Restoration of old-growth forest structure is an emerging silvicultural goal, especially in those regions where old-growth abundance falls below the historic range of variability. However, longitudinal studies of old-growth dynamics that can inform silvicultural and policy options are few. We analyzed the change in structure, including stand density, diameter distribution, and the abundance of large live, standing dead, and downed dead trees on 58 late-successional and old-growth plots in Maine, USA, and compared these to regional data from the U.S. Forest Inventory and Analysis program. Structural dynamics on the late-successional plots reflected orderly change associated with density-dependent growth and mortality, but dynamics on the old-growth plots were more variable. Some plots experienced heavy mortality associated with beech bark disease. Diameter distributions conformed poorly to a classic exponential distribution, and did not converge toward such a distribution at the plot scale. Although large live trees showed a broad trend of increasing density in regional forests, recent harvesting patterns offset a considerable fraction of those gains, while mean diameter was static and the number of large dead trees was weakly declining. Even though forests of the northeast are aging, changes in silviculture and forest policy are necessary to accelerate restoration of old-growth structure.
References
[1]
Harmon, M.E.; Ferrell, W.K.; Franklin, J.F. Effects on carbon storage of conversion of old-growth forests to young forests. Science 1990, 247, 699–702.
[2]
Luyssaert, S.; Schulze, E.; B？rner, A. Old-growth forests as global carbon sinks. Nature 2008, 455, 213–215, doi:10.1038/nature07276.
[3]
Whitman, A.A.; Hagan, J.M. An index to identify late-successional forest in temperate and boreal zones. For. Ecol. Manag. 2007, 246, 144–154, doi:10.1016/j.foreco.2007.03.004.
[4]
Reed, W.J. The decision to conserve or harvest old-growth forest. Ecol. Econ. 1993, 8, 45–69.
[5]
Freedman, B.; Zelazny, V.; Beaudette, D.; Fleming, T.; Fleming, S.; Forbes, G.; Gerrow, J.; Johnson, G.; Woodley, S. Biodiversity implications of changes in quantity of dead organic matter in managed forests. Environ. Rev. 1996, 4, 238–265, doi:10.1139/a96-013.
[6]
Recher, H. Conservation and Management of Eucalypt Forest Vertebrates. In Conservation of Faunal Diversity in Forested Landscapes; DeGraaf, R.M., Miller, R.I., Eds.; Chapman and Hall: New York, NY, USA, 1996; pp. 339–388.
[7]
Lindenmayer, D.B.; Franklin, J.F. Conserving Forest Biodiversity: A Comprehensive Multiscaled Approach; Island Press: Washington, DC, USA, 2002.
[8]
Irland, L.C. The Northeast’s Changing Forests; Harvard University Press: Cambridge, MA, USA, 1999.
[9]
Foster, D.R. Forests in Time: The Environmental Consequences of 1,000 Years of Change in New England; Yale University Press: New Haven, CT, USA, 2006.
[10]
Pan, Y.; Chen, J.-M.; Birdsey, R.; McCullough, K.; He, L.; Deng, F. Age structure and disturbance legacy of North American forests. Biogeosciences 2011, 8, 715–732, doi:10.5194/bg-8-715-2011.
[11]
Lorimer, C.G. The Presettlement forest and natural disturbance cycle of northeastern maine. Ecology 1977, 58, 139–148, doi:10.2307/1935115.
[12]
Cogbill, C.V. Vegetation of the presettlement forests of northern New England and New York. Rhodora 2000, 102, 250–276.
[13]
Lorimer, C.G. Historical and ecological roles of disturbance in eastern North American forests: 9,000 years of change. Wildl. Soc. Bull. 2001, 29, 425–439.
[14]
Lorimer, C.G.; White, A.S. Scale and frequency of natural disturbances in the northeastern U.S.: Implications for early-successional forest habitats and regional age distributions. For. Ecol. Manag. 2003, 185, 41–64, doi:10.1016/S0378-1127(03)00245-7.
[15]
Davis, M.B. Eastern old-Growth Forests: Prospects for Rediscovery and Recovery; Island Press: Covelo, CA, USA, 1996.
[16]
Keeton, W.S. Managing for late-successional/old-growth characteristics in northern hardwood-conifer forests. For. Ecol. Manag. 2006, 235, 129–142.
[17]
North, M.P.; Keeton, W.S. Emulating Natural Disturbance Regimes: An Emerging Approach for Sustainable Forest ManagementChapter 17. In Patterns and Processes in Forest Landscapes; Lafortezza, R., Sanesi, G., Chen, J., Eds.; Springer Netherlands: Houten, The Netherlands, 2008; pp. 341–372.
[18]
Bauhus, J.; Puettmann, K.; Messier, C. Silviculture for old-growth attributes. For. Ecol. Manag. 2009, 258, 525–537, doi:10.1016/j.foreco.2009.01.053.
[19]
D’Amato, A.W.; Catanzaro, P.C. Restoring Old-Growth Characteristics. In Cooperative Extension Landowner Outreach Pamphlet; University of Massachusetts: Amherst, MA, USA, 2007; p. 18.
[20]
Burrascano, S.; Keeton, W.S.; Sabatini, F.M.; Blasi, C. Commonality and variability in the structural attributes of moist temperate old-growth forests: A global review. For. Ecol. Manag. 2013, 291, 458–479.
[21]
Gore, J.A.; Patterson, W.A., III. Mass of downed wood in northern hardwood forests in New Hampshire: Potential effects of forest management. Can. J. For. Res. 1986, 16, 335–339, doi:10.1139/x86-057.
[22]
Leak, W.B. Characteristics of Five Climax Stands in New Hampshire; USDA Forest Service, Northeastern Forest Experiment Station: Broomall, PA, USA, 1987.
[23]
Odum, E.P. The strategy of ecosystem development. Science 1969, 164, 262–270.
[24]
Jarvis, P.G. Atmospheric carbon dioxide and forests. Philos. Trans. R. Soc. Biol. Sci. 1989, 324, 369–392.
[25]
Keith, H.; Mackey, B.G.; Lindenmayer, D.B. Re-evaluation of forest biomass carbon stocks and lessons from the world’s most carbon-dense forests. Proc. Natl. Acad. Sci. USA 2009, 106, 11635–11640.
[26]
Keeton, W.; Whitman, A.; McGee, G.G.; Goodale, C.L. Late-successional biomass development in northern hardwood-conifer forests of the northeastern United States. For. Sci. 2011, 57, 489–505.
[27]
Galik, C.S.; Jackson, R.B. Risks to forest carbon offset projects in a changing climate. For. Ecol. Manag. 2009, 257, 2209–2216, doi:10.1016/j.foreco.2009.03.017.
Henry, J.D.; Swan, J.M.A. Reconstructing forest history from live and dead plant material--an approach to the study of forest succession in southwest New Hampshire. Ecology 1974, 55, 772–783, doi:10.2307/1934413.
[32]
Ziegler, S.S. Composition, structure, and disturbance history of old-growth and second-growth forests in Adirondack Park, New York. Phys. Geogr. 2004, 25, 152–169, doi:10.2747/0272-3646.25.2.152.
[33]
D’Amato, A.W.; Orwig, D.A.; Foster, D.R. The influence of successional processes and disturbance on the structure of Tsuga canadensis forests. Ecol. Appl. 2004, 18, 1182–1199.
[34]
Foster, D.R. Disturbance history, community organization and vegetation dynamics of the old-growth Pisgah Forest, southwestern New Hampshire, USA. J. Ecol. 1988, 76, 105–134, doi:10.2307/2260457.
[35]
Filip, S.M.; Marquis, D.A.; Leak, W.B. Development of Old-Growth Northern Hardwoods on Bartlett Experimental Forest—A 22-Year Record; U.S. Forest Service: Washington, DC, USA, 1960.
[36]
Woods, K.D. Dynamics in late-successional hemlock-hardwood forests over three decades. Ecology 2000, 81, 110–126.
[37]
Woods, K.D. Long-term change and spatial pattern in a late-successional hemlock–northern hardwood forest. J. Ecol. 2000, 88, 267–282, doi:10.1046/j.1365-2745.2000.00448.x.
[38]
Lorimer, C.G.; Dahir, S.E.; Nordheim, E.V. Tree mortality rates and longevity in mature and old-growth hemlock-hardwood forests. J. Ecol. 2001, 89, 960–971, doi:10.1111/j.1365-2745.2001.00619.x.
[39]
Busing, R.T. Tree mortality, canopy turnover, and woody detritus in old cove forests of the southern Appalachians. Ecology 2005, 86, 73–84, doi:10.1890/04-0410.
[40]
Hagan, J.M.; Grove, S.L. Coarse woody debris: Humans and nature competing for trees. J. For. 1999, 97, 6–11.
[41]
Gunn, J.S.; Hagan, J.M. Woodpecker abundance and tree use in uneven-aged managed, and unmanaged, forest in northern Maine. For. Ecol. Manag. 2000, 126, 1–12, doi:10.1016/S0378-1127(99)00078-X.
[42]
Gunn, J.S.; Ducey, M.J.; Whitman, A.A. Late-successional and old-growth forest carbon temporal dynamics in the northern forest (northeastern USA). For. Ecol. Manag. 2013. in press.
[43]
Whitman, A. Manomet Center for Conservation Sciences, Brunswick, ME, USA. Unpublished Data, 2013.
[44]
Fraver, S.; White, A.S.; Seymour, R.S. Natural disturbance in an old-growth landscape of northern Maine, USA. J. Ecol. 2009, 97, 289–298, doi:10.1111/j.1365-2745.2008.01474.x.
[45]
Tyrrell, L.E.; Nowacki, G.J.; Crow, T.R.; Buckley, D.S.; Nauertz, E.A.; Niese, J.N.; Rollinger, J.L.; Zasada, J.C. Information about Old Growth for Selected Forest Type Groups in the Eastern United States; USDA Forest Service, North Central Forest Experiment Station: St. Paul, MN, USA, 1998.
[46]
Gove, J.H.; van Deusen, P.C. On fixed-area plot sampling for downed coarse woody debris. Forestry 2011, 84, 109–117, doi:10.1093/forestry/cpq049.
[47]
Reineke, L.H. Perfecting a stand-density index for even-aged forests. J. Agric. Res. 1933, 46, 627–638.
[48]
Ducey, M.J.; Knapp, R.A. A stand density index for complex mixed species forests in the northeastern United States. For. Ecol. Manag. 2010, 260, 1613–1622, doi:10.1016/j.foreco.2010.08.014.
[49]
Gingrich, S.F. Measuring and evaluating stocking and stand density in upland hardwood forests in the Central States. For. Sci. 1967, 13, 38–53.
[50]
Meyer, H.A.; Stevenson, D.D. The structure and growth of virgin beech-birch-maple-hemlock forests in northern Pennsylvania. J. Agric. Res. 1943, 67, 465–484.
[51]
Meyer, H.A. Structure, growth, and drain in uneven-aged forests. J. For. 1952, 50, 85–92.
[52]
De Liocourt, F. De l’amanagement des sapinieres. In Bulletin Trimestriel; Société forestière de Franche-Comté et Belfort, Ed.; Impr. et litographie de Paul Jacquin: Besancon, France, 1898. (in French).
[53]
Leak, W.B. The J-shaped probability distribution. For. Sci. 1965, 11, 405–409.
[54]
Leak, W.B. An expression of diameter distribution for unbalanced, uneven-aged stands and forests. For. Sci. 1964, 10, 39–50.
[55]
Goff, F.G.; West, D. Canopy-understory interaction effects on forest population structure. For. Sci. 1975, 21, 98–108.
[56]
Bailey, R.L.; Dell, T.R. Quantifying diameter distributions with the Weibull function. For. Sci. 1973, 19, 97–104.
[57]
Stacy, E.W. A generalization of the gamma distribution. Ann. Math. Stat. 1962, 33, 1187–1192, doi:10.1214/aoms/1177704481.
[58]
McDonald, J.B. Some generalized functions for the size distribution of income. Econometrica 1984, 52, 647–664, doi:10.2307/1913469.
[59]
Akaike, H. A new look at the statistical model identification. IEEE Trans. Autom. Control 1974, 19, 716–723, doi:10.1109/TAC.1974.1100705.
[60]
Burnham, K.P.; Anderson, D.R. Model Selection and Multimodel Inference: A Practical Information-Theoretic Approach, 2nd ed. ed.; Springer-Verlag: New York, NY, USA, 2002.
[61]
The Enhanced Forest Inventory and Analysis Program—National Sampling Design and Estimation Procedures; Bechtold, W.A., Patterson, P.L., Eds.; USDA Forest Service, Southern Research Station: Asheville, NC, USA, 2005.
[62]
R Core Team. R: A Language and Environment for Statistical Computing, R Foundation for Statistical Computing, Vienna, Austria, 2012.
[63]
Beaudet, M.; Brisson, J.; Gravel, D.; Messier, C. Effect of a major canopy disturbance on the coexistence of Acer saccharum and Fagus grandifolia in the understorey of an old-growth forest. J. Ecol. 2007, 95, 458–467, doi:10.1111/j.1365-2745.2007.01219.x.
[64]
Keeton, W.S.; Kraft, C.E.; Warren, D.R. Mature and old-growth riparian forests: Structure, dynamics, and effects on Adirondack stream habitats. Ecol. Appl. 2007, 17, 852–868, doi:10.1890/06-1172.
[65]
Hoover, C.M.; Leak, W.B.; Keel, B.G. Benchmark carbon stocks from old-growth forests in northern New England, USA. For. Ecol. Manag. 2012, 266, 108–114, doi:10.1016/j.foreco.2011.11.010.
[66]
McGarrigle, E.; Kershaw, J.A.; Ducey, M.J.; Lavigne, M.B. A new approach to modeling stand-level dynamics based on informed random walks: Influence of bandwidth and sample size. Forestry 2013, 86, 377–389, doi:10.1093/forestry/cpt008.
[67]
Oliver, C.D.; Larson, B.C. Forest Stand Dynamics; Wiley: New York, NY, USA, 1996.
[68]
Franklin, J.F.; Spies, T.A.; van Pelt, R.; Carey, A.; Thornburgh, D.; Berg, D.R.; Lindenmayer, D.; Harmon, M.; Keeton, W.S.; Shaw, D.C.; et al. Disturbances and the structural development of natural forest ecosystems with some implications for silviculture. For. Ecol. Manag. 2002, 155, 399–423, doi:10.1016/S0378-1127(01)00575-8.
[69]
Morin, R.S.; Liebhold, A.M.; Tobin, P.C.; Gottschalk, K.W.; Luzader, E. Spread of beech bark disease in the eastern United States and its relationship to regional forest composition. Can. J. For. Res. 2007, 37, 726–736, doi:10.1139/X06-281.
[70]
Kasson, M.T.; Livingston, W.H. Relationships among beech bark disease, climate, radial growth response and mortality of American beech in northern Maine, USA. For. Pathol. 2012, 42, 199–212, doi:10.1111/j.1439-0329.2011.00742.x.
Canham, C.D.; Rogers, N.; Buchholz, T. Regional variation in forest harvest regimes in the northeastern United States. Ecol. Appl. 2013, 23, 515–522, doi:10.1890/12-0180.1.
[73]
Zheng, D.L.; Heath, L.S.; Ducey, M.J.; Butler, B. Relationships between ownership, forest aboveground biomass distributions and landscape dynamics in the New England states of USA. Environ. Manag. 2010, 45, 377–386, doi:10.1007/s00267-009-9408-3.
[74]
Benjamin, J.; Lilieholm, R.J.; Damery, D. Challenges and opportunities for the northeastern forest bioindustry. J. For. 2009, 107, 125–131.
[75]
Franklin, J.F. Toward a new forestry. Am. For. 1989, 95, 37–44.
[76]
Orwig, D.A.; Cogbill, C.V.; Foster, D.R.; O’Keefe, J.F. Variations in old-growth structure and definitions: Forest dynamics on Wachusett Mountain, Massachusetts. Ecol. Appl. 2001, 11, 437–452, doi:10.1890/1051-0761(2001)011[0437:VIOGSA]2.0.CO;2.
[77]
Leak, W.B.; Solomon, D.S.; de Bald, P.S. Silvicultural Guide to Northern Hardwood Types in the Northeast, Revised; USDA Forest Service, Northeastern Forest Research Station: Broomall, PA, USA, 1987.
[78]
Frank, R.M.; Bjorkbom, J.C. A Silvicultural Guide for Spruce-Fir in the Northeast; USDA Forest Service, Northeastern Forest Research Station: Upper Darby, PA, USA, 1973.
[79]
Gronewald, C.A.; D’Amato, A.W.; Palik, B.J. The influence of cutting cycle and stocking level on the structure and composition of managed old-growth northern hardwoods. For. Ecol. Manag. 2010, 259, 1151–1160, doi:10.1016/j.foreco.2010.01.001.
[80]
Smith, D.M.; Larson, B.C.; Kelty, M.J.; Ashton, P.M.S. The Practice of Silviculture: Applied Forest Ecology, 9th ed. ed.; John Wiley and Sons: New York, NY, USA, 2007.
[81]
Smith, D.M. Concluding Remarks. In The Ecology and Silviculture of Mixed-Species Forests; Kelty, M.J., Oliver, C.D., Larson, B.C., Eds.; Kluwer Academic Publishers: Dordrecht, The Netherlands, 1992; pp. 281–287.
[82]
O’Hara, K.L.; Gersonde, R.F. Stocking control concepts in uneven-aged silviculture. Forestry 2004, 77, 131–143, doi:10.1093/forestry/77.2.131.
[83]
Ducey, M.J. The Reverse-J and Beyond: Developing Practical, Effective Marking Guides. Proceedings of Implementing Uneven-Aged Management in New England: Is It Practical? Fox Research and Demonstration Forest; U.N.H. Cooperative Extension: Durham, NH, USA, 2006. Available online: http://extension.unh. edu/resources/files/Resource002220_Rep3279.pdf (accessed on 1 October 2013).
[84]
Goodburn, J.M.; Lorimer, C.G. Population structure in old-growth and managed northern hardwoods: An examination of the balanced diameter distribution concept. For. Ecol. Manag. 1999, 118, 11–29, doi:10.1016/S0378-1127(98)00478-2.
[85]
Rouvinen, S.; Kuuluvainen, T. Tree diameter distributions in natural and managed old Pinus sylvestris-dominated forests. For. Ecol. Manag. 1995, 208, 45–61, doi:10.1016/j.foreco.2004.11.021.
[86]
Schütz, J.P. Modelling the demographic sustainability of pure beech plenter forests in Eastern Germany. Ann. For. Sci. 2006, 63, 93–100, doi:10.1051/forest:2005101.
[87]
Gove, J.H.; Ducey, M.J.; Leak, W.B.; Zhang, L. Rotated sigmoid structures in managed uneven-aged northern hardwood stands: A look at the Burr Type III distribution. Forestry 2008, 81, 161–176, doi:10.1093/forestry/cpm025.
[88]
Allesandrini, A.; Biondi, F.; di Fillippo, A.; Ziaco, E.; Piovesan, G. Tree size distribution at increase spatial scales converges to the rotated sigmoid curve in two old-growth beech stands of the Italian Apennines. For. Ecol. Manag. 2011, 262, 1950–1962, doi:10.1016/j.foreco.2011.08.025.
[89]
Singer, M.T.; Lorimer, C.G. Crown release as a potential old-growth restoration approach in northern hardwoods. Can. J. For. Res. 1997, 27, 1222–1232, doi:10.1139/x97-071.
[90]
Ma, Z.; Kittredge, D.B. How family forest owners consider timber harvesting, land sale, and conservation easement decisions: Insights from Massachusetts, USA. Int. J. For. Res. 2011, 2011, 1–13.
[91]
Majumdar, I.; Laband, D.; Teeter, L.; Butler, B.J. Motivations and land-use intentions of nonindustrial private forest landowners: Comparing inheritors to noninheritors. For Sci. 2009, 55, 423–432.
[92]
Maine Forest Practices Act, 29 July 2004. Available online: http://www.maine.gov/doc/mfs/pubs/htm/fpa_04.html (accessed on 26 June 2013).
[93]
State of Maine, Department of Conservation, Maine Forest Service Silvicultural Activities Reports 1996–2011. Available online: http://www.maine.gov/doc/mfs/pubs.htm (accessed on 26 June 2013).
[94]
State of Maine. An Act to Amend the Maine Tree Growth Tax Law and the Open Space Tax Law. Public Law Chapter 618, H.P. 844-L.D. 1138. 2012. Available online: http://www. mainelegislature.org/legis/bills/getPDF.asp?paper=HP0844&item=4&snum=125 (accessed on 29 June 2013).
[95]
Sustainable Forestry Initiative 2010–2014 Standard. Available online: http://www.sfiprogram.org/files/pdf/section2sfirequirements2010-2014pdf/ (accessed on 26 June 2013).
[96]
American Forest Foundation 2010–2015 Standards of Sustainability for Forest Certification. Available online: http://www.treefarmsystem.org/certification-american-tree-farm-standards (accessed on 26 June 2013).
[97]
FSC-US Forest Management Standard, Ver. 1.0. Available online: http://us.fsc.org/download.fsc-us-forest-management-standard-v1-0.95.pdf (accessed on 27 June 2013).
[98]
Gunn, J.S.; Saah, D.S.; Fernholz, K.; Ganz, D.J. Carbon credit eligibility under area regulation of harvest levels in Northern Minnesota. For. Sci. 2011, 57, 470–478.
[99]
Nunery, J.S.; Keeton, W.S. Forest carbon storage in the northeastern United States: Net effects of harvesting frequency, post-harvest retention, and wood products. For. Ecol. Manag. 2010, 259, 1363–1375, doi:10.1016/j.foreco.2009.12.029.
[100]
Climate Action Reserve Forest Project Protocol Version 3.2. Available online: http://www.climateactionreserve.org/how/protocols/forest/dev/version-3-2/ (accessed on 27 June 2013).
[101]
American Carbon Registry Forest Carbon Project Standard v2.1. Available online: http://americancarbonregistry.org/carbon-accounting/carbon-accounting/forest-carbon-project-standard-v2.0 (accessed on 27 June 2013).
[102]
Fishburn, I.S.; Kareiva, P.; Gaston, K.J.; Armsworth, P.R. The growth of easements as a conservation tool. PLoS One 2009, 4, e4996.
