 2:2105–10.CrossRefGoogle Scholar
  40. Pedlar JH, McKenney DW, Aubin I, Beardmore T, Beaulieru J, Iverson LR, O’Neill GA, Winder RS, Ste-Marie C. 2012. Placing forestry in the assisted migration debate. Bioscience 62:835–42.CrossRefGoogle Scholar
  41. Petrice TR, Haack RA. 2011. Effects of cutting time, stump height, and herbicide application on ash (Fraxinus spp.) stump sprouting and colonization by emerald ash borer (Agrilus planipennis). Northern J Appl Forest 28:79–83.Google Scholar
  42. Pimentel D, McNair S, Janecka J, Wightman J, Simmonds C, O’Connell C, Wong E, Russel L, Zern J, Aquino T, Tsomondo T. 2001. Economic and environmental threats of alien plant, animal, and microbe invasions. Agric Ecosyst Environ 84:1–20.CrossRefGoogle Scholar
  43. Prasad A, Iverson L, Peters M, Bossenbroek J, Matthews SN, Sydnor D, Schwartz M. 2010. Modeling the invasive emerald ash borer risk of spread using a spatially explicit cellular model. Landsc Ecol 25:353–69.CrossRefGoogle Scholar
  44. Prasad AM, Iverson LR, Liaw A. 2006. Newer classification and regression tree techniques: bagging and random forests for ecological prediction. Ecosystems 9:181–99.CrossRefGoogle Scholar
  45. PRISM Climate Group. 2013. PRISM climate data. Oregon State University. Accessed 7 Dec 2013.
  46. Pryor SC, Scavia D, Downer C, Gladen M, Iverson L, Nordstrom R, Patz J, Robertson GP. 2014. Ch. 18: Midwest. Climate change impacts in the United States: the third national climate assessment. In: Melillo JM, Melillo TR, Richmond T, Yohe GW (eds) Washington, DC: US Global Change Research Program, pp 418–440.Google Scholar
  47. Reich PB, Sendall KM, Rice K, Rich RL, Stefanski A, Hobbie SE, Montgomery RA. 2015. Geographic range predicts photosynthetic and growth response to warming in co-occurring tree species. Nat Clim Change 5:148–52.CrossRefGoogle Scholar
  48. Saunders S, Findlay D, Easley T, Spencer T. 2012. Doubled trouble: more Midwestern extreme storms. Louisville: The Rocky Mountain Climate Organization and The Natural Resources Defense Council.Google Scholar
  49. Schwartz MW, Hellmann JJ, Jason MM, Sax D, Borevitz JO, Brennan J, Camacho AE, Ceballos G, Clark JR, Doremus H, Early R, Etterson JR, Fielder D, Gill JL, Gonzalez P, Green N, Hannah L, Jamieson D, Javeline D, Minteer BA, Odenbaugh J, Polasky S, Richardson DM, Root T, Safford HD. 2012. Managed relocation: integrating the scientific, regulatory, and ethical challenges. Bioscience 62:732–43.CrossRefGoogle Scholar
  50. Slesak RA, Lenhart CF, Brooks KN, D’Amato AW, Palik BJ. 2014. Water table response to harvesting and simulated emerald ash borer mortality in black ash wetlands in Minnesota, USA. Can J For Res 44:961–8.CrossRefGoogle Scholar
  51. Smith A, Herms DA, Long RP, Gandhi KJK. 2015. Community composition and structure had no effect on forest susceptibility to invasion by the emerald ash borer (Coleoptera: Buprestidae). Can Entomol 147:318–28.CrossRefGoogle Scholar
  52. Sobek-Swant S, Crosthwaite J, Lyons DB, Sinclair B. 2012. Could phenotypic plasticity limit an invasive species? Incomplete reversibility of mid-winter deacclimation in emerald ash borer. Biol Invasions 14:115–25.CrossRefGoogle Scholar
  53. Stern N. 2006. Stern Review: The Economics of Climate Change. London: HM Treasury.Google Scholar
  54. Stohlgren TJ, Resnik JR, Plumb GE. 2014. Climate change and ‘alien species in National Parks’: Revisited. In: Ziska LH, Dukes S, Eds. Invasive species and global climate change. Cambridge: CABI Publishing. Google Scholar
  55. Sydnor TD, Smith K, Heiligmann R. 2005. Ash replacements for urban and woodland plantings. Columbus: Ohio State University Extension. p 82p.Google Scholar
  56. University of Minnesota Extension and Minnesota Department of Natural Resources. 2011. Ash Management Guidelines for Private Forest Landowners. University of Minnesota Extension Service.
  57. Venette RC, Abrahamson M. 2010. Cold hardiness of emerald ash borer, Agrilus planipennis: a new perspective. In: Black ash symposium: Proceedings of the meeting, May. pp 25–27.Google Scholar
  58. Washington WM, Weatherly JW, Meehl GA, Semtner AJ Jr, Bettge TW, Craig AP, Strand WG Jr, Arblaster JM, Wayland VB, James R, Zhang Y. 2000. Parallel climate model (PCM) control and transient simulations. Clim Dyn 16:755–74.CrossRefGoogle Scholar
  59. Wei XIA, Wu YUN, Reardon R, Sun T-H, Lu MIN, Sun JH. 2007. Biology and damage traits of emerald ash borer (Agrilus planipennis Fairmaire) in China. Insect Sci 14:367–73.CrossRefGoogle Scholar
  60. Williams MI, Dumroese RK. 2013. Preparing for climate change: forestry and assisted migration. J For 111:287–97.Google Scholar
  61. Wilson BT, Lister AJ, Riemann RI. 2012. A nearest-neighbor imputation approach to mapping tree species over large areas using forest inventory plots and moderate resolution raster data. For Ecol Manag 271:182–98.CrossRefGoogle Scholar
  62. Wright JW, Rauscher HM. 1990. Fraxinus nigra Marsh, black ash. In: Burns RM, Honkala BH, Eds. Silvics of North America: 2. Hardwoods. US Department of Agriculture, Forest Service, Agricultural Handbook. Newtown Square: US Department of Agriculture, Forest Service. p 688–93.Google Scholar

Copyright information

© Springer Science+Business Media New York (outside the USA) 2015

Authors and Affiliations

  • Louis Iverson
    • 1
  • Kathleen S. Knight
    • 1
  • Anantha Prasad
    • 1
  • Daniel A. Herms
    • 2
  • Stephen Matthews
    • 1
    • 3
  • Matthew Peters
    • 1
  • Annemarie Smith
    • 4
  • Diane M. Hartzler
    • 2
  • Robert Long
    • 5
  • John Almendinger
    • 6
  1. 1.Northern Research StationUSDA Forest ServiceDelawareUSA
  2. 2.Department of Entomology, Ohio Agricultural Research and Development CenterThe Ohio State UniversityWoosterUSA
  3. 3.School of Natural ResourcesThe Ohio State UniversityColumbusUSA
  4. 4.US Green Building CouncilCentral Ohio ChapterColumbusUSA
  5. 5.Northern Research StationUSDA Forest ServiceIrvineUSA
  6. 6.Division of ForestryMinnesota Department of Natural ResourcesGrand RapidsUSA

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