New Forests

, Volume 48, Issue 4, pp 491–512 | Cite as

Growth, survival, and competitive ability of chestnut (Castanea Mill.) seedlings planted across a gradient of light levels

  • Cornelia C. Pinchot
  • Scott E. Schlarbaum
  • Stacy L. Clark
  • Arnold M. Saxton
  • Ami M. Sharp
  • Callie J. Schweitzer
  • Frederick V. Hebard


There has been an increased interest in tree breeding for resistance to exotic pests and pathogens, however relatively little research has focused on the reintroduction of these tree species. Understanding the durability of resistance in field settings and the field performance of improved trees is critical for successful species reintroduction. To evaluate methods for reintroducing American chestnut [Castanea dentata (Marsh.) Borkh] to managed forests on the Cumberland Plateau, we quantified four-year survival and growth and three-year competitive ability of chestnut seedlings planted on the Daniel Boone National Forest in southeastern Kentucky, USA. We used a split-plot design to compare chestnut response among three silvicultural treatments spanning a gradient of light levels; midstory removal, thinning, and shelterwood with reserves (2, 24, and 65% available photosynthetically active radiation, respectively) and three chestnut breeding types; American, Chinese (C. mollissima Blume.), and BC2F3 hybrid. One of two hybrid families planted had similar survival to American chestnuts, 21 and 27% survival, respectively, while the other had better survival, 57%. Chinese chestnut survival was better than the other breeding generations (90%). High mortality among American and hybrid chestnut seedlings was likely caused by infection from Phytophthora cinnamomi Rands. Incidence of blight infection was low. While chestnut seedling growth was greatest in the high-light treatment, competitive ability of chestnut, evaluated by comparing planted seedling height to height of understory competitors, was maximized in the intermediate light treatment. These results demonstrate the importance of evaluating competition pressure from co-occurring vegetation and field performance of resistant genotypes when assessing methods for reintroducing tree species to forested settings.


Castanea Species restoration Silvicultural treatment Competitive ability 



We thank Tracy Powers and Dave Griffin, University of Tennessee Tree Improvement Program, for assistance with study establishment and field measurements. Thank you to Dr. Jennifer Franklin and Dr. David Buckley, Department of Forestry, Wildlife and Fisheries, University of Tennessee, for guidance in methods development. We are grateful for assistance and guidance from the Cold Hill Ranger District of the Daniel Boone National Forest, particularly for help from Robbie Sitzlar, Silviculturist. Thanks to Dr. Steve Jeffers and Inga Meadows, College of Agriculture, Forestry and Life Sciences, Clemson University, for Phytophthora cinnamomi testing. Dr. Sandra Anagnostakis, Dr. Melissa Thomas Van-Gundy, and two anonymous reviewers provided excellent feedback on early drafts of this paper.


This work was supported by the USDA Forest Service, Northern Research Station, Southern Research Station and by a Joint Venture Agreement between the University of Tennessee and the USDA Forest Service, Southern Research Station [10-JV-11330134-066].

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

11056_2017_9577_MOESM1_ESM.docx (19 kb)
Supplementary material 1 (DOCX 19 kb)


  1. Anagnostakis SL (2001) The effect of multiple importations of pests and pathogens on a native tree. Biol Invasions 3:245–254CrossRefGoogle Scholar
  2. Anagnostakis SL (2006) A history of Phytophthora cinnamomi in the United States. Aust Nutgrow 20:36–38Google Scholar
  3. Anagnostakis SL (2012) Chestnut breeding in the United States for disease and insect resistance. Plant Dis 96:1392–1403CrossRefGoogle Scholar
  4. Ashe WW (1911) Chestnut in Tennessee. Tennessee Geological Survey Bulletin 10-B. Baird-Ward Printing Company, NashvilleGoogle Scholar
  5. 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 329:21–29. doi: 10.1016/j.foreco.2014.06.011 CrossRefGoogle Scholar
  6. Boring LR, Monk CD, Swank WT (1981) Early regeneration of a clear-cut southern Appalachian forest. Ecology 62(5):1244–1253. doi: 10.2307/1937289 CrossRefGoogle Scholar
  7. Boyd IL, Freer-Smith PH, Gilligan CA, Godfray HC (2013) The consequence of tree pests and diseases for ecosystem services. Science 342(6160):1235773. doi: 10.1126/science.1235773 CrossRefPubMedGoogle Scholar
  8. Braun L (1950) Deciduous forests of eastern North America. Hafner, New YorkGoogle Scholar
  9. Brown CE, Bailey BG, Saunders MR, Jacobs DF (2014) Effects of root competition on development of chestnut and oak regeneration following midstory removal. Forestry 87(4):562–570. doi: 10.1093/forestry/cpu014 CrossRefGoogle Scholar
  10. Buckley D, Isebrands JG, Sharik TL (1999) Practical field methods of estimating canopy cover, PAR, and LAI in Michigan oak and pine stands. North J Appl For 16(1):25–32Google Scholar
  11. Burnham CR (1988) The restoration of the American chestnut: mendelian genetics may solve a problem that has resisted other approaches. Am Sci 76(5):478–487Google Scholar
  12. Campbell FT, Schlarbaum SE (1994) Fading forests: North American trees and the threat of exotic pests. Natural Resources Defense Council Report. Accessed 24 Jan 2017
  13. Campbell FT, Schlarbaum SE (2002) Fading forests II. Trading away North America’s natural heritage? Healing Stones Foundation Publication. Accessed 24 Jan 2017
  14. Campbell FT, Schlarbaum SE (2014) Fading forests III: American forests. What choice will we make? The Nature Conservancy and The University of Tennessee. Accessed 24 Jan 2017
  15. 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(4):1017–1033. doi: 10.3390/f3041017 CrossRefGoogle Scholar
  16. Clark SL, Schlarbaum SE, Pinchot CC, Anagnostakis SL, Saunders MR, Thomas-Van Gundy M, Schaberg PG, McKenna J, Bard J, Berrang P, Casey DM, Casey CE, Crane B, Jackson B, Kochenderfer J, Lewis R, MacFarlane R, Makowski R, Miller M, Rodrigue J, Stelock J, Thornton C, Williamson T (2014) Reintroduction of American Chestnut in the National Forest System. J For 112(5):501–512Google Scholar
  17. 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(2):243–270. doi: 10.1007/s11056-015-9512-6 CrossRefGoogle Scholar
  18. Crandall BS, Gravatt GF, Ryan MM (1945) Root disease of Castanea species and some coniferous and broadleaf nursery stocks caused by Phytophthora cinnamomi. Phytopathology 35:162–180Google Scholar
  19. Decagon Devices, Inc. (2007) AccuPAR Linear PAR/LAI ceptometer, Model PAR-80. Operator’s Manual, Version 3.4. Pullman, WAGoogle Scholar
  20. Dey DC, Parker WC (1997) Overstory density affects field performance of underplanted red oak (Quercus rubra L.) in Ontario. North J Appl For 14(3):120–125. doi: 10.1093/treephys/28.5.797 Google Scholar
  21. Diamond SJ, Giles RH, Kirkpatrick RL, Griffin GJ (2000) Hard mast production before and after the chestnut blight. South J Appl For 24(4):196–201Google Scholar
  22. Emerson GB (1846) A report on the trees and shrubs growing naturally in the forests of Massachusetts. Little, Brown and Company, BostonGoogle Scholar
  23. Frothingham EH (1924) Some silvicultural aspects of the chestnut blight situation. J For 22(8):861–872Google Scholar
  24. Gauthier MM, Zellers KE, Lof M, Jacobs DF (2013) Inter- and intra-specific competitiveness of plantation-grown American chestnut (Castanea dentata). For Ecol Manag 291:289–299. doi: 10.1016/j.foreco.2012.11.014 CrossRefGoogle Scholar
  25. Gingrich SF (1967) Measuring and evaluating stocking and stand density in upland hardwood forests in the Central States. For Sci 13(1):38–53Google Scholar
  26. Griffin GJ (1989) Incidence of chestnut blight and survival of American chestnut in forest clear-cut and neighboring understory sites. Plant Dis 73:123–127. doi: 10.1094/pd-73-0123 CrossRefGoogle Scholar
  27. Griffin GJ, Hebard FV, Wendt RW, Elkins JR (1983) Survival of American chestnut trees: evaluation of blight resistance and hypovirulence in Endothia parasitica. Phytopathology 73(7):1084–1092. doi: 10.1094/phyto-73-1084 CrossRefGoogle Scholar
  28. Griffin GJ, Smith HC, Dietz A, Elkins JR (1991) Importance of hardwood competition to American chestnut survival, growth, and blight development in forest clearcuts. Can J Bot 69(8):1804–1809. doi: 10.1139/b91-229 CrossRefGoogle Scholar
  29. Hawley RC, Hawes AF (1912) Forestry in New England: manual of forestry for the northeastern United States, vol 1, 2nd edn. Wiley, New YorkGoogle Scholar
  30. Hebard FV (2001) Backcross breeding program produces blight-resistant American chestnuts. Ecol Restor 19:252–254CrossRefGoogle Scholar
  31. Hebard FV (2012) The American Chestnut Foundation Breeding Program. In: Proceedings of the 4th international workshop on the genetics of host-parasite interactions in forestry. July 2012. Edited by Sniezko RA, Yanchuk AD, Kliejunas JT, Palmieri KM, Alexander JM, Frankel SJ. USDA Forest Service, Albany, California, Gen Tech Rep PSW-GTR-240, pp 221–234Google Scholar
  32. Hosmer DW Jr, Lemeshow S (2004) Applied logistic regression. Wiley, HobokenGoogle Scholar
  33. Jacobs DF (2007) Toward development of silvical strategies for forest restoration of American chestnut (Castanea dentata) using blight-resistant hybrids. Biol Conserv 137(4):497–506. doi: 10.1016/j.biocon.2007.03.013 CrossRefGoogle Scholar
  34. Jacobs DF, Severeid LR (2004) Dominance of interplanted American chestnut (Castanea dentata) in southwestern Wisconsin, USA. For Ecol Manag 191(1):111–120. doi: 10.1016/j.foreco.2003.11.015 CrossRefGoogle Scholar
  35. 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(2):378–393. doi: 10.1111/nph.12020 CrossRefPubMedGoogle Scholar
  36. Jeffers SN, James JB, Sisco PH (2009) Screening for resistance to Phytophthora cinnamomi in hybrid seedlings of American chestnut. In: Proceedings of the fourth meeting of the International Union of Forest Research Organizations (IUFRO) Working Party 7.02.09, Phytophthora in Forests and Natural Ecosystems. August 2007. Edited by Gohee EM, Frankel SJ. USDA Forest Service, Albany, California, Gen Tech Rep PSW-GTR-221, pp 188–194Google Scholar
  37. Joesting HM, McCarthy BC, Brown KJ (2009) Determining the shade tolerance of American chestnut using morphological and physiological leaf parameters. For Ecol Manag 257(1):280–286. doi: 10.1016/j.foreco.2008.09.009 CrossRefGoogle Scholar
  38. Johnson PS (1971) Growth and survival of interplanted hardwoods in southern Wisconsin oak clearcuttings. USDA Forest Service Research Note NC-11, St. Paul, MinnesotaGoogle Scholar
  39. Johnson PS (1976) Eight-year performance of interplanted hardwoods in southern Wisconsin oak clearcuts. USDA Forest Service Research Paper NC-126, St. Paul, MinnesotaGoogle Scholar
  40. Johnson PS (1984) Responses of planted northern red oak to three overstory treatments. Can J For Res 14(4):536–542. doi: 10.1139/x84-099 CrossRefGoogle Scholar
  41. Klinka K, Wang Q, Kayahara GJ (1992) Light-growth response relationships in Pacific silver fir (Aibes amabilis) and subalpine fir (Abies lasiocarpa). Can J Bot 70(10):1919–1930. doi: 10.1139/b92-239 CrossRefGoogle Scholar
  42. Kormanik PP, Sung SS, Kormanik TL (1994) Toward a single nursery protocol for oak seedlings. In: Proceedings of the 22nd southern forest tree improvement conference. June 1993. Edited by Lantz CW, Moorhead DJ. USDA Forest Service, Atlanta, Georgia, pp 89–98Google Scholar
  43. Kormanik PP, Sung SS, Kas DJ, Schlarbaum SE (1997) Effect of seedling size and first order lateral roots on early development of northern red oak on mesic sites. In: Proceedings of the ninth biennial southern silviculture research conference. February 1997. Edited by Waldrop TA. USDA Forest Service, Asheville, North Carolina, Gen Tech Rep SRS-48, pp 332–337Google Scholar
  44. Latham RE (1992) Co-occurring tree species change rank in seedling performance with resources varied experimentally. Ecology 73(6):2129–2144. doi: 10.2307/1941461 CrossRefGoogle Scholar
  45. Lemmon PE (1956) A spherical densiometer for estimating forest overstory density. For Sci 2(4):314–320Google Scholar
  46. Lhotka JM, Loewenstein EF (2006) Indirect measures for characterizing light along a gradient of mixed-hardwood riparian forest canopy structures. For Ecol Manag 226(1):310–318. doi: 10.1016/j.foreco.2006.01.043 CrossRefGoogle Scholar
  47. Loftis DL (1990a) A shelterwood method for regenerating red oak in the Southern Appalachians. For Sci 36(4):917–929Google Scholar
  48. Loftis DL (1990b) Predicting post-harvest performance of advance red oak reproduction in the southern Appalachians. For Sci 36(4):908–916Google Scholar
  49. Mattoon FE (1909) The origin and early development of chestnut sprouts. For Q 7(1):34–37Google Scholar
  50. Maunder M (1992) Plant reintroduction: an overview. Biodivers Conserv 1(1):51–61. doi: 10.1007/bf00700250 CrossRefGoogle Scholar
  51. McCament CL, McCarthy BC (2005) Two-year response of American chestnut (Castanea dentata) seedlings to shelterwood harvesting and fire in mixed-oak forest ecosystem. Can J For Res 35(3):740–749. doi: 10.1139/x05-002 CrossRefGoogle Scholar
  52. McGee CE, Loftis DL (1986) Planted oaks perform poorly in North Carolina and Tennessee. North J Appl For 3(3):114–115Google Scholar
  53. McNab HW (2003) Early results from a pilot test of American chestnut seedlings under a forest canopy. Am Chestnut Found J 16:32–41Google Scholar
  54. Meadows IM, Zwart DC, Jeffers SN, Waldrop TA, Bridges WC Jr (2011) Effects of fuel reduction treatments on incidence of Phytophthora species in soil of a southern Appalachian Mountain forest. Plant Dis 5(7):811–820. doi: 10.1094/pdis-07-10-0505 CrossRefGoogle Scholar
  55. Murrill WA (1906) A serious chestnut disease. J N Y Bot Gard 7:143–153Google Scholar
  56. Novoselov VS (1960) A closed volumeter for plant root systems. Fiziol Rast 7:243–244Google Scholar
  57. Oldfield SF (2009) Botanic gardens and the conservation of tree species. Trends Plant Sci 14(11):581–583. doi: 10.1016/j.tplants.2009.08.013 CrossRefPubMedGoogle Scholar
  58. Ostry ME, Moore M (2008) Response of butternut selections to inoculation with Sirococcus clavigignenti-juglandacearum. Plant Dis 92(9):1336–1338. doi: 10.1094/pdis-92-9-1336 CrossRefGoogle Scholar
  59. Paillet FL (1984) Growth-form and ecology of American chestnut sprout clones in northeastern Massachusetts. Bull Torrey Bot Club 111:316–328. doi: 10.2307/2995913 CrossRefGoogle Scholar
  60. Paquette A, Bouchard A, Cogliastro A (2006) Survival and growth of under-planted trees: a meta-analysis across four biomes. Ecol Appl 16(4):1575–1589. doi:10.1890/1051-0761(2006)016[1575:SAGOUT]2.0.CO;2 CrossRefPubMedGoogle Scholar
  61. Parent S, Messier C (1996) A simple and efficient method to estimate microsite light availability under a forest canopy. Can J For Res 26(1):151–154. doi: 10.1139/x26-017 CrossRefGoogle Scholar
  62. Rhoades CC, Brosi SL, Dattilo AJ, Vincelli P (2003) Effect of soil compaction and moisture on incidence of phytophthora root rot on American chestnut (Castanea dentata) seedlings. For Ecol Manag 184(1):47–54. doi: 10.1016/S0378-1127(03)00147-6 CrossRefGoogle Scholar
  63. Rhoades C, Loftis D, Lewis J, Clark SL (2009) The influence of silviculture treatments and site conditions on American chestnut (Castanea dentata) seedling establishment in eastern Kentucky, USA. For Ecol Manag 258(7):1211–1218. doi: 10.1016/j.foreco.2009.06.014 CrossRefGoogle Scholar
  64. Santini A, La Porta N, Ghelardini L, Mittempergher L (2008) Breeding against Dutch elm disease adapted to the Mediterranean climate. Euphytica 163(1):45–56. doi: 10.1007/s10681-007-9573-5 CrossRefGoogle Scholar
  65. Santini A, Ghelardini L, De Pace C, Desprez-Loustau ML, Capretti P, Chandelier A, Cech T, Chira D, Diamandis S, Gaitniekis T, Hantula J, Holdenrieder O, Jankovsky L, Jung T, Jurc D, Kirisits T, Kunca A, Lygis V, Malecka M, Marcais B, Schmitz S, Schumacher J, Solheim H, Solla A, Szabό I, Tsopelas P, Vannini A, Vettraino AM, Webber J, Woodward S, Stenlid J (2013) Biogeographical patterns and determinants of invasion by forest pathogens in Europe. New Phytol 197(1):238–250. doi: 10.1111/j.1469-8137.2012.04364.x CrossRefPubMedGoogle Scholar
  66. SAS Institute Inc. (2011) SAS/STAT© 9.3 User’s Guide, Version 9.3. SAS Institute, Inc., Cary, North CarolinaGoogle Scholar
  67. Schenck CA (1912) The art of the second growth of American sylviculture, 3rd revised edn. Brandow Printing Co., AlbanyCrossRefGoogle Scholar
  68. Schlarbaum SE (1990) Returning the American chestnut to eastern North America. In: McGee CE (ed) Proceedings of the 1989 southern Appalachian mast management workshop. August 1989. United States Forest Service, Washington, DC, pp 66–70Google Scholar
  69. Schuler JL, Robison DJ (2010) Performance of northern red oak enrichment plantings in naturally regenerating southern Appalachian hardwood stands. New For 40(1):119–130. doi: 10.1007/s11056-009-9187-y CrossRefGoogle Scholar
  70. Schweitzer CJ, Clark SL, Gottschalk KW, Stringer JW, Sitzlar RL (2014) Proactive Restoration: planning, implementation, and early results of silvicultural strategies for increasing resilience against gypsy moth infestation in upland oak forests on the Daniel Boone National Forest, Kentucky. J For 112(5):401–411. doi: 10.5849/jof.13-085 Google Scholar
  71. Seddon PJ (2010) From reintroduction to assisted colonization: moving along the conservation translocation spectrum. Restor Ecol 18(6):796–802. doi: 10.1111/j.1526-100X.2010.00724.x CrossRefGoogle Scholar
  72. Smith NJ (1991) Specific leaf area: clues to how salal (Gualtheria shallon Pursh) responds to stand conditions. Can J For Res 21(3):300–305. doi: 10.1139/x91-037 CrossRefGoogle Scholar
  73. Sniezko RA (2006) Resistance breeding against nonnative pathogens in forest tree—current successes in North America. Can J Plant Pathol 28(S1):S270–S279. doi: 10.1080/07060660609507384 CrossRefGoogle Scholar
  74. Spetich MA, Dey DC, Johnson PS, Graney DL (2002) Competitive capacity of Quercus rubra L. planted in Arkansas’ Boston Mountains. For Sci 48(3):504–517Google Scholar
  75. 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. doi: 10.1007/s11056-016-9561-5 Google Scholar
  76. Teclaw RM, Isebrands JG (1993) An artificial regeneration system for establishing northern red oak on dry-mesic sites in the Lake States, USA. Ann Des Sci For 50(6):543–552. doi: 10.1051/forest:19930603 CrossRefGoogle Scholar
  77. Thompson LM, Van Manen FT, Schlarbaum SE, DePoy M (2006) A spatial modeling approach to identify potential butternut restoration sites in Mammoth Cave National Park. Restor Ecol 14(2):289–296. doi: 10.1111/j.1526-100x.2006.00131.x CrossRefGoogle Scholar
  78. Townsend AM, Douglass LW (2001) Variation among American elm clones in long-term dieback, growth, and survival following Ophiostoma inoculation. J Environ Hortic 19:100–103Google Scholar
  79. Townsend AM, Bentz SE, Douglass LW (2005) Evaluation of 19 American elm clones for tolerance to Dutch elm disease. J Environ Hortic 23:21–24Google Scholar
  80. 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(1):173–180. doi: 10.1016/j.foreco.2005.12.063 CrossRefGoogle Scholar
  81. Wang GG, Knapp BO, Clark SL, Mudder BT (2013) The Silvics of Castanea dentata (Marsh.) Borkh., American Chestnut, Fagaceae (Beech Family). USDA Forest Service Gen Tech Rep SRS-GTR-173, Asheville, North CarolinaGoogle Scholar
  82. Wendel GW (1980) Growth and survival of planted northern red oak seedlings in West Virginia. South J Appl For 4(1):49–54Google Scholar
  83. Zaczek JJ, Steiner KC, Bowersox TW (1997) Northern red oak planting stock: 6-year results. New For 13(1–3):177–191CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media Dordrecht (outside the USA) 2017

Authors and Affiliations

  • Cornelia C. Pinchot
    • 1
  • Scott E. Schlarbaum
    • 2
  • Stacy L. Clark
    • 3
  • Arnold M. Saxton
    • 4
  • Ami M. Sharp
    • 2
  • Callie J. Schweitzer
    • 5
  • Frederick V. Hebard
    • 6
  1. 1.US Department of Agriculture Forest Service, Northern Research StationDelawareUSA
  2. 2.Department of Forestry, Wildlife and FisheriesThe University of TennesseeKnoxvilleUSA
  3. 3.US Department of Agriculture Forest Service, Southern Research StationKnoxvilleUSA
  4. 4.Department of Animal ScienceThe University of TennesseeKnoxvilleUSA
  5. 5.US Department of Agriculture Forest Service, Southern Research StationHuntsvilleUSA
  6. 6.The American Chestnut FoundationMeadowviewUSA

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