Silviculture for Eastern Old Growth in the Context of Global Change

  • William S. Keeton
  • Craig G. Lorimer
  • Brian J. Palik
  • Frédérik Doyon


When management for old-growth characteristics in eastern forests first began to be discussed in the late twentieth century, there was skepticism from some quarters as to whether it was a desirable or even a feasible idea. Old growth will recover on its own. Why not just let nature take its course? There were also those who saw little value in managing for old-growth features, perceiving this as a threat to more traditional management objectives (Puettmann et al. 2015). Since that time, concepts of managing for stand structural complexity, in ways that encourage some characteristics of old-growth forests, have caught on in a variety of contexts (Bauhus et al. 2009; Puettmann et al. 2009). In many ways this shift mirrors how the profession has grown to embrace multifunctional forestry broadly defined (Gustafsson et al. 2012). Old-growth silviculture increasingly has a place within this framework, filling the niche of enhancing the representation of late successional forests on landscapes where they are now vastly underrepresented relative to their abundance on landscapes prior to Euro-American settlement (Lorimer and White 2003; Rhemtulla et al. 2007). The working hypothesis is that this type of management will contribute to sustainable forest practices focused on providing a broad array of ecosystem goods and services, including those associated with late successional systems. And in recent decades there has been increasing interest in old-growth restoration more narrowly and management for older forest characteristics in working forests generally, both in terms of experimental research (e.g., Keeton 2006; Gronewold et al. 2010; Forrester et al. 2013; Palik et al. 2014) and practical applications (Hagenbuch et al. 2013; Fassnacht et al. 2015).


  1. Angers, V. A., C. Messier, M. Beaudet, and A. Leduc. 2005. “Comparing composition and structure in old-growth and harvested (selection and diameter-limit cuts) northern hardwood stands in Quebec” Forest Ecology and Management217: 275–293.CrossRefGoogle Scholar
  2. Aubin, I., A. D. Munson, F. Cardou, P. J. Burton, N. Isabel, J. H. Pedlar, and C. Messier. 2016. “Traits to stay, traits to move: a review of functional traits to assess sensitivity and adaptive capacity of temperate and boreal trees to climate change” Environmental Reviews24: 164–186.CrossRefGoogle Scholar
  3. Bauhus, J., K. Puettmann, and C. Messier. 2009. “Silviculture for old-growth attributes” Forest Ecology and Management4: 525–537.CrossRefGoogle Scholar
  4. Boal, C. W., D. E. Andersen, and P. L. Kennedy. 2005. “Foraging and nesting habitat of breeding male northern goshawks in the Laurentian mixed forest province, Minnesota” Journal of Wildlife Management69: 1516–1527.CrossRefGoogle Scholar
  5. Bolton, N. W., and A. W. D’Amato 2011. “Regeneration responses to gap size and coarse woody debris within natural disturbance-based silvicultural systems in northeastern Minnesota, USA” Forest Ecology and Management262: 1215–1222.Google Scholar
  6. Bormann, F. H., and G. E. Likens. 1979. Pattern and Process in a Forested Ecosystem. New York: Springer Verlag.CrossRefGoogle Scholar
  7. Brang, P., P. Spathelf, J. B. Larsen, J. Bauhus, A. Bonccˇìna, C. Chauvin, L. Drössler, et al. 2014. “Suitability of close-to-nature silviculture for adapting temperate European forests to climate change” Forestry87: 492–503.CrossRefGoogle Scholar
  8. Burrascano, S., W. S. Keeton, F. M. Sabatini, and C. Blasi. 2013. “Commonality and variability in the structural attributes of moist temperate old-growth forests: A global review” Forest Ecology and Management291: 458–479.CrossRefGoogle Scholar
  9. Choi, J., C. G. Lorimer, and J. M. Vanderwerker. 2007. “A simulation of the development and restoration of old-growth structural features in northern hardwoods” Forest Ecology and Management249: 204–220.CrossRefGoogle Scholar
  10. Cogbill, C., J. Burk, and G. Motzkin. 2003. “The forests of presettlement New England, USA: spatial and compositional patterns based on town proprietor surveys” Journal of Biogeography29: 1279–1304.CrossRefGoogle Scholar
  11. Crow, T. R., R. D. Jacobs, R. R. Oberg, and C. H. Tubbs. 1981. “Stocking and structure for maximum growth in sugar maple selection stands.” Research Paper NC-199. North Central Forest Experiment Station. St. Paul, MN: USDA Forest Service.Google Scholar
  12. Curzon, M. T., and W. S. Keeton. 2010. “Spatial characteristics of canopy disturbances in riparian old-growth hemlock-northern hardwood forests, Adirondack Mountains, New York, USA” Canadian Journal of Forest Research40: 13–25.CrossRefGoogle Scholar
  13. D’Amato, A. W., J. B. Bradford, S. Fraver, and B. J. Palik. 2011. “Forest management for mitigation and adaptation to climate change: insights from long-term silviculture experiments” Forest Ecology and Management262: 803–816.CrossRefGoogle Scholar
  14. D’Amato, A. W., P. F. Catanzaro, and L. S. Fletcher. 2015. “Early regeneration and structural responses to patch selection and structural retention in second-growth northern hardwoods” Forest Science61: 183–189.CrossRefGoogle Scholar
  15. D’Amato, A. W., and D. A. Orwig. 2008. “Stand and landscape-level disturbance dynamics in old-growth forests in western Massachusetts” Ecological Monographs78: 507–522.CrossRefGoogle Scholar
  16. DeGraaf, R. M., and M. Yamasaki. 2001. New England Wildlife. Hanover and Lebanon, NH: University Press of New England.Google Scholar
  17. Della-Bianca, L., and D. E. Beck. 1985. “Selection management in southern Appalachian hardwoods” Southern Journal of Applied Forestry9: 191–196.Google Scholar
  18. Després, T., H. Asselin, F. Doyon, and Y. Bergeron. 2014. “Structural and spatial characteristics of old-growth temperate deciduous forests at their northern distribution limit” Forest Science60: 871–880.CrossRefGoogle Scholar
  19. Diffenbaugh, N. S., M. Scherer, and R. J. Trapp. 2013. “Robust increases in severe thunderstorm environments in response to greenhouse forcing” Proceedings of the National Academy of Sciences110: 16361–16366.CrossRefGoogle Scholar
  20. Donato, D. C., J. L. Campbell, and J. F. Franklin. 2012. “Multiple successional pathways and precocity in forest development: can some forests be born complex?” Journal of Vegetation Science 23: 576–584.CrossRefGoogle Scholar
  21. Dove, N. C., and W. S. Keeton. 2015. “Structural complexity enhancement increases fungi diversity in northern hardwood forests” Fungal Ecology13: 181–192.CrossRefGoogle Scholar
  22. Doyon, F., D. Gagnon, and J. F. Giroux. 2005. “Effects of strip and single-tree selection cutting on birds and their habitat in a southwestern Quebec northern hardwood forest” Forest Ecology and Management209: 101–116.CrossRefGoogle Scholar
  23. Fahey, R. T., B. Alveshere, J. I. Burton, A. D’Amato, Y. L. Dickinson, W. S. Keeton, C. C. Kern, et al. 2018. “Shifting conceptions of complexity in forest management and silviculture.” Forest Ecology and Management. In Press.CrossRefGoogle Scholar
  24. Fahey, R. T., and C. G. Lorimer. 2013. “Restoring a midtolerant pine species as a component of late-successional forests: results of gap-based planting trials” Forest Ecology and Management292: 139–149.CrossRefGoogle Scholar
  25. Fassnacht, K. S., D. R. Bronson, B. J. Palik, A. W. D’Amato, C. Lorimer, G., and K. J. Martin. 2015. “Accelerating the development of old-growth characteristics in second-growth northern hardwoods.” General Technical Report NRS-144. Northern Research Station. Newtown Square, PA: USDA Forest Service.CrossRefGoogle Scholar
  26. Fassnacht, K. S., and T. W. Steele. 2016. “Snag dynamics in northern hardwood forests under different management scenarios” Forest Ecology and Management363: 267–276.CrossRefGoogle Scholar
  27. FEMAT [Forest Ecosystem Management Assessment Team]. 1993. Forest Ecosystem Management: An Ecological, Economic, and Social Assessment. Portland, OR: USDA Forest Service.Google Scholar
  28. Ford, S. E., and W. S. Keeton. 2017. “Enhanced carbon storage through management for old-growth characteristics in northern hardwoods” Ecosphere8: 1–20.Google Scholar
  29. Forrester, J. A., D. J. Mladenoff, and S. T. Gower. 2013. “Experimental manipulation of forest structure: near-term effects on gap and stand scale C dynamics” Ecosystems16: 1455–1472.CrossRefGoogle Scholar
  30. Franklin, J. F., D. R. Berg, D. A. Thornburgh, and J. C. Tappeiner. 1997. “Alternative silvicultural approaches to timber harvesting: variable retention harvest system.” In Creating a Forestry for the 21st Century: The Science of Ecosystem Management, edited by K. A. Kohm, and J. F. Franklin, 111–140. Washington, DC: Island Press.Google Scholar
  31. Franklin, J. F., R. J. Mitchell, and B. Palik. 2007. “Natural disturbance and stand development principles for ecological forestry.” General Technical Report NRS-19. Northern Research Station. Newton Square, PA: USDA Forest Service.CrossRefGoogle Scholar
  32. Franklin, J. F., T. A. Spies, R. Van Pelt, A. B. Carey, D. A. Thornburgh, D. R. Berg, D. B. Lindenmayer, M. E. Harmon, W. S. Keeton, D. C. Shaw, K. Bible, and J. Chen. 2002. “Disturbances and structural development of natural forest ecosystems with silvicultural implications, using Douglas-fir forests as an example” Forest Ecology and Management155: 399–423.CrossRefGoogle Scholar
  33. Frey, S., A. Hadley, S. Johnson, M. Schulze, J. Jones, and M. Betts. 2016. “Spatial models reveal the microclimatic buffering capacity of old-growth forests.” Science Advances 22: e1501392.CrossRefGoogle Scholar
  34. Gauthier, M-M., M-C. Lambert, and S. Bedard. 2016. “Effects of harvest gap size, soil scarification, and vegetation control on regeneration dynamics in sugar maple-yellow birch stands” Forest Science62: 237–246.CrossRefGoogle Scholar
  35. Goodburn, J. M. 1996. “Comparison of forest habitat structure and composition in old-growth and managed northern hardwoods in Wisconsin and Michigan.” Master’s thesis, University of Wisconsin–Madison.Google Scholar
  36. Goodburn, J. M., and C. G. Lorimer. 1999. “Population structure in old-growth and managed northern hardwoods: an examination of the balanced diameter distribution concept” Forest Ecology and Management118: 11–29.CrossRefGoogle Scholar
  37. Gottesman, A., and W. S. Keeton. 2017. “Regeneration responses to management for old-growth characteristics in northern hardwood-conifer forests” Forests8: 1–21.CrossRefGoogle Scholar
  38. Gronewald, C. A., A. W. D’Amato, and B. J. Palik. 2010. “The influence of cutting cycle and stocking level on the structure and composition of managed old-growth northern hardwoods” Forest Ecology and Management259: 1151–1160.CrossRefGoogle Scholar
  39. Gustafson, E. J., and T. R. Crow. 1996. “Simulating the effects of alternative forest management strategies on landscape structure” Journal of Environmental Management47: 77–94.CrossRefGoogle Scholar
  40. Gustafsson, L., S. C. Baker, J. Bauhus, W. J. Beese, A. Brodie, J. Kouki, D. B. Lindenmayer, et al. 2012. “Retention forestry to maintain multifunctional forests: a world perspective” Bioscience62: 633–645.CrossRefGoogle Scholar
  41. Hagenbuch, S., K. Manaras, N. Patch, J. Shallow, K. Sharpless, M. Snyder, and K. Thompson 2013. Managing Your Woods with Birds in Mind: A Vermont Landowner’s Guide. Huntington and Waterbury, VT: Audubon Vermont and Vermont Department of Forests, Parks, and Recreation.Google Scholar
  42. Hale, C. M., J. Pastor, and K. A. Rusterholtz. 1999. “Comparison of structural and compositional characteristics in old-growth and mature, managed hardwood forests of Minnesota, U.S.A” Canadian Journal of Forest Research29: 1479–1489.CrossRefGoogle Scholar
  43. Halpin, C. R., and C. G. Lorimer. 2016. “Trajectories and resilience of stand structure in response to variable disturbance severities in northern hardwoods” Forest Ecology and Management365: 69–82.CrossRefGoogle Scholar
  44. Halpin, C. R., and C. G. Lorimer. 2017. “Predicted long-term effects of group selection on species composition and stand structure in northern hardwood forests” Forest Ecology and Management400: 677–691.CrossRefGoogle Scholar
  45. Hanson, J. J., and C. G. Lorimer. 2007. “Forest structure and light regimes following moderate wind storms: Implications for multi-cohort management” Ecological Applications17: 1325–1340.PubMedCrossRefPubMedCentralGoogle Scholar
  46. Hanson, J. J., C. G. Lorimer, C. R. Halpin, and B. J. Palik. 2012. “Ecological forestry in an uneven-aged, late-successional forest: simulated effects of contrasting treatments on structure and yield” Forest Ecology and Management270: 94–107.CrossRefGoogle Scholar
  47. Hardt, R. A., and W. T. Swank. 1997. “A comparison of structural and compositional characteristics of southern Appalachian young second-growth, maturing second-growth, and old-growth stands” Natural Areas Journal17: 42–52.Google Scholar
  48. Heinselman, M. L. 1973. “Fire in the virgin forests of the Boundary Waters Canoe Area, Minnesota.” Quaternary Research 3: 329–382.CrossRefGoogle Scholar
  49. Hunter, W. C., D. A. Buechler, R. A. Canterbury, J. L. Confer, and P. B. Hamel. 2001. “Conservation of disturbance dependent birds in eastern North America” Wildlife Society Bulletin29: 425–439.Google Scholar
  50. Keeton, W. S. 2006. “Managing for late-successional/old-growth characteristics in northern hardwood-conifer forests” Forest Ecology and Management235: 129–142.CrossRefGoogle Scholar
  51. Keeton, W. S. 2007. “Role of managed forestlands and models for sustainable forest management: perspectives from North America” George Wright Forum24: 38–53.Google Scholar
  52. Keeton, W. S., E. M. Copeland, and M. C. Watzin. 2017. “Towards flood resilience: exploring linkages between riparian forest structure and geomorphic condition in northeastern U.S. streams” Canadian Journal of Forest Research47: 476–487.CrossRefGoogle Scholar
  53. Keeton, W. S., C. E. Kraft, and D. R. Warren. 2007. “Mature and old-growth riparian forests: structure, dynamics, and effects on Adirondack stream habitats” Ecological Applications17: 852–868.Google Scholar
  54. Keeton, W. S. and A. R. Troy. 2006. “Balancing ecological and economic objectives while managing for late-successional forest structure.” In Ecologisation of Economy as a Key Prerequisite for Sustainable Development. Proceedings of the International Conference, Sept. 22–23, 2005, edited by L. Zahvoyska, 22–23. L’viv, Ukraine: Ukrainian National Forestry University.Google Scholar
  55. Keeton, W. S., A. A. Whitman, G. G. McGee, and C. L. Goodale. 2011. “Late-successional biomass development in northern hardwood-conifer forests of the northeastern United States” Forest Science57: 489–505.Google Scholar
  56. Kerchner, C. and W. S. Keeton. 2015. “California’s regulatory forest carbon market: panacea or Pandora’s box for northeastern landowners?” Forest Policy and Economics 50: 70–81.Google Scholar
  57. Kern, C. C., J. Burton, P. Raymond, A. D’Amato, W. S. Keeton, A. A. Royo, M. B. Walters, C. R. Webster, and J. L. Willis. 2016. “Challenges facing gap-based silviculture and possible solutions for mesic northern forests in North America” Forestry90: 4–17.Google Scholar
  58. Kern, C. C., A. W. D’Amato, and T. F. Strong. 2013. “Diversifying the composition and structure of managed, late-successional forests with harvest gaps: what is the optimal gap size?” Forest Ecology and Management 304: 110–120.CrossRefGoogle Scholar
  59. Kern, C. C., P. B. Reich, R. A. Montgomery, and T. F. Strong. 2012. “Do deer and shrubs override canopy gap size effects on growth and survival of yellow birch, northern red oak, eastern white pine, and eastern hemlock seedlings?” Forest Ecology and Management 267: 134–143.CrossRefGoogle Scholar
  60. Klockow, P. A., A. W. D’Amato, and J. B. Bradford. 2013. “Impacts of post-harvest slash and live-tree retention on biomass and nutrient stocks in Populus tremuloides Michx.-dominated forests, northern Minnesota USA” Forest Ecology and Management291: 278–288.CrossRefGoogle Scholar
  61. Leak, W. B. 1999. “Species composition and structure of a northern hardwood stand after 61 years of group/patch selection” Northern Journal of Applied Forestry16: 151–153.Google Scholar
  62. Leak, W. B., and P. E. Sendak. 2002. “Changes in species, grade, and structure over 48 years in a managed New England northern hardwood stand” Northern Journal of Applied Forestry19: 25–27.Google Scholar
  63. Lorimer, C. G., and C. R. Halpin. 2014. “Classification and dynamics of developmental stages in late-successional temperate forests” Forest Ecology and Management334: 344–357.CrossRefGoogle Scholar
  64. Lorimer, C. G., and A. S. White. 2003. “Scale and frequency of natural disturbances in the northeastern U.S.: Implications for early-successional forest habitats and regional age distributions” Forest Ecology and Management185: 41–64.CrossRefGoogle Scholar
  65. McGarvey, J. C., J. R. Thompson, H. E. Epstein, and H. H. Shugart, Jr. 2015. “Carbon storage in old-growth forests of the Mid-Atlantic: toward better understanding the eastern forest carbon sink” Ecology96: 311–317.PubMedCrossRefPubMedCentralGoogle Scholar
  66. McGee, G. G., D. J. Leopold, and R. D. Nyland. 1999. “Structural characteristics of old-growth, maturing, and partially cut northern hardwood forests” Ecological Applications9: 1316–1329.CrossRefGoogle Scholar
  67. McKenny, H. C., W. S. Keeton, and T. M. Donovan. 2006. “Effects of structural complexity enhancement on eastern red-backed salamander (Plethodon cinereus) populations in northern hardwood forests” Forest Ecology and Management230: 186–196.CrossRefGoogle Scholar
  68. McLachlan, J. S., D. R. Foster, and F. Menalled. 2000. “Anthropogenic ties to late-successional structure and composition in four New England hemlock stands” Ecology81: 717–733.CrossRefGoogle Scholar
  69. Meigs, G. W. and W. S. Keeton. 2018. “Intermediate-severity wind disturbance in mature temperate forests: effects on legacy structure, carbon storage, and stand dynamics.” Ecological Applications. doi:10.1002/eap.1691.CrossRefPubMedPubMedCentralGoogle Scholar
  70. Messier, C., K. Puettmann, and D. Coates. 2013. Managing Forests as Complex Adaptive Systems: Building Resilience to the Challenge of Global Change. New York, NY: Routledge.Google Scholar
  71. Millar, C. I., N. L. Stephenson, and S. L. Stephens. 2007. “Climate change and forests of the future: managing in the face of uncertainty” Ecological Applications17: 2145–2151.PubMedCrossRefPubMedCentralGoogle Scholar
  72. Miller, G. W., J. N. Kochenderfer, and D. B. Fekedulegn. 2006. Influence of individual reserve trees on nearby reproduction in two-aged Appalachian hardwood stands. Forest Ecology and Management 224: 241–251.CrossRefGoogle Scholar
  73. Mladenoff, D. J., M. A. White, T. R. Crow, and J. Pastor. 1994. “Applying principles of landscape design and management to integrate old-growth forest enhancement and commodity use” Conservation Biology8: 752–762.CrossRefGoogle Scholar
  74. Montgomery, R. A., B. J. Palik, S. B. Boyden, and P. B. Reich. 2013. “New cohort growth and survival in variable retention harvests of a pine ecosystem in Minnesota, USA” Forest ecology and management310: 327–335.CrossRefGoogle Scholar
  75. North, M. P. and W. S. Keeton. 2008. “Emulating natural disturbance regimes: an emerging approach for sustainable forest management. In Patterns and Processes in Forest Landscapes - Multiple Use and Sustainable Management, edited by R. Lafortezza, J. Chen, G. Sanesi, and T. R. Crow, 341–372. The Netherlands: Springer Verlag.CrossRefGoogle Scholar
  76. Nunery, J. S. and W. S. Keeton. 2010. “Forest carbon storage in the northeastern United States: Net effects of harvesting frequency, post-harvest retention, and wood products” Forest Ecology and Management259: 1363–1375.CrossRefGoogle Scholar
  77. Odum, E. P. 1969. “The strategy of ecosystem development.” Science 164: 262 – 270.PubMedCrossRefPubMedCentralGoogle Scholar
  78. O’Hara, K. L. 1998. “Silviculture for structural diversity: a new look at multi-aged systems” Journal of Forestry96: 4–10.Google Scholar
  79. Owen, S. F., M. A. Menzel, and J. W. Edwards. 2004. “Bat activity in harvested and intact forest stands in the Allegheny Mountains” Northern Journal of Applied Forestry21: 154–159.Google Scholar
  80. Palik, B. J., and A. W. D’Amato. 2017. “Ecological forestry: much more than retention harvesting.” Journal of Forestry 115: 51.CrossRefGoogle Scholar
  81. Palik, B. J., R. A. Montgomery, P. B. Reich, and S. B. Boyden. 2014. “Biomass growth response to spatial pattern of variable-retention harvesting in a northern Minnesota pine ecosystem” Ecological Applications24: 2078–2088.PubMedCrossRefPubMedCentralGoogle Scholar
  82. Palik, B. J., and J. Zasada. 2003. “An ecological context for regenerating multi-cohort, mixed species red pine forests.” Research Note NC-382. North Central Research Station. St. Paul, MN: USDA Forest Service.CrossRefGoogle Scholar
  83. Puettmann, K., D. Coates, and C. Messier. 2009. A Critique of Silviculture: Managing for Complexity. Washington, DC: Island Press.Google Scholar
  84. Puettmann, K. J., S. M. Wilson, S. C. Baker, P. Donoso, L. Drossler, G. Amente, B. D. Harvey, et al. 2015. “Silvicultural alternatives to conventional even-aged forest management—What limits global adoption?” Forest Ecosystems 2: 1–16.CrossRefGoogle Scholar
  85. Raymond, P., S. Bédard, V. Roy, C. Larouche, and S. Tremblay. 2009. “The irregular shelterwood system: review, classification, and potential application to forests affected by partial disturbances” Journal of Forestry107: 405–413.Google Scholar
  86. Rhemtulla, J. M., D. J. Mladenoff, and M. K. Clayton. 2007. “Regional land-cover conversion in the US Upper Midwest: magnitude of change and limited recovery (1850–1935–1993)” Landscape Ecology22: 57–75.CrossRefGoogle Scholar
  87. Rhemtulla, J. M., D. J. Mladenoff, and M. K. Clayton. 2009. “Historical forest baselines reveal potential for continued carbon sequestration.” Proceedings of the National Academy of Sciences 106: 6082–6087. CrossRefGoogle Scholar
  88. Runkle, J. R., and T. C. Yetter. 1987. “Treefalls revisited: gap dynamics in the southern Appalachians” Ecology68: 417–424.CrossRefGoogle Scholar
  89. Schuler, T. M. 2004. “Fifty years of partial harvesting in a mixed mesophytic forest: composition and productivity” Canadian Journal of Forest Research34: 985–997.CrossRefGoogle Scholar
  90. Schwartz, N. B., D. L. Urban, P. S. White, A. Moody, and R. N. Klein. 2016. “Vegetation dynamics vary across topographic and fire severity gradients following prescribed burning in Great Smoky Mountains National Park” Forest Ecology and Management365: 1–11.CrossRefGoogle Scholar
  91. Selva, S. B. 2003. “Using calicioid lichens and fungi to assess ecological continuity in the Acadian Forest Ecoregion of the Canadian Maritimes” The Forestry Chronicle79: 550–558.CrossRefGoogle Scholar
  92. Senécal, J-F, F. Doyon, and C. Messier. 2018. “Management implications of varying gap detection height thresholds and other canopy dynamics processes in temperate deciduous forests.” Forest Ecology and Management. In Press.Google Scholar
  93. Seymour, R. S., A. S. White, and P. H. deMaynadier. 2002. “Natural disturbance regimes in northeastern North America: evaluating silvicultural systems using natural scales and frequencies” Forest Ecology and Management155: 357–367.CrossRefGoogle Scholar
  94. Singer, M. T., and C. G. Lorimer. 1997. “Crown release as a potential old-growth restoration approach in northern hardwoods” Canadian Journal of Forest Research27: 1222–1232.CrossRefGoogle Scholar
  95. Smith, K. J., W. S. Keeton, M. Twery, and D. Tobi. 2008. “Understory plant response to alternative forestry practices in northern hardwood-conifer forests.” Canadian Journal of Forest Research 38: 1–17.CrossRefGoogle Scholar
  96. Spies, T. A. 1997. “Forest stand structure, composition, and function.” In Creating a Forestry for the 21st century, edited by K. A. Kohm, and J. F. Franklin, 11–30. Washington, DC: Island Press.Google Scholar
  97. Stribling, H. L., H. R. Smith, and R. H. Yahner. 1990. “Bird community response to timber stand improvement and snag retention” Northern Journal of Applied Forestry7: 35–38.Google Scholar
  98. Tyrrell, L. E., and T. R. Crow. 1994. “Structural characteristics of old-growth hemlock-hardwood forests in relation to age” Ecology75: 370–386.CrossRefGoogle Scholar
  99. Urbano, A. R. and W. S. Keeton. 2017. “Forest structural development, carbon dynamics, and co-varying habitat characteristics as influenced by land-use history and reforestation approach” Forest Ecology and Management392: 21–35.CrossRefGoogle Scholar
  100. Walters, M. B., E. J. Farinosi, J. L. Willis, and K. W. Gottschalk. 2016. “Managing for diversity: harvest gap size drives complex light, vegetation, and deer herbivory impacts on tree seedlings.” Ecosphere 7: e01397.CrossRefGoogle Scholar
  101. Warren, D. R., W. S. Keeton, P. M. Kiffney, M. J. Kaylor, H. A. Bechtold, and J. Magee. 2016. “Changing forests-changing streams: riparian forest stand development and ecosystem function in temperate headwaters.” Ecosphere 7 (8). doi:10.1002/ecs2.1435.CrossRefGoogle Scholar
  102. Webster, C. R. and C. G. Lorimer. 2005. “Minimum opening sizes for canopy recruitment of midtolerant tree species: a retrospective approach” Ecological Applications15: 1245–1262.CrossRefGoogle Scholar
  103. Woods, K. D. 2000. “Dynamics in late-successional hemlock-hardwood forests over three decades” Ecology81: 110–126.Google Scholar
  104. Zenner, E. K., S. A. Acker, and W. H. Emmingham. 1998. “Growth reduction in harvest-age, coniferous forests with residual trees in the western central Cascade Range of Oregon” Forest Ecology and Management102: 75–88.CrossRefGoogle Scholar
  105. Ziegler, S. S. 2000. “A comparison of structural characteristics between old-growth and post-fire second-growth hemlock-hardwood in Adirondack Park, New York, U.S.A” Global Ecology and Biogeography9: 373–389.CrossRefGoogle Scholar
  106. Ziegler, S. S. 2002. “Disturbance of hemlock-dominated old-growth in northern New York, USA” Canadian Journal of Forest Research32: 2106–2115.CrossRefGoogle Scholar

Copyright information

© Andrew M. Barton and William S. Keeton 2018

Authors and Affiliations

  • William S. Keeton
  • Craig G. Lorimer
  • Brian J. Palik
  • Frédérik Doyon

There are no affiliations available

Personalised recommendations