Climate change is projected to profoundly influence vegetation patterns and community compositions, either directly through increased species mortality and shifts in species distributions or indirectly through disturbance dynamics such as increased wildfire activity and extent, shifting fire regimes, and pathogenesis. Mountainous landscapes have been shown to be particularly sensitive to climate changes and are likely to experience significant impacts under predicted future climate regimes. Western white pine ( Pinus monticola), a five-needle pine species that forms the most diverse of the white pine forest cover types in the western United States, is vulnerable to an interacting suite of threats that includes climate change, fire suppression, white pine blister rust ( Cronartium ribicola), and mountain pine beetles ( Dendroctonus ponderosae) that have already caused major changes in species distribution and abundance. We used the mechanistic simulation model FireBGCv2 to simulate effects of climate change and fire management on western white pines in a mountainous watershed in Glacier National Park, Montana, USA. Our results suggest that warming temperatures favor increased abundance of western white pine over existing climax and shade tolerant species in the study area, mainly because warmer conditions potentiate fire dynamics, including increased wildfire frequency and extent, which facilitates regeneration. Suppression of wildfires reduced the area dominated by western white pine, but fire suppression was less effective at limiting burned area extent and fire frequency in a warmer and drier climate. Wildfires created canopy gaps that allowed for western white pine regeneration at a high enough rate to escape local extirpation from white pine blister rust. Western white pine appears to be a resilient species even under fairly extreme warming trajectories and shifting fire regimes, and may provide a hedge against vegetation community shifts away from forest types and toward grass and shrublands.
References
[1]
Lenihan, J.M.; Drapek, R.; Bachelet, D.; Neilson, R.P. Climate change effects on vegetation distribution, carbon, and fire in California. Ecol. Appl. 2003, 13, 1667–1681.
[2]
Bentz, B.; Régnière, J.; Fettig, C.; Hansen, E.; Hayes, J.; Hicke, J.; Kelsey, R.; Negrón, J.; Seybold, S. Climate change and bark beetles of the western United States and Canada: Direct and indirect effects. BioScience 2010, 60, 602–613.
[3]
McKenzie, D.; Gedalof, Z.; Peterson, D.L.; Mote, P. Climatic change, wildfire, and conservation. Conserv. Biol. 2004, 18, 890–902.
[4]
Flannigan, M.; Stocks, B.; Wotton, B. Climate change and forest fires. Sci. Total Environ. 2000, 262, 221–229.
Fagre, D.B.; Peterson, D.L. Ecosystem dynamics and disturbance in mountain wildernesses: Assessing vulnerability of natural resources to change. In Wilderness Science in a Time of Change, Vol. 3: Wilderness as a Place for Scientific Inquiry; McCool, S.F., Cole, D.N., Borrie, W.T., O'Loughlin, J., Eds.; Rocky Mountain Research Station, USDA Forest Service: Ogden, UT, USA, 2000; pp. 74–81.
[7]
Rehfeldt, G.E.; Crookston, N.L.; Warwell, M.V.; Evans, J.S. Empirical analyses of plant-climate relationships for the Western United States. Int. J. Plant Sci. 2006, 167, 1123–1150.
[8]
Van Mantgem, P.J.; Stephenson, N.L.; Byrne, J.C.; Daniels, L.D.; Franklin, J.F.; Fule, P.Z.; Harmon, M.E.; Larson, A.J.; Smith, J.M.; Taylor, A.H.; Veblen, T.T. Widespread increase of tree mortality rates in the Western United States. Science 2009, 323, 521–524.
[9]
Tomback, D.; Achuff, P. Blister rust and western forest biodiversity: Ecology, values and outlook for white pines. For. Pathol. 2010, 40, 186–225.
[10]
Keane, R.E.; Loehman, R.A.; Holsinger, L.M. The FireBGCv2 Landscape Fire and Succession Model: A Research Simulation Platform for Exploring Fire and Vegetation Dynamics; US Department of Agriculture, Forest Service, Rocky Mountain Research Station: Fort Collins, CO, USA, 2011; p. 137.
[11]
Bowman, D.M.J.S.; Balch, J.K.; Artaxo, P.; Bond, W.J.; Carlson, J.M.; Cochrane, M.A.; D'Antonio, C.M.; DeFries, R.S.; Doyle, J.C.; Harrison, S.P. Fire in the earth system. Science 2009, 324, 481–484.
[12]
Westerling, A.L.; Turner, M.G.; Smithwick, E.A.H.; Romme, W.H.; Ryan, M.G. Continued warming could transform Greater Yellowstone fire regimes by mid-21st century. Proc. Natl. Acad. Sci. USA 2011, 108, 13165–13170.
[13]
Keane, R.E.; Veblen, T.; Ryan, K.C.; Logan, J.; Allen, C.; Hawkes, B. The cascading effects of fire exclusion in the Rocky Mountains. In Rocky Mountain Futures: An Ecological Perspective; Baron, J., Ed.; Island Press: Washington, DC, USA, 2002; pp. 133–153.
[14]
Millar, C.I.; Westfall, R.D.; Delany, D.L.; King, J.C.; Graumlich, L.J. Response of subalpine conifers in the Sierra Nevada, California, USA, to 20th-century warming and decadal climate variability. Arct. Antarct. Alp. Res. 2004, 36, 181–200.
[15]
Climate Change 2007: Synthesis Report. Contribution of Working Groups I, II, and III to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change; IPCC: Geneva, Switzerland, 2007; p. 104.
[16]
Littell, J.; McKenzie, D.; Peterson, D.; Westerling, A. Climate and wildfire area burned in western US ecoprovinces, 1916–2003. Ecol. Appl. 2009, 19, 1003–1021.
[17]
Westerling, A.L.; Hidalgo, H.G.; Cayan, D.R.; Swetnam, T.W. Warming and earlier spring increase Western US forest wildfire activity. Science 2006, 313, 940–943.
[18]
Heyerdahl, E.K.; McKenzie, D.; Daniels, L.D.; Hessl, A.E.; Littell, J.S.; Mantua, N.J. Climate drivers of regionally synchronous fires in the inland Northwest (1651-1900). Int. J. Wildland Fire 2008, 17, 40–49.
[19]
Schoennagel, T.; Veblen, T.T.; Romme, W.H. The interaction of fire, fuels, and climate across Rocky Mountain forests. BioScience 2004, 54, 661–676.
[20]
Heimann, M.; Reichstein, M. Terrestrial ecosystem carbon dynamics and climate feedbacks. Nature 2008, 451, 289–292.
[21]
Boisvenue, C.; Running, S.W. Simulations show decreasing carbon stocks and potential for carbon emissions in Rocky Mountain forests over the next century. Ecol. Appl. 2010, 20, 1302–1319.
[22]
Giorgi, F.; Mearns, L.O. Calculation of average, uncertainty range, and reliability of regional climate changes from AOGCM simulations via the “reliability ensemble averaging” (REA) method. J. Clim. 2002, 15, 1141–1158.
[23]
Christensen, J.H.; Hewitson, B.; Busuioc, A.; Chen, A.; Gao, X.; Held, I.; Jones, R.; Kolli, R.K.; Kwon, W.-T.; Laprise, R.; et al. Regional climate projections. In Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change; Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K.B., Tignor, M., Miller, H.L., Eds.; Cambridge University Press: Cambridge, UK; New York, NY, USA, 2007; pp. 848–940.
[24]
Flannigan, M.D.; Amiro, B.D.; Logan, K.A.; Stocks, B.J.; Wotton, B.M. Forest fires and climate change in the 21st century. Mitig. Adapt. Strat. Glob. Chang. 2006, 11, 847–859.
[25]
Torn, M.S.; Fried, J.S. Predicting the impacts of global warming on wildland fire. Clim. Chang. 1992, 21, 257–274.
Kinloch, B.B., Jr. White pine blister rust in North America: Past and prognosis. Phytopathology 2003, 93, 1044–1047.
[29]
Kim, M.S.; Richardson, B.A.; McDonald, G.I.; Klopfenstein, N.B. Genetic diversity and structure of western white pine (Pinus monticola) in North America: A baseline study for conservation, restoration, and addressing impacts of climate change. Tree Genet. Genomes 2010, 7, 11–21.
[30]
Fowells, H.A. Silvics of Forest Trees of the United States; US Department of Agriculture: Washington, DC, USA, 1965; p. 762.
[31]
Little, E.L., Jr. Digital Representation of “Atlas of United States Trees”. U.S. Geological Survey Professional Paper 1650; U.S. Geological Survey: Reston, VA, USA, 1999.
[32]
Little, E.L., Jr. Atlas of United States Trees: Volume 1. Conifers and Important Hardwoods. US Department Agriculture, Miscellaneous Publication No. 1146; US Government Printing Office: Washington, DC, USA, 1971.
[33]
Graham, R.T. Pinus monticola Dougl ex D. Don, Western White Pine. In Silvics of North America; Conifers; Burns, R.M., Honkala, B.H., Eds.; US Department of Agriculture, Forest Service: Washington, DC, USA, 1990; Volume 1, pp. 385–394.
[34]
Griffith, R.S. Pinus Monticola. In Fire Effects Information System; US Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory: Washington, DC, USA, 1992.
[35]
Habeck, J.R. Forest succession in the Glacier Park cedar-hemlock forests. Ecology 1968, 49, 872–880.
[36]
Habeck, J.R. The Vegetation of Glacier National Park, Montana. Final Report on file at Glacier National Park; US Department of Interior National Park Service: West Glacier, MT, USA, 1970; p. 132.
[37]
Habeck, J.R.; Mutch, R.W. Fire-dependent forests in the northern Rocky Mountains. Quat. Res. 1973, 3, 408–424.
[38]
Harvey, A.E.; Byler, J.W.; McDonald, G.I.; Neuenschwander, L.F.; Tonn, J.R. Death of an Ecosystem: Perspectives on Western White Pine Ecosystems of North America at the End of the Twentieth Century; US Department of Agriculture, Forest Service, Rocky Mountain Research Station: Fort Collins, CO, USA, 2008; p. 10.
[39]
Ryan, M.G.; Harmon, M.E.; Birdsey, R.A.; Giardina, C.P.; Heath, L.S.; Houghton, R.A.; Jackson, R.B.; McKinley, D.C.; Morrison, J.F.; Murray, B.C. A synthesis of the science on forests and Carbon for US Forests. Issues Ecol. 2010, 13, 1–16.
[40]
Zeglen, S.; Pronos, J.; Merler, H. Silvicultural management of white pines in western North America. For. Pathol. 2010, 40, 347–368.
[41]
Ketcham, D.E.; Wellner, C.A.; Evans, S.S. Western white pine management programs realigned on Northern Rocky Mountain National Forests. J. For. 1968, 66, 329–332.
[42]
Lowery, D. Western White Pine; US Department of Agriculture, Forest Service: Washington, DC, USA, 1984; p. 7.
[43]
Schwandt, J.W.; Lockman, I.B.; Kliejunas, J.T.; Muir, J.A. Current health issues and management strategies for white pines in the western United States and Canada. For. Pathol. 2010, 40, 226–250.
[44]
Samman, S.; Schwandt, J.W.; Wilson, J.L. Managing for Healthy White Pine Ecosystems in the United States to Reduce the Impacts of White Pine Blister Rust; US Department of Agriculture, Forest Service: Missoula, MT, USA, 2003; p. 10.
[45]
Harvey, A.E.; Hessburg, P.F.; Byler, J.W.; McDonald, G.I.; Weatherby, J.C.; Wickman, B.E. Health declines in western interior forests: Symptoms and solutions. Proceedings of Ecosystem Management in Western Interior Forests, Spokane, WA, USA, 3–5 May 1994; Cooperative Extension, Washington State University: Pullman, WA, USA, 1994.
[46]
McDonald, G.I.; Hoff, R.J.; Tomback, D.F.; Arno, S.F.; Keane, R.E. Blister rust: An introduced plague. In Whitebark Pine Communities. Ecology and Restoration; Tomback, D.F., Arno, S.F., Keane, R.E., Eds.; Island Press: Washington, DC, USA, 2001; pp. 193–220.
[47]
Maloy, O.C. White pine blister rust control in North America: A case history. Annu. Rev. Phytopathol. 1997, 35, 87–109.
[48]
Zambino, P.J. Biology and pathology of ribes and their implications for management of white pine blister rust. For. Pathol. 2010, 40, 264–291.
[49]
Graham, R.T.; Harvey, A.E.; Jain, T.B.; Tonn, J.R. The Effects of Thinning and Similar Stand Treatments on Fire Behavior in Western Forests. General Technical Report PNW-GTR-463; Pacific Northwest Research Station, USDA Forest Service: Washington, DC, USA, 1999.
[50]
Keane, R.E.; Morgan, P.; White, J.D. Temporal patterns of ecosystem processes on simulated landscapes in Glacier National Park, Montana, USA. Landsc. Ecol. 1999, 14, 311–329.
[51]
Keane, R.E.; Ryan, K.; Finney, M. Simulating the consequences of fire and climate regimes on a complex landscape in Glacier National Park, USA. Tall Timbers 1998, 20, 310–324.
[52]
Keane, R.E.; Ryan, K.C.; Running, S.W. Simulating effects of fire on northern Rocky Mountain landscapes with the ecological process model FIRE-BGC. Tree Physiol. 1996, 16, 319–331.
[53]
Running, S.W.; Hunt, E.R., Jr. Generalization of a forest ecosystem process model for other biomes, BIOME-BGC, and an application for global-scale models. In Scaling Physiological Processes: Leaf to Globe; Ehleringer, J.R., Ed.; Academic Press: London, UK, 1993; pp. 141–158.
[54]
Running, S.W.; Coughlan, J.C. A general model of forest ecosystem processes for regional applications I. Hydrologic balance, canopy gas exchange and primary production processes. Ecol. Model. 1988, 42, 125–154.
[55]
McDonald, G.I.; Hoff, R.J.; Wykoff, W. Computer Simulation of White Pine Blister Rust Epidemics: Model Formulation; US Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: Washington, DC, USA, 1981.
[56]
Howell, B.; Burns, K.S.; Kearns, H.S.J.; Witcosky, J.J.; Cross, F.J. Biological evaluation of a model for predicting presence of white pine blister rust in Colorado based on climatic variable and susceptible white pine species distribution. Biological Evaluation R2-06-04. 2006, 1–15.
[57]
King, J.N.; David, A.; Noshad, D.; Smith, J. A review of genetic approaches to the management of blister rust in white pines. For. Pathol. 2010, 40, 292–313.
[58]
Finklin, A.I. A Climatic Handbook for Glacier National Park: With Data for Waterton Lakes National Park; US Department of Agriculture, Forest Service, Intermountain Research Station: Ogden, UT, USA, 1986; p. 55.
[59]
Habeck, J.; Mutch, R. Fire-dependent forests in the northern Rocky Mountains. Quat. Res. 1973, 3, 408–424.
[60]
Kessell, S.R. Gradient Modeling: Resource and Fire Management; DeSanto, R.S., Ed.; Springer-Verlag: New York, NY, USA, 1979; p. 432.
[61]
Habeck, J.R.; Choate, C.M. An analysis of Krummholz Communities at Logan Pass, Glacier National Park. Northwest Sci. 1963, 37, 165–166.
[62]
Barrett, S.W.; Arno, S.F.; Key, C.H. Fire regimes of western larch-lodgepole pine forests in Glacier National Park, Montana. Can. J. For. Res. 1991, 21, 1711–1720.
[63]
Barrett, S.W. Fire History of Glacier National Park: Middle Fork Flathead River Drainage, in National Park Service, Glacier National Park, West Glacier, MT; US Department of Agriculture, Forest Service, Intermountain Research Station: Missoula, MT, USA, 1986.
[64]
Habeck, J.R. Fire Ecology Investigations in Glacier National Park—Historical Considerations and Current Observations. Final Report on file at Glacier National Park; US Department of Interior National Park Service: West Glacier, MT, USA, 1970; p. 80.
[65]
NCDC. Daily Surface Data; US Department of Commerce National Climatic Data Center: Asheville, NC, USA, 2011.
[66]
Hungerford, R.D.; Nemani, R.R.; Running, S.W.; Coughlan, J.C. MTCLIM: A Mountain Microclimate Simulation Model; US Department of Agriculture, Forest Service, Intermountain Research Station: Ogden, UT, USA, 1989; pp. 1–52.
[67]
Running, S.W.; Nemani, R.R.; Hungerford, R.D. Extrapolation of synoptic meteorological data in mountainous terrain and its use for simulating forest evapotranspiration and photosynthesis. Can. J. For. Res. 1987, 17, 472–483.
[68]
Mote, P. CLIMET Downscaled HadCM 3 Model Output, SRES A2 and B2, Northern Rocky Mountain Region. 2003.
[69]
Nakicenovic, N.; Alcamo, J.; Davis, G.; de Vries, B.; Fenhann, J.; Gaffin, S.; Gregory, K.; Grubler, A.; Jung, T.Y.; Kram, T.; et al. Special Report on Emissions Scenarios: A Special Report of Working Group III of the Intergovernmental Panel on Climate Change; Cambridge University Press: Cambridge, UK, 2000; p. 612.
[70]
Hood, S.M. Mitigating Old Tree Mortality in Long-Unburned, Fire-Dependent Forests: A Synthesis; US Department of Agriculture, Forest Service, Rocky Mountain Research Station: Fort Collins, CO, USA, 2010; p. 71.
[71]
Stephens, S.L.; Ruth, L.W. Federal forest-fire policy in the United States. Ecol. Appl. 2005, 15, 532–542.
[72]
R Development Core Team. R: A Language and Environment for Statistical Computing; R Foundation for Statistical Computing: Vienna, Austria, 2010.
[73]
Morgan, P.; Heyerdahl, E.; Gibson, C. Multi-season climate synchronized forest fires throughout the 20th century, northern Rockies, USA. Ecology 2008, 89, 717–728.
[74]
Kitzberger, T.; Brown, P.; Heyerdahl, E.; Swetnam, T.; Veblen, T. Contingent Pacific–Atlantic Ocean influence on multicentury wildfire synchrony over western North America. Proc. Natl. Acad. Sci. USA 2007, 104, 543.
[75]
Heikkinen, R.; Luoto, M.; Araújo, M.; Virkkala, R.; Thuiller, W.; Sykes, M. Methods and uncertainties in bioclimatic envelope modelling under climate change. Prog. Phys. Geogr. 2006, 30, 751.
[76]
Hampe, A. Bioclimate envelope models: What they detect and what they hide. Glob. Ecol. Biogeogr. 2004, 13, 469–471.
[77]
Sinclair, S.J.; White, M.D.; Newell, G.R. How useful are species distribution models for managing biodiversity under future climates? Ecol. Soc. 2010, 15, 8.
[78]
Millar, C.I.; Stephenson, N.L.; Stephens, S.L. Climate change and forests of the future: Managing in the face of uncertainty. Ecol. Appl. 2007, 17, 2145–2151.
[79]
Schrag, A.M.; Bunn, A.G.; Graumlich, L.J. Influence of bioclimatic variables on tree-line conifer distribution in the Greater Yellowstone Ecosystem: Implications for species of conservation concern. J. Biogeogr. 2008, 35, 698–710.
[80]
Perry, G.L.W.; Millington, J.D.A. Spatial modelling of succession-disturbance dynamics in forest ecosystems: Concepts and examples. Perspect. Plant Ecol. Evol. Syst. 2008, 9, 191–210.
[81]
Taylor, A.R.; Chen, H.Y.H.; van Damme, L. A review of forest succession models and their suitability for forest management planning. For. Sci. 2009, 55, 23–36.
[82]
He, H.S.; Keane, R.E.; Iverson, L.R. Forest landscape models, a tool for understanding the effect of the large-scale and long-term landscape processes. For. Ecol. Manag. 2008, 254, 371–374.
[83]
Keane, R.E.; Cary, G.J.; Davies, I.D.; Flannigan, M.D.; Gardner, R.H.; Lavorel, S.; Lenihan, J.M.; Li, C.; Rupp, T.S. A classification of landscape fire succession models: Spatial simulations of fire and vegetation dynamics* 1. Ecol. Model. 2004, 179, 3–27.
Littell, J.S.; McKenzie, D.; Peterson, D.L.; Westerling, A.L. Climate and wildfire area burned in western US ecoprovinces, 1916–2003. Ecol. Appl. 2009, 19, 1003–1021.
[86]
Whitlock, C.; Shafer, S.L.; Marlon, J. The role of climate and vegetation change in shaping past and future fire regimes in the northwestern US and the implications for ecosystem management. For. Ecol. Manag. 2003, 178, 5–21.
[87]
Brown, T.J.; Hall, B.L.; Westerling, A.L. The impact of twenty-first century climate change on wildland fire danger in the western United States: An applications perspective. Clim. Chang. 2004, 62, 365–388.
[88]
Flannigan, M.; Stocks, B.; Turetsky, M.; Wotton, M. Impacts of climate change on fire activity and fire management in the circumboreal forest. Glob. Chang. Biol. 2008, 14, 1–12.
[89]
Graham, R.T.; Harvey, A.E.; Jain, T.B.; Tonn, J.R. The effects of thinning and similar stand treatments on fire behavior in Western forests. General Technical Report PNW-GTR-463, USDA Forest Service; Pacific Northwest Research Station: Portland, OR, USA, 1999.
[90]
Hall, M.H.P.; Fagre, D.B. Modeled climate-induced glacier change in Glacier National Park, 1850-2100. BioScience 2003, 53, 131–140.
[91]
Fagre, D.B.; Peterson, D.L.; Hessl, A.E. Taking the pulse of mountains: Ecosystem responses to climatic variability. Clim. Chang. 2003, 59, 263–282.
[92]
White, J.D.; Running, S.W.; Thornton, P.E.; Keane, R.E.; Ryan, K.C.; Fagre, D.B.; Key, C.H. Assessing simulated ecosystem processes for climate variability research at glacier national park, USA. Ecol. Appl. 1998, 8, 805–823.
[93]
Lenihan, J.M.; Bachelet, D.; Neilson, R.P.; Drapek, R. Response of vegetation distribution, ecosystem productivity, and fire to climate change scenarios for California. Clim. Chang. 2008, 87, 215–230.
[94]
Agee, J. Fire Ecology of Pacific Northwest Forests; Island Press: Washington, DC, USA, 1996; p. 493.
[95]
Schoettle, A.W.; Sniezko, R.A. Proactive intervention to sustain high-elevation pine ecosystems threatened by white pine blister rust. J. For. Res. 2007, 12, 327–336.
[96]
Gibson, C.E.; Morgan, P. Atlas of Digital Polygon Fire Extents for Idaho and Western Montana (1889-2003); US Department of Agriculture, Forest Service, Rocky Mountain Research Station: Fort Collins, CO, USA, 2009. Available online: http://www.fs.fed.us/rm/data_archive (accessed on 23 September 2011).
[97]
Iverson, L.R.; Prasad, A.M.; Matthews, S.N.; Peters, M. Estimating potential habitat for 134 eastern US tree species under six climate scenarios. For. Ecol. Manag. 2008, 254, 390–406.
[98]
Morin, X.; Thuiller, W. Comparing niche-and process-based models to reduce prediction uncertainty in species range shifts under climate change. Ecology 2009, 90, 1301–1313.
[99]
Walck, J.L.; Hidayati, S.N.; Dixon, K.W.; Thompson, K.E.N.; Poschlod, P. Climate change and plant regeneration from seed. Glob. Chang. Biol. 2011, 17, 2145–2161.
[100]
Pilkey, O.H.; Pilkey-Jarvis, L. Useless Arithmetic: Why Environmental Scientists can't Predict the Future; Columbia University Press: New York, NY, USA, 2007.
[101]
Noss, R.F. Beyond Kyoto: Forest management in a time of rapid climate change. Conserv. Biol. 2001, 15, 578–590.