Skip to main content

Resurrecting the Lost Flames of American Chestnut

Ecosystems volume 22, pages 995–1006 (2019)Cite this article

Abstract

Loss of native foundation tree species to introduced pests profoundly alters the structure and function of many forest ecosystems. Recent advances to resurrect or prevent the loss of species by developing resistant hybrids hold promise, but uncertainty remains about the potential impacts of introducing a novel genotype on ecological processes, such as fire. A classic example of a non-native, pathogen-caused loss of a foundation species is American chestnut (Castanea dentata), a species now functionally extinct from the eastern US but undergoing experimental trials to resurrect the species with a putatively disease-resistant genotype. We compared the litter flammability among American chestnut, Chinese chestnut (C. mollisima), and a population of first intercross, third-generation backcrossed hybrid (BC3F2) using laboratory burning experiments. Litter flammability of American chestnut was consistently greater than Chinese chestnut, whereas the hybrid had an intermediate flammability or more closely resembled Chinese chestnut flammability by some measures. Greater flammability in American chestnut was associated with a longer leaf structure with greater curling when dry. American chestnut had flammability properties consistent with other pyrophytic species present in contemporary fire-prone ecosystems. The loss of American chestnut may have altered litter flammability of some eastern US forest ecosystems, a result more commonly associated with compositional changes in conjunction with fire exclusion and other disturbances. Resurrecting American chestnut with a hybrid genotype may mitigate this change in areas where less flammable species have replaced American chestnut. Resurrection of lost foundation species through introduction of resistant genotypes may represent a resounding ecological success story, but unanticipated changes to ecological processes, such as fire, should be considered.

This is a preview of subscription content, access via your institution.

Access options

Buy single article

Instant access to the full article PDF.

39,95 €

Price includes VAT (Finland)

Figure 1
Figure 2
Figure 3
Figure 4

References

  • Agrawal A, Stephenson SL. 1995. Recent successional changes in a former chestnut-dominated forest in Southwestern Virginia. Castanea:107–113.

  • Aldrich SR, Lafon CW, Grissino-Mayer HD, DeWeese GG. 2014. Fire history and its relations with land use and climate over three centuries in the central Appalachian Mountains, USA. Williams J, editor. J Biogeogr 41:2093–104.

  • Anagnostakis SL. 1987. Chestnut blight: the classical problem of an introduced pathogen. Mycologia 79:23.

    Article  Google Scholar 

  • Ashe WW. 1912. Chestnut in Tennessee. Nashville, Tennessee: Baird-Ward Printing Co.

    Google Scholar 

  • Aukema JE, McCullough DG, Von Holle B, Liebhold AM, Britton K, Frankel SJ. 2010. Historical accumulation of nonindigenous forest pests in the continental United States. BioScience 60:886–97.

    Article  Google Scholar 

  • Bailey JK, Schweitzer JA, Rehill BJ, Lindroth RL, Martinsen GD, Whitham TG. 2004. Beavers as molecular geneticists: a genetic basis to the foraging of an ecosystem engineer. Ecology 85:603–8.

    Article  Google Scholar 

  • Beckage B, Platt WJ, Gross LJ. 2009. Vegetation, fire, and feedbacks: A disturbance-mediated model of savannas. Am Nat 174:805–18.

    Article  PubMed  Google Scholar 

  • Belair ED, Saunders MR, Bailey BG. 2014. Four-year response of underplanted American chestnut (Castanea dentata) and three competitors to midstory removal, root trenching, and weeding treatments in an oak-hickory forest. For Ecol Manag 329:21–9.

    Article  Google Scholar 

  • Belair ED, Saunders MR, Landhäusser SM. 2018. Growth traits of juvenile American chestnut and red oak as adaptations to disturbance. Restor Ecol 26:712–19.

    Article  Google Scholar 

  • Brooks ML, D’antonio CM, Richardson DM, Grace JB, Keeley JE, DiTomaso JM, Hobbs RJ, Pellant M, Pyke D. 2004. Effects of invasive alien plants on fire regimes. BioScience 54:677–88.

    Article  Google Scholar 

  • Brose PH, Dey DC, Waldrop TA. 2014. The fire—oak literature of eastern North America: synthesis and guidelines. Delaware, OH, USA: USDA Forest Service Northern Research Station http://www.academia.edu/download/42004891/The_fire-oak_literature_of_eastern_North20160203-30232-10ioxmj.pdf. Last accessed 27/06/2017.

  • Brown CE, Mickelbart MV, Jacobs DF. 2014. Leaf physiology and biomass allocation of backcross hybrid American chestnut (Castanea dentata) seedlings in response to light and water availability. Tree Physiol 34:1362–75.

    Article  CAS  PubMed  Google Scholar 

  • Clark S, McNab H, Loftis D, Zarnoch S. 2012. American chestnut growth and survival five years after planting in two silvicultural treatments in the southern Appalachians, USA. Forests 3:1017–33.

    Article  Google Scholar 

  • Clark SL, Schlarbaum SE, Pinchot CC, Anagnostakis SL, Saunders MR, Thomas-Van Gundy M, Schaberg P, McKenna J, Bard JF, Berrang PC, Casey DM, Casey CE, Crane B, Jackson BD, Kochenderfer JD, Lewis RF, MacFarlane R, Makowski R, Miller MD, Rodrigue JA, Stelick J, Thornton CD, Williamson TS. 2014. Reintroduction of American Chestnut in the National Forest System. J For 112:502–12.

    Google Scholar 

  • Clark SL, Schlarbaum SE, Saxton AM, Hebard FV. 2016. Establishment of American chestnuts (Castanea dentata) bred for blight (Cryphonectria parasitica) resistance: influence of breeding and nursery grading. New For 47:243–70.

    Article  Google Scholar 

  • Dayton PK. 1972. Toward an understanding of community resilience and the potential effects of enrichments to the benthos at McMurdo Sound, Antarctica. Lawrence, KS: Allen Press.

    Google Scholar 

  • Diskin M, Steiner KC, Hebard FV. 2006. Recovery of American chestnut characteristics following hybridization and backcross breeding to restore blight-ravaged Castanea dentata. For Ecol Manag 223:439–47.

    Article  Google Scholar 

  • Elliott KJ, Swank WT. 2008. Long-term changes in forest composition and diversity following early logging (1919–1923) and the decline of American chestnut (Castanea dentata). Plant Ecol 197:155–72.

    Article  Google Scholar 

  • Ellison AM, Bank MS, Clinton BD, Colburn EA, Elliott K, Ford CR, Foster DR, Kloeppel BD, Knoepp JD, Lovett GM, Mohan J, Orwig DA, Rodenhouse NL, Sobczak WV, Stinson KA, Stone JK, Swan CM, Thompson J, Von Holle B, Webster JR. 2005. Loss of foundation species: consequences for the structure and dynamics of forested ecosystems. Front Ecol Environ 3:479–86.

    Article  Google Scholar 

  • Elton CS. 1958. The Ecology of Invasions by Animals and Plants. Chicago, Illinois, USA: University of Chicago Press.

    Book  Google Scholar 

  • Engber EA, Varner JM. 2012. Patterns of flammability of the California oaks: the role of leaf traits. Can J For Res 42:1965–75.

    Article  Google Scholar 

  • Fonda RW. 2001. Burning characteristics of needles from eight pine species. For Sci 47:390–6.

    Google Scholar 

  • Forrestel AB, Moritz MA, Stephens SL. 2011. Landscape-scale vegetation change following fire in Point Reyes, California, USA. Fire Ecol 7:114–28.

    Article  Google Scholar 

  • Fosberg MA, Lancaster JW, Schroeder MJ. 1970. Fuel moisture response- drying relationships under standard and field conditions. For Sci 16:121–8.

    Google Scholar 

  • Foster DR, Clayden S, Orwig DA, Hall B, Barry S. 2002. Oak, chestnut and fire: climatic and cultural controls of long-term forest dynamics in New England, USA. J Biogeogr 29:1359–79.

    Article  Google Scholar 

  • Fraedrich SW, Harrington TC, Rabaglia RJ, Ulyshen MD, Mayfield Iii AE, Hanula JL, Eickwort JM, Miller DR. 2008. A fungal symbiont of the redbay ambrosia beetle causes a lethal wilt in redbay and other Lauraceae in the southeastern United States. Plant Dis 92:215–24.

    Article  CAS  PubMed  Google Scholar 

  • Guyette RP, Stambaugh MC, Dey DC, Muzika R-M. 2012. Predicting fire frequency with chemistry and climate. Ecosystems 15:322–35.

    Article  Google Scholar 

  • Hawkins TS. 2006. A forest transect of Pine Mountain, Kentucky: changes since E. Lucy Braun and chestnut blight. J Ky Acad Sci 67:73–80.

    Google Scholar 

  • Hessl AE, Saladyga T, Schuler T, Clark P, Wixom J. 2011. Fire history from three species on a central Appalachian ridgetop. Can J For Res 41:2031–9.

    Article  Google Scholar 

  • Jacobs D. 2007. Toward development of silvical strategies for forest restoration of American chestnut (Castanea dentata) using blight-resistant hybrids. Biol Conserv 137:497–506.

    Article  Google Scholar 

  • Jacobs DF, Dalgleish HJ, Nelson CD. 2013. A conceptual framework for restoration of threatened plants: the effective model of American chestnut (Castanea dentata) reintroduction. New Phytol 197:378–93.

    Article  Google Scholar 

  • Kane JM, Varner JM, Hiers JK. 2008. The burning characteristics of southeastern oaks: Discriminating fire facilitators from fire impeders. For Ecol Manag 256:2039–45.

    Article  Google Scholar 

  • Keever C. 1953. Present composition of some stands of the former oak-chestnut forest in the southern Blue Ridge Mountains. Ecology 34:44–54.

    Article  Google Scholar 

  • Kendra PE, Montgomery WS, Niogret J, Epsky ND. 2013. An Uncertain Future for American Lauraceae: A Lethal Threat from Redbay Ambrosia Beetle and Laurel Wilt Disease (A Review). Am J Plant Sci 04:727–38.

    Article  Google Scholar 

  • Knapp BO, Wang GG, Clark SL, Pile LS, Schlarbaum SE. 2014. Leaf physiology and morphology of Castanea dentata (Marsh.) Borkh., Castanea mollissima Blume, and three backcross breeding generations planted in the southern Appalachians. USA. New For 45:283–93.

    Article  Google Scholar 

  • Korstian CF, Stickel PW. 1927. The natural replacement of blight-killed chestnut in the hardwood forests of the northeast. J Agric Res 34:631–48.

    Google Scholar 

  • Kreye JK, Varner JM, Hamby GW, Kane JM. 2018. Mesophytic litter dampens flammability in fire-excluded pyrophytic oak-hickory woodlands. Ecosphere 9:e02078.

    Article  Google Scholar 

  • Kreye JK, Varner JM, Hiers JK, Mola J. 2013. Toward a mechanism for eastern North American forest mesophication: differential litter drying across 17 species. Ecol Appl 23:1976–86.

    Article  Google Scholar 

  • Kuljian H, Varner JM. 2013. Foliar consumption across a sudden oak death chronosequence in laboratory fires. Fire Ecol 9:33–44.

    Article  Google Scholar 

  • Lafon CW, Naito AT, Grissino-Mayer HD, Horn SP, Waldrop TA. 2017. Fire history of the Appalachian region: a review and synthesis. Asheville, NC: USDA Forest Service, Southern Research Station http://trace.tennessee.edu/utk_geogpubs/14/. Last accessed 31/07/2017.

  • Levine JM, D’Antonio CM. 2003. Forecasting biological invasions with increasing international trade. Conserv Biol 17:322–6.

    Article  Google Scholar 

  • Liebhold AM, MacDonald WL, Bergdahl D, Mastro VC. 1995. Invasion by exotic forest pests: a threat to forest ecosystems. For Sci Monogr 30:1–49.

    Google Scholar 

  • Lovett GM, Canham CD, Arthur MA, Weathers KC, Fitzhugh RD. 2006. Forest ecosystem responses to exotic pests and pathogens in eastern North America. BioScience 56:395–405.

    Article  Google Scholar 

  • Mauseth JD. 2000. Theoretical aspects of surface-to-volume ratios and water-storage capacities of succulent shoots. Am J Bot 87:1107–15.

    Article  CAS  PubMed  Google Scholar 

  • McCormick JF, Platt RB. 1980. Recovery of an Appalachian forest following the chestnut blight or Catherine Keever-you were right!. Am Midl Nat 104:264.

    Article  Google Scholar 

  • McEwan RW, Dyer JM, Pederson N. 2011. Multiple interacting ecosystem drivers: toward an encompassing hypothesis of oak forest dynamics across eastern North America. Ecography 34:244–56.

    Article  Google Scholar 

  • Meentemeyer RK, Cunniffe NJ, Cook AR, Filipe JAN, Hunter RD, Rizzo DM, Gilligan CA. 2011. Epidemiological modeling of invasion in heterogeneous landscapes: spread of sudden oak death in California (1990–2030). Ecosphere 2:art17.

  • Metz MR, Frangioso KM, Meentemeyer RK, Rizzo DM. 2011. Interacting disturbances: wildfire severity affected by stage of forest disease invasion. Ecol Appl 21:313–20.

    Article  PubMed  Google Scholar 

  • Metz MR, Varner JM, Frangioso KM, Meentemeyer RK, Rizzo DM. 2013. Unexpected redwood mortality from synergies between wildfire and an emerging infectious disease. Ecology 94:2152–9.

    Article  PubMed  Google Scholar 

  • Mola JM, Varner JM, Jules ES, Spector T. 2014. Altered community flammability in Florida’s Apalachicola Ravines and implications for the persistence of the endangered conifer Torreya taxifolia. Bond-Lamberty B, editor. PLoS ONE 9:e103933.

  • Muggeo VM. 2008. Segmented: an R package to fit regression models with broken-line relationships. R News 8:20–5.

    Google Scholar 

  • Murrill WA. 1906. A new chestnut disease. Torreya 6:186–9.

    Google Scholar 

  • Myers BR, Walck JL, Blum KE. 2004. Vegetation change in a former chestnut stand on the Cumberland Plateau of Tennessee during an 80-year period (1921–2000). Castanea 69:81–91.

    Article  Google Scholar 

  • Nelson RM, Hiers JK. 2008. The influence of fuelbed properties on moisture drying rates and timelags of longleaf pine litter. Can J For Res 38:2394–404.

    Article  Google Scholar 

  • Niinemets Ü, Valladares F. 2006. Tolerance to shade, drought, and waterlogging of temperate Northern Hemisphere trees and shrubs. Ecol Monogr 76:521–47.

    Article  Google Scholar 

  • Nowacki GJ, Abrams MD. 2008. The demise of fire and “mesophication” of forests in the eastern United States. BioScience 58:123–38.

    Article  Google Scholar 

  • Oksanen J, Blanchet G, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens HH, Szoecs E, Wagner H. 2017. Vegan: Community Ecology Package. R package version 2.4-4. https://cran.r-project.org/package=vegan.

  • Paillet FL. 2002. Chestnut: history and ecology of a transformed species. J Biogeogr 29:1517–30.

    Article  Google Scholar 

  • Pinchot C, Clark S, Schlarbaum S, Saxton A, Sung S-J, Hebard F. 2015. Effects of temporal dynamics, nut weight and nut size on growth of American chestnut, Chinese chestnut and backcross generations in a commercial nursery. Forests 6:1537–56.

    Article  Google Scholar 

  • R Development Core Team. 2017. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.

    Google Scholar 

  • Rhoades C, Loftis D, Lewis J, Clark S. 2009. The influence of silvicultural treatments and site conditions on American chestnut (Castanea dentata) seedling establishment in eastern Kentucky, USA. For Ecol Manag 258:1211–18.

    Article  Google Scholar 

  • Rizzo DM, Garbelotto M. 2003. Sudden oak death: endangering California and Oregon forest ecosystems. Front Ecol Environ 1:197–204.

    Article  Google Scholar 

  • Rothermel RC. 1983. How to predict the spread and intensity of forest and range fires. Ogden, UT: USDA Forest Service Intermountain Forest and Range Experiment Station.

    Book  Google Scholar 

  • Saterberg T, Sellman S, Ebenman B. 2013. High frequency of functional extinctions in ecological networks. Nature 499:468–70.

    Article  CAS  PubMed  Google Scholar 

  • Scarff FR, Westoby M. 2006. Leaf litter flammability in some semi-arid Australian woodlands. Funct Ecol 20:745–52.

    Article  Google Scholar 

  • Schweitzer JA, Bailey JK, Rehill BJ, Martinsen GD, Hart SC, Lindroth RL, Keim P, Whitham TG. 2004. Genetically based trait in a dominant tree affects ecosystem processes: Plant genetics impact ecosystems. Ecol Lett 7:127–34.

    Article  Google Scholar 

  • Shapiro B. 2017. Pathways to de-extinction: how close can we get to resurrection of an extinct species? Seddon P, editor. Funct Ecol 31:996–1002.

  • Sherkow JS, Greely HT. 2013. What if extinction is not forever? Science 340:32–3.

    Article  CAS  PubMed  Google Scholar 

  • Sniezko RA. 2006. Resistance breeding against nonnative pathogens in forest trees—current successes in North America. Can J Plant Pathol 28:S270–9.

    Article  Google Scholar 

  • Stambaugh MC, Varner JM, Noss RF, Dey DC, Christensen NL, Baldwin RF, Guyette RP, Hanberry BB, Harper CA, Lindblom SG, Waldrop TA. 2015. Clarifying the role of fire in the deciduous forests of eastern North America: reply to Matlack: Fire in Deciduous Forests. Conserv Biol 29:942–6.

    Article  PubMed  Google Scholar 

  • Steiner KC, Carlson JE, editors. 2006. Restoration of American chestnut to forest lands: Proceedings of a conference and workshop. In: Restoration of American chestnut to forest lands: proceedings of a conference and workshop. Washington DC, USA: National Park Service http://sfr.psu.edu/public/chestnut/information/conference-2004/conference/conference-proceedings.

  • Steiner KC, Westbrook JW, Hebard FV, Georgi LL, Powell WA, Fitzsimmons SF. 2017. Rescue of American chestnut with extraspecific genes following its destruction by a naturalized pathogen. New For 48:317–36.

    Article  Google Scholar 

  • Sturrock RN, Frankel SJ, Brown AV, Hennon PE, Kliejunas JT, Lewis KJ, Worrall JJ, Woods AJ. 2011. Climate change and forest diseases: Climate change and forest diseases. Plant Pathol 60:133–49.

    Article  Google Scholar 

  • Thomas-Van Gundy M, Bard J, Kochenderfer J, Berrang P. 2017. Mortality, early growth, and blight occurrence in hybrid, Chinese, and American chestnut seedlings in West Virginia. In: Proceedings 20th Central Hardwood Forest Conference. Vol. GTR NRS-P-167. Columbia, MO, USA: USDA Forest Service, Northern Research Station. pp 222–39.

  • Varner JM, Arthur MA, Clark SL, Dey DC, Hart JL, Schweitzer CJ. 2016. Fire in eastern North American oak ecosystems: filling the gaps. Fire Ecol 12:1–5.

    Article  Google Scholar 

  • Varner JM, Kane JM, Banwell EM, Kreye JK. 2015a. Flammability of litter from southeastern trees: A preliminary assessment. In: Holley GA, Connor KF, Haywood JD, editors. Proceedings of the 17th Southern Silvicultural Research Conference. Vol. eGTR-SRS-203. Ashville, NC: USDA Forest Service, Southern Research Station. pp 183–7.

  • Varner JM, Kane JM, Kreye JK, Engber E. 2015b. The flammability of forest and woodland litter: a synthesis. Curr For Rep 1:91–9.

    Google Scholar 

  • Varner JM, Kuljian HG, Kreye JK. 2017. Fires without tanoak: the effects of a non-native disease on future community flammability. Biol Invasions 19:2307–17.

    Article  Google Scholar 

  • Veldman JW, Mattingly WB, Brudvig LA. 2013. Understory plant communities and the functional distinction between savanna trees, forest trees, and pines. Ecology 94:424–34.

    Article  PubMed  Google Scholar 

  • Wang GG, Bauerle WL, Mudder BT. 2006. Effects of light acclimation on the photosynthesis, growth, and biomass allocation in American chestnut (Castanea dentata) seedlings. For Ecol Manag 226:173–80.

    Article  Google Scholar 

  • Wang GG, Knapp BO, Clark SL, Mudder BT. 2013. The silvics of Castanea dentata (Marsh.) Borkh., American chestnut, Fagaceae (beech family). Asheville, NC, USA: USDA Forest Service Southern Research Station https://www.researchgate.net/profile/Stacy_Clark3/publication/260779839_The_Silvics_of_Castanea_dentata_Marsh_Borkh_American_Chestnut_Fagaceae_Beech_Family/links/54806fa40cf2ccc7f8bd0477.pdf. Last accessed 24/08/2017.

  • Weed AS, Ayres MP, Hicke JA. 2013. Consequences of climate change for biotic disturbances in North American forests. Ecol Monogr 83:441–70.

    Article  Google Scholar 

  • Whitham TG, Bailey JK, Schweitzer JA, Shuster SM, Bangert RK, LeRoy CJ, Lonsdorf EV, Allan GJ, DiFazio SP, Potts BM, Fischer DG, Gehring CA, Lindroth RL, Marks JC, Hart SC, Wimp GM, Wooley SC. 2006. A framework for community and ecosystem genetics: from genes to ecosystems. Nat Rev Genet 7:510–23.

    Article  CAS  PubMed  Google Scholar 

  • Whitham TG, Young WP, Martinsen GD, Gehring CA, Schweitzer JA, Shuster SM, Wimp GM, Fischer DG, Bailey JK, Lindroth RL, Woolbright S, Kuske CR. 2003. Community and ecosystem genetics: a consequence of the extended phenotype. Ecology 84:559–73.

    Article  Google Scholar 

  • Youngs RL. 2000. A right smart little jolt: loss of the chestnut and a way of life. J For 98:17–21.

    Google Scholar 

Download references

Acknowledgements

Laboratory burning and moisture trials were assisted by L. Rios, A. Cowen, K. Dedrick, T. Bradley, and J. Avitia. Field collections of samples and leaf scans were conducted by S. Greenler. Discussions with J. Kreye and D. Orwig were helpful in advancing our thoughts on the topic. Two anonymous reviewers also provided helpful suggestions and comments to an earlier draft. Financial support for this research was provided by the USDA National Institute of Food and Agriculture, McIntire-Stennis Cooperative Forestry Research Program at Humboldt State University (Project #CALZ-157) and Purdue University (IND011557MS), the National Fire Plan, and Hardwood Tree Improvement and Regeneration Center.

Author information

Authors and Affiliations

  1. Department of Forestry and Wildland Resources, Wildland Fire Laboratory, Humboldt State University, 1 Harpst Street, Arcata, California, 95521, USA

    Jeffrey M. Kane

  2. Pacific Wildland Fire Sciences Laboratory, United States Department of Agriculture, Forest Service, 400 N 34th Street, Suite 201, Seattle, Washington, 98103, USA

    J. Morgan Varner

  3. Department of Forestry and Natural Resources, Purdue University, 715 W State Street, West Lafayette, Indiana, 47907, USA

    Michael R. Saunders

Authors
  1. Jeffrey M. Kane
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. J. Morgan Varner
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. Michael R. Saunders
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Jeffrey M. Kane.

Additional information

Author Contributions

JMK, JMV, and MRS all contributed to the study concept and design and writing of the manuscript, JMK and MRS performed the research, and JMK analyzed the data.

Data will be hosted at the Humboldt State University data sharing repository.

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Kane, J.M., Varner, J.M. & Saunders, M.R. Resurrecting the Lost Flames of American Chestnut. Ecosystems 22, 995–1006 (2019). https://doi.org/10.1007/s10021-018-0318-1

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10021-018-0318-1

Keywords

  • fire ecology
  • disease resistance
  • global change
  • litter flammability
  • mesophication
  • restoration
  • resurrection ecology