Awareness of geographic patterns and stand variables that influence tree growth will help forest managers plan appropriate management and monitoring strategies. We quantified influences of stand location, species composition, stand density, and tree size on aspen tree growth and vigor around the Lake Tahoe Basin in the Sierra Nevada Mountains of California and Nevada, USA. Radial growth data were taken from increment cores. Aspen trees on the south and west sides of the lake grew 20–25% faster than aspen in north and east side stands. Diameter growth at 2,400?m elevation was 58% of growth at 1,900?m near lake level. Aspen grew faster with less competition from neighbor trees. At any level of competition, aspen growth was slower beside conifer neighbors and correlated with crown ratio (CR: length of live crown relative to total tree height, a proxy for tree vigor). Analysis of independent CR data for 707 aspen trees in nine additional stands indicated that aspen had smaller crowns in the presence of greater competition, and that composition of neighbor trees also affected CR: aspen trees had shorter crowns in the presence of conifer at higher stand densities. Taken collectively, our analyses point towards a cascading decline in aspen growth and vigor incited by succession of aspen stands to conifers. Our findings suggest that conifer removal and stand density control in aspen-conifer stands at Lake Tahoe will enhance aspen growth and vigor. 1. Introduction Quaking aspen (Populus tremuloides Michx.) communities are being replaced by conifers throughout many forests and rangelands of North America [1–9]. Succession to coniferous species is expected in the absence of disturbances such as wildfires that kill young conifers establishing within aspen stands [10]. Pioneering features such as lightweight seed, shade intolerance, and rapid growth of vegetative root-sucker regeneration indicate that aspen is adapted to disturbance [10, 11]. Having the widest distribution of any native tree in North America suggests that aspen has benefitted from a long history of natural disturbances. Fire suppression throughout much of the 20th Century has lengthened fire return intervals, allowing conifers time to establish and develop thicker fire-resistant bark with advancing size and age. Conifers eventually overtop aspen and suppress aspen regeneration and herbaceous vegetation. Loss of aspen forest area merits concern because aspen is considered a keystone species [12, 13], and aspen is the foundation species in stands that are “hotspots” of biodiversity [14]. At the
References
[1]
W. D. Shepperd, D. L. Bartos, and S. A. Mata, “Above- and below-ground effects of aspen clonal regeneration and succession to conifers,” Canadian Journal of Forest Research, vol. 31, no. 5, pp. 739–745, 2001.
[2]
T. G. Wall, R. F. Miller, and T. J. Svejcar, “Juniper encroachment into aspen in the northwest Great Basin,” Journal of Range Management, vol. 54, no. 6, pp. 691–698, 2001.
[3]
A. L. Gallant, A. J. Hansen, J. S. Councilman, D. K. Monte, and D. W. Betz, “Vegetation dynamics under fire exclusion and logging in a Rocky Mountain watershed, 1856–1996,” Ecological Applications, vol. 13, no. 2, pp. 385–403, 2003.
[4]
A. P. Di Orio, R. Callas, and R. J. Schaefer, “Forty-eight year decline and fragmentation of aspen (Populus tremuloides) in the South Warner Mountains of California,” Forest Ecology and Management, vol. 206, no. 1–3, pp. 307–313, 2005.
[5]
B. E. Jones, T. H. Rickman, A. Vazquez, Y. Sado, and K. W. Tate, “Removal of encroaching conifers to regenerate degraded aspen stands in the Sierra Nevada,” Restoration Ecology, vol. 13, no. 2, pp. 373–379, 2005.
[6]
M. W. Kaye, D. Binkley, and T. J. Stohlgren, “Effects of conifers and elk browsing on quaking aspen forests in the central Rocky Mountains, USA,” Ecological Applications, vol. 15, no. 4, pp. 1284–1295, 2005.
[7]
A. E. Smith and F. W. Smith, “Twenty-year change in aspen dominance in pure aspen and mixed aspen/conifer stands on the Uncompahgre Plateau, Colorado, USA,” Forest Ecology and Management, vol. 213, no. 1–3, pp. 338–348, 2005.
[8]
J. D. Bates, R. F. Miller, and K. W. Davies, “Restoration of quaking aspen woodlands invaded by western juniper,” Rangeland Ecology and Management, vol. 59, no. 1, pp. 88–97, 2006.
[9]
W. J. Calder, K. J. Horn, and S. B. St. Clair, “Conifer expansion reduces the competitive ability and herbivore defense of aspen by modifying light environment and soil chemistry,” Tree Physiology, vol. 31, no. 6, pp. 582–591, 2011.
[10]
D. A. Perala, “Quaking aspen (Populus tremuloides Michx.),” in Silvics of North America, I. I. Deciduous, R. M. Burns, and B. H. Honkala, Eds., no. 654, pp. 555–569, United States Department of Agriculture Handbook, 1990.
[11]
B. V. Barnes, “The clonal growth habit of American aspens,” Ecology, vol. 47, pp. 439–447, 1966.
[12]
D. L. Bartos, “Landscape dynamics of aspen and conifer forests,” in Sustaining Aspen in Western Landscapes: Symposium Proceedings, W. D. Shepperd, D. Binkley, D. L. Bartos, T. J. Stohlgren, and L. G. Eskew, Eds., pp. 5–14, U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, RMRS-P-18, Grand Junction, Colo, USA, 2001.
[13]
W. D. Shepperd, P. C. Rogers, D. Burton, and D. Bartos, “Ecology, biodiversity, management, and restoration of aspen in the Sierra Nevada,” U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, RMRS-GTR-178, Fort Collins, Colo, USA, 2006.
[14]
T. J. Stohlgren, D. Binkley, G. W. Chong et al., “Exotic plant species invade hot spots of native plant diversity,” Ecological Monographs, vol. 69, no. 1, pp. 25–46, 1999.
[15]
T. J. Kuhn, H. D. Safford, B. E. Jones, and K. W. Tate, “Aspen (Populus tremuloides) stands and their contribution to plant diversity in a semiarid coniferous landscape,” Plant Ecology, vol. 212, no. 9, pp. 1451–1463, 2011.
[16]
T. W. Richardson and S. K. Heath, “Effects of conifers on aspen-breeding bird communities in the Sierra Nevada,” Transactions of the Western Section of the Wildlife Society, vol. 40, pp. 68–81, 2004.
[17]
D. L. Bartos and R. B. Campbell, “Decline of quaking aspen in the interior west-examples from Utah,” Rangelands, vol. 20, no. 1, pp. 17–24, 1998.
[18]
D. Binkley, “Age distribution of aspen in Rocky Mountain National Park, USA,” Forest Ecology and Management, vol. 255, no. 3-4, pp. 797–802, 2008.
[19]
G. E. Dixon, “Essential FVS: a user’s guide to the forest vegetation simulator (Revised 2011),” Internal Report, U.S. Department of Agriculture, Forest Service, Forest Management Service Center, Fort Collins, Colo, USA, 2002.
[20]
C. E. Keyser and G. E. Dixon, “Western Sierra Nevada (WS) variant overview—forest vegetation simulator (Revised 2011),” Internal Report, U.S. Department of Agriculture, Forest Service, Forest Management Service Center, Fort Collins, Colo, USA, 2008.
[21]
R. Boone, J. Tardif, and R. Westwood, “Radial growth of oak and aspen near a coal-fired station, Manitoba, Canada,” Tree-Ring Research, vol. 60, no. 1, pp. 45–58, 2004.
[22]
E. H. Hogg, J. P. Brandt, and B. Kochtubajda, “Factors affecting interannual variation in growth of western Canadian aspen forests during 1951–2000,” Canadian Journal of Forest Research, vol. 35, no. 3, pp. 610–622, 2005.
[23]
SAS Institute Inc, SAS/STAT 9.1 User’s Guide, vol. 1–7, SAS Institute Inc, Cary, NC, USA, 2nd edition, 2004.
[24]
R. C. Littell, G. A. Milliken, W. W. Stroup, R. D. Wolfinger, and O. Schabenberger, SAS for Mixed Models, SAS Institute Inc, Cary, NC, USA, 2nd edition, 2006.
[25]
J. N. Long and T. W. Daniel, “Assessment of growing stock in uneven-aged stands,” Western Journal of Applied Forestry, vol. 5, pp. 93–96, 1990.
[26]
J. D. Shaw, “Application of stand density index to irregularly structured stands,” Western Journal of Applied Forestry, vol. 15, no. 1, pp. 40–42, 2000.
[27]
R. F. Gersonde and K. L. O'Hara, “Comparative tree growth efficiency in Sierra Nevada mixed-conifer forests,” Forest Ecology and Management, vol. 219, no. 1, pp. 95–108, 2005.
[28]
L. Dolislager, A. Lashgari, J. Pederson, and T. VanCuren, “Lake Tahoe Atmospheric Deposition Study (LTADS),” Final Report, Chapter 2: Atmospheric Processes. Staff Report, California Environmental Protection Agency Air Resources Board, Sacramento, Calif, USA, 2006.
[29]
E. H. Hogg, J. P. Brandt, and M. Michaelian, “Impacts of a regional drought on the productivity, dieback, and biomass of western Canadian aspen forests,” Canadian Journal of Forest Research, vol. 38, no. 6, pp. 1373–1384, 2008.
[30]
K. Brown, A. J. Hansen, R. E. Keane, and L. J. Graumlich, “Complex interactions shaping aspen dynamics in the Greater Yellowstone Ecosystem,” Landscape Ecology, vol. 21, no. 6, pp. 933–951, 2006.
[31]
V. D. Hipkins and J. H. Kitzmiller, “Genetic variation and clonal distribution of quaking aspen in the central Sierra Nevada,” Transactions of the Western Section of the Wildlife Society, vol. 40, pp. 32–44, 2004.
[32]
H. Temesgen, V. LeMay, and S. J. Mitchell, “Tree crown ratio models for multi-species and multi-layered stands of southeastern British Columbia,” Forestry Chronicle, vol. 81, no. 1, pp. 133–141, 2005.
[33]
M. R. Holdaway, “Modeling tree crown ratio,” Forestry Chronicle, vol. 10, pp. 451–455, 1986.
[34]
M. W. Ritchie and D. W. Hann, Equations for predicting height to crown base for fourteen tree species in southwest Oregon, Oregon State University Forest Research Laboratory Research Paper 50, Corvallis, Ore, USA, 1987.
[35]
R. K. Tew, “Root carbohydrate reserves in vegetative reproduction of aspen,” Forest Science, vol. 16, pp. 318–320, 1970.
[36]
N. V. DeByle and R. P. Winokur, Aspen: Ecology and Management in the Western United States, U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, GTR-RM-119, Fort Collins, Colo, USA, 1985.