Abstract
Tsuga canadensis (L.) Carr. forests of the southern Appalachian Mountains are currently facing imminent decline induced by a nonnative insect pest, the hemlock woolly adelgid (Adelges tsugae Annand). To effectively manage these forest systems now and in the future, land managers need baseline data on forest structure and dynamics prior to large-scale Tsuga canadensis mortality. Most of our knowledge concerning the dynamics of Tsuga canadensis forests comes from more northern locations such as the Great Lakes region and New England and, therefore, may not pertain to the ecological systems found within the southern Appalachian Mountains. We examined the structure and canopy dynamics of four Tsuga canadensis forest stands within the Cataloochee watershed, in the far eastern part of Great Smoky Mountains National Park (GSMNP). We characterized the environmental settings and vertical forest layers, as well as the diameter and age-structures of each Tsuga canadensis forest stand. These environmental and structural data showed that there were indeed differences between forest stands with and without successful Tsuga canadensis regeneration. The two forest stands exhibiting successful Tsuga canadensis regeneration were located above 1,000 m in elevation on well-drained, moderately steep slopes and had the greatest canopy openness. Structural data from these two forest stands indicated a history of more continuous Tsuga canadensis regeneration. We also constructed disturbance chronologies detailing the history of canopy response to disturbance events and related these to Tsuga canadensis regeneration within each forest stand. Student t-tests adjusted for unequal variances indicated significant differences in the number of release events per tree between forest stands with and without successful Tsuga canadensis regeneration. While forest stands with successful Tsuga canadensis regeneration were more frequently disturbed by minor to major canopy disturbances, events of moderate intensity were found to be most significant in terms of regeneration. These data will be of value to land managers maintaining stands of Tsuga canadensis where treatment for hemlock woolly adelgid infestation has been successful. In areas where treatment is impractical or unsuccessful, land managers will be able to use these data to restore Tsuga canadensis forests after the wave of hemlock woolly adelgid induced mortality has passed.
Similar content being viewed by others
References
Abrams MD, Orwig DA (1996) A 300-year history of disturbance and canopy recruitment for co-occurring white pine and hemlock on the Allegheny Plateau, USA. J Ecol 84:353–363
Abrams MD, van de Gevel S, Dodson RC, Copenheaver CA (2000) The dendroecology and climatic impacts for old-growth white pine and hemlock on the extreme slopes of the Berkshire Hills, Massachusetts, U.S.A. Can J Bot 78:851–861
Barnes BV, Zak DR, Denton SR, Spurr SH (1998) Forest ecology. John Wiley and Sons, Inc., New York
Bormann FH, Likens GE (1979) Pattern and process in a forested ecosystem. Springer-Verlag, New York
Busing RT (2005) Tree mortality, canopy turnover, and woody detritus in old cove forests of the southern Appalachian Mountains. Ecology 86:73–84
Clark JS, Royall PS (1995) Transformations of a northern hardwood forest by aboriginal (Iroquois) fire: charcoal evidence from Crawford Lake, Ontario, Canada. Holocene 5:1–9
Clark SL, Hallgren SW (2004) Age estimation of Quercus marilandica and Quercus stellata: applications for interpreting stand dynamics. Can J For Res 34:1353–1358
Davis HC (1999) Reflections of Cataloochee Valley and its vanished people in the Great Smoky Mountains. Hattie Caldwell Davis, Maggie Valley
Dobbs MM, Parker AJ (2004) Evergreen understory dynamics in Coweeta Forest, North Carolina. Phys Geogr 25:481–498
Frelich LE, Lorimer CG (1991) Natural disturbance regimes in hemlock-hardwood forests of the upper Great Lakes region. Ecol Monogr 61:145–164
Frelich LE, Graumlich LJ (1994) Age class distribution and spatial patterns in an old- growth hemlock-hardwood forest. Can J For Res 24:1939–1947
Frelich LE (2002) Forest dynamics and disturbance regimes. Cambridge University Press, Cambridge
Grissino-Mayer HD (2001) Evaluating crossdating accuracy: a manual and tutorial for the computer program COFECHA. Tree-Ring Res 57:205–221
Harmon ME, Bratton SP, White PS (1983) Disturbance and vegetation response in relation to environmental gradients in the Great Smoky Mountains. Vegetatio 55:129–139
Hart JL, Shankman D (2005) Disjunct eastern hemlock (Tsuga canadensis) stands at its southern range boundary. J Torrey Bot Soc 132:602–612
Hedman CW, Van Lear DH (1995) Vegetative structure and composition of southern Appalachian riparian forests. Bull Torrey Bot Club 122:134–144
Hett JM, Loucks OL (1976) Age structure models of balsam fir and eastern hemlock. J Ecol 64:1029–1044
Holmes RL (1983) Computer-assisted quality control in tree-ring dating and measurement. Tree-Ring Bull 43:69–78
Houk R (1993) Great Smoky Mountains. Houghton Mifflin Company, New York
Johnson EA, Fryer JI (1989) Population dynamics in Lodgepole pine-Engelmann spruce forests. Ecology 70:1335–1345
Kavanagh K, Kellman M (1986) Performance of Tsuga canadensis (L.) Carr. at the centre and northern edge of its range: a comparison. J Biogeogr 13:145–157
Kemp S (1998) Trees of the smokies. Great Smoky Mountains Natural History Association, Gatlinburg
Kincaid JA (2008) Spatial structure and environmental controls of Tsuga canadensis (L.) Carr. regeneration in the southern Appalachian Mountains (submitted)
Knebel LG (1999) Changes in old growth hemlock forests of the Cataloochee watershed in the Great Smoky Mountains National Park: preparing for the arrival of the hemlock woolly adelgid (Adelges tsugae). Masters Thesis, Western Carolina University
Lorimer CG (1977) The presettlement forest and natural disturbance cycle in northeastern Maine. Ecology 58:139–148
Lorimer CG (1980) Age structure and disturbance history of a southern Appalachian virgin forest. Ecology 61:1169–1184
Lorimer CG, Krug AG (1983) Diameter distributions in even-aged stands of shade-tolerant and mid-tolerant tree species. Am Midl Nat 109:331–345
Lorimer CG, Frelich LE (1989) A methodology for estimating canopy disturbance frequency and intensity in dense temperate forests. Can J For Res 19:651–663
Lorimer CG (1995) Dynamics and structural characteristics of eastern hemlock stands. In: Mroz G, Martin J (eds) Hemlock ecology and management: proceedings of a regional conference on ecology and management of eastern hemlock. University of Wisconsin, Madison, pp 43–59
Madden M, Welch R, Jordan T, Jackson P, Seavey R, Seavey J (2004) Digital vegetation maps for the Great Smoky Mountains National Park. Final report submitted to USDI National Park Service, Great Smoky Mountains National Park, Cooperative Agreement No. 1443-CA-5460-98-019, pp 1–44
McLachlan JS, Foster DR, Menalled F (2000) Anthropogenic ties to late-successional structure and composition in four New England hemlock stands. Ecology 81:717–733
Mladenoff DJ, Stearns F (1993) Eastern hemlock regeneration and deer browsing in the northern Great Lakes region: a re-examination and model simulation. Conserv Biol 7:889–900
Monk CD, McGinty DT, Day FP (1985) The ecological importance of Kalmia latifolia and Rhododendron maximum in the deciduous forests of the southern Appalachians. Bull Torrey Bot Club 112:87–193
Nienstaedt H, Olson JS (1955) Heredity and environment: short-cut study shows how both affect hemlock growth. Front Plant Sci 7:7
North Carolina State Climate Office (2006) Available via http://www.ncclimate.ncsu.edu/cronos/normals.php?station=311564. Cited July 2006
NOAA (2006) Climate visualization website. Available via http://www.ncdc.noaa.gov/oa/climate/onlineprod/drought/xmgrg3.html. Cited July 2006
Nowacki GJ, Abrams MD (1997) Radial-growth averaging criteria for reconstructing disturbance histories from presettlement-origin oaks. Ecol Monogr 67:225–249
Orwig DA, Cogbill CV, Foster DR, O’Keefe JF (2001) Variations in old-growth structure and definitions: forest dynamics on Wachusett Mountain, Massachusetts. Ecol Appl 11:437–452
Parker AJ (1982) The topographic relative moisture index: an approach to soil-moisture assessment in mountain terrain. Phys Geogr 3:60–169
Parker AJ (1993) Structural variation and dynamics of lodgepole pine forests in Lassen Volcanic National Park, California. Ann Assoc Am Geogr 83:613–629
Parker AJ, Parker KC, McCay DH (2001) Disturbance-mediated variation in stand structure between varieties of Pinus clausa (sand pine). Ann Assoc Am Geogr 91:28–47
Parshall T (1995) Canopy mortality and stand-scale change in a northern hemlock-hardwood forest. Can J For Res 25:1466–1478
Phipps RL (1985) Collecting, preparing, crossdating, and measuring increment cores. US Geological Survey Report 85-4148
Ploucher AE, Carvell KL (1987) Population dynamics of rosebay rhododendron thickets in the southern Appalachian Mountains. Bull Torrey Bot Club 114:121–126
Pyle C (1988) The type and extent of anthropogenic vegetation disturbance in the Great Smoky Mountains before National Park acquisition. Castanea 53:183–196
Rankin WT, Tramer EJ (2002) The gap dynamics of canopy trees of a Tsuga canadensis forest community. Northeast Nat 9:391–406
Rogers RS (1978) Forests dominated by hemlock (Tsuga canadensis): distribution as related to site and postsettlement history. Can J Bot 56:843–854
Rooney TP, Waller DM (1998) Local and regional variation in hemlock seedling establishment in forests of the upper Great Lakes region, USA. For Ecol Manage 111:211–224
Rooney TP, McCormick RJ, Solheim SL, Waller DM (2000) Regional variation in recruitment of hemlock seedlings and saplings in the upper Great Lakes, USA. Ecol Appl 10:1119–1132
Runkle JR (1981) Gap regeneration in some old-growth forests of the eastern United States. Ecology 62:1041–1051
Runkle JR (1982) Patterns of disturbance in some old-growth mesic forests of eastern North America. Ecology 63:1533–1546
Runkle JR, Yetter TC (1987) Treefalls revisited: gap dynamics in the southern Appalachians. Ecology 68:417–424
R-Well (2005) Desktop mapping system (DMS) 5.0. R-WEL, Inc., Athens
Shanks RE (1954) Climates of the Great Smoky Mountains. Ecology 35:354–361
Singer MT, Lorimer CG (1997) Crown release as a potential old-growth restoration approach in northern hardwoods. Can J For Res 27:1222–1232
StataCorp (2003) Stata statistical software: release 8.0. Stata Corporation, College Station
Stokes MA, Smiley TL (1996) An introduction to tree-ring dating. The University of Arizona Press, Tucson
Tyrell LE, Crow TR (1994) Structural characteristics of old-growth hemlock-hardwood forests in relation to age. Ecology 75:370–386
Van Lear DH, Vandermast DB, Rivers CT, Baker TT, Hedman CW, Clinton DB, Waldrop TA (2002) American chestnut, rhododendron, and the future of Appalachian cove forests. In: Outcalt KW (ed) Proceedings of the eleventh biennial southern silvicultural research conference. USDA Forest Service, Southern Research Station, General Technical Report SRS-48, pp 214–220
Waller DM, Alverson WS, Solheim SL (1996) Local and regional factors influencing the regeneration of eastern hemlock. In: Mroz G, Martin J (eds) Hemlock ecology and management: proceedings of a regional conference on ecology and management of eastern hemlock. Department of Forestry, Michigan Technical University, Houghton, pp 73–90
Webster CR, Lorimer CG (2002) Single-tree versus group selection in hemlock-hardwood forests: are smaller openings less productive? Can J For Res 32:591–604
White PS (1979) Pattern, process, and natural disturbance in vegetation. Bot Rev 45:229–299
White PS (1996) Wildflowers of the smokies. Great Smoky Mountains Natural History Association, Gatlinburg
Whitney GG (1990) The history and status of the hemlock-hardwood forests of the Allegheny Plateau. J Ecol 78:443–458
Whittaker RH (1956) Vegetation of the Great Smoky Mountains. Ecol Monogr 26:1–80
Woods KD (2000a) Dynamics of late-successional hemlock-hardwood forests over three decades. Ecology 81:110–126
Woods KD (2000b) Long-term change in spatial pattern in a late-successional hemlock-hardwood forest. J Ecol 88:267–282
Woods FW, Shanks RE (1959) Natural replacement of chestnut by other species in the Great Smoky Mountains National Park. Ecology 40:349–361
Yamaguchi DK (1991) A simple method for cross-dating increment cores from living trees. Can J For Res 21:414–416
Yost EC, Johnson KS, Blozan WF (1994) Old-growth project: stand delineation and disturbance rating, Great Smoky Mountains National Park. Technical Report NPS/SERGSM/NATR, United States Department of the Interior, National Park Service
Ziegler SS (2000) A comparison of structural characteristics between old-growth and postfire second-growth hemlock-hardwood forests in Adirondack Park, New York, U.S.A. Glob Ecol Biogeogr 9:373–389
Ziegler SS (2002) Disturbance regimes of hemlock-dominated old-growth forests in northern New York, U.S.A. Can J For Res 32:2106–2115
Acknowledgments
This research was supported by a Carlos C. Campbell Memorial Fellowship from the Great Smoky Mountains Conservation Association. We would like to thank Kristine Johnson and Mike Jenkins of GSMNP for information regarding the location of Tsuga candensis forest stands within the park. We thank Paul Knapp of the Department of Geography, University of North Carolina-Greensboro for access to his tree-ring laboratory. For technical and graphical assistance, we thank Joel Sabin, Thomas Jordan, and Heather Phares. Three anonymous reviewers provided suggestions that greatly improved the manuscript.
Author information
Authors and Affiliations
Corresponding author
Additional information
As of August 2008, Joshua A. Kincaid will be a member of the Environmental Studies program at Shenandoah University in Winchester, Virginia, USA
Rights and permissions
About this article
Cite this article
Kincaid, J.A., Parker, A.J. Structural characteristics and canopy dynamics of Tsuga canadensis in forests of the southern Appalachian Mountains, USA. Plant Ecol 199, 265–280 (2008). https://doi.org/10.1007/s11258-008-9431-0
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11258-008-9431-0