Climatic and biophysical controls on conifer species distributions in mountain forests of Washington State, USA

Aim The purpose of this study was to quantify relationships between conifer species distributions and climatic and biophysical variables, in order to provide better insight into the potential for redistribution of species on the landscape in response to climatic change. Location Data are from 10,653 georeferenced sites in Washington State, USA, along a longitudinal gradient from west of the crest of the Cascade Range to the beginnings of the western slope of the Rocky Mountains, and across two physiographic provinces, the Northern Cascades, characterized by steep, rugged topography, and the Okanogan Highlands, presenting moderate slopes and broad rounded summits. Methods Tree data were drawn from the USDA Forest Service Area Ecology Program database, collected in mature, undisturbed stands. We compared simple climatic variables (annual temperature, growing-degree days, annual and seasonal precipitation) to biophysical variables (soil, hydrologic, and solar radiation) derived from climatic variables. Climatic and biophysical variables were taken from the output of climatological and hydrological simulation models and estimated for each plot in the tree database. Generalized linear models were used, for each of fourteen tree species, at multiple spatial extents, to estimate the probability of occurrence of that species as a function of climatic and biophysical predictors. Models were validated by a combination of bootstrapping and estimating receiver operating characteristic (ROC) curves. Results For the majority of species, we were able to fit variables representing both moisture and temperature gradients, and in all but a few cases these models identified a unimodal response of species occurrence to these gradients. In some cases the ecological/ environmental niche of a species had been clearly captured by the model, whereas in others a longer gradient in the predictor variable(s) would be needed. Responses of most species were consistent across three spatial scales.

[1]  Eric R. Ziegel,et al.  Generalized Linear Models , 2002, Technometrics.

[2]  F. Holtmeier,et al.  Regeneration of trees in the treeline ecotone: northern Finnish Lapland , 2003 .

[3]  J. Moen,et al.  Timberline patterns in northernmost Fennoscandia. Relative importance of climate and grazing , 1995 .

[4]  James A. Young,et al.  Seeds of woody plants in North America , 1993 .

[5]  Niklaus E. Zimmermann,et al.  Risks of global warming on montane and subalpine forests in Switzerland – a modeling study , 2000 .

[6]  M. Austin,et al.  Models for the analysis of species' response to environmental gradients , 2004, Vegetatio.

[7]  Mike P. Austin,et al.  The potential contribution of vegetation ecology to biodiversity research , 1999 .

[8]  S. Walsh,et al.  Snow-avalanche Paths: Conduits from the Periglacial-Alpine to the Subalpine-Depositional Zone , 1992 .

[9]  Francis Juanes,et al.  INFERRING ECOLOGICAL RELATIONSHIPS FROM THE EDGES OF SCATTER DIAGRAMS: COMPARISON OF REGRESSION TECHNIQUES , 1998 .

[10]  J. B. Benedict.  Rates of Tree‐Island Migration, Colorado Rocky Mountains, USA , 1984 .

[11]  Stéphane Mallat,et al.  Multifrequency channel decompositions of images and wavelet models , 1989, IEEE Trans. Acoust. Speech Signal Process..

[12]  Lisa J. Graumlich,et al.  A 1000-Year Record of Temperature and Precipitation in the Sierra Nevada , 1993, Quaternary Research.

[13]  John Bell,et al.  A review of methods for the assessment of prediction errors in conservation presence/absence models , 1997, Environmental Conservation.

[14]  K. K. Mohanty.  The wavelet transform for local image enhancement , 1997 .

[15]  Lynn M. Resler,et al.  Variability in an edaphic indicator in alpine tundra , 2002 .

[16]  Jerry F. Franklin,et al.  Invasion of Subalpine Meadows by Trees in the Cascade Range, Washington and Oregon , 1971 .

[17]  R. Neilson.  A Model for Predicting Continental‐Scale Vegetation Distribution and Water Balance , 1995 .

[18]  D. Peterson.  Climate, limiting factors and environmental change in high-altitude forests of Western North America , 1998 .

[19]  I. Moore,et al.  Digital terrain modelling: A review of hydrological, geomorphological, and biological applications , 1991 .

[20]  T. Virtanen,et al.  How can the dynamics of the tundra-taiga boundary be remotely monitored? , 2002, Ambio.

[21]  H. Bugmann,et al.  Impacts of interannual climate variability on past and future forest composition , 2000 .

[22]  O. Hoegh‐Guldberg,et al.  Ecological responses to recent climate change , 2002, Nature.

[23]  A. Taylor.  Forest expansion and climate change in the mountain hemlock (Tsuga mertensiana) zone , 1995 .

[24]  Martin Beniston,et al.  Climate change and its impacts on glaciers and permafrost in the Alps , 1998 .

[25]  R. Neilson,et al.  Climate Change Effects on Vegetation Distribution and Carbon Budget in the United States , 2001, Ecosystems.

[26]  L. Kullman.  Change and stability in the altitude of the birch tree-limit in the southern Swedish Scandes 1915-1975 , 1979 .

[27]  Thomas A. Spies,et al.  REGIONAL GRADIENT ANALYSIS AND SPATIAL PATTERN OF WOODY PLANT COMMUNITIES OF OREGON FORESTS , 1998 .

[28]  A. Lloyd.  Patterns and Processes of Treeline Forest Response to Late Holocene Climate Change in the Sierra Nevada, California , 1996 .

[29]  K. Hall,et al.  Zoogeomorphology in the Alpine: some observations on abiotic–biotic interactions , 2003 .

[30]  A. H. Murphy,et al.  A General Framework for Forecast Verification , 1987 .

[31]  D. Butler.  Geomorphic process-disturbance corridors: a variation on a principle of landscape ecology , 2001 .

[32]  C. E. Woodcock,et al.  Geostatistical Estimation of Resolution- Dependent Variance in Remotely Sensed Images , 1999 .

[33]  G. Malanson,et al.  Non-equilibrium geomorphic processes and patterns on avalanche paths in the northern Rocky Mountains, U.S.A , 1990 .

[34]  Hong S. He,et al.  Study of landscape change under forest harvesting and climate warming-induced fire disturbance , 2002 .

[35]  D. Bamber.  The area above the ordinal dominance graph and the area below the receiver operating characteristic graph , 1975 .

[36]  L. Graumlich,et al.  HOLOCENE DYNAMICS OF TREELINE FORESTS IN THE SIERRA NEVADA , 1997 .

[37]  S. Wood.  Holocene stratigraphy and chronology of Mountain Meadows, Sierra Nevada, California , 1975 .

[38]  Daniel B. Botkin,et al.  Sensitivity of Cool-Temperate Forests and their Fossil Pollen Record to Rapid Temperature Change , 1985, Quaternary Research.

[39]  David H. Peterson,et al.  1976 step in the Pacific climate: forty environmental changes between 1968-1975 and 1977-1984 , 1991 .

[40]  P. Wardle.  An explanation for alpine timberline , 1971 .

[41]  Stephen J. Walsh,et al.  Lithologic, Structural, and Topographic Influences on Snow-Avalanche Path Location, Eastern Glacier National Park, Montana , 1990 .

[42]  Shoshiro Minobe,et al.  Resonance in bidecadal and pentadecadal climate oscillations over the North Pacific: Role in climatic regime shifts , 1999 .

[43]  L. Kullman.  20th Century Climate Warming and Tree-limit Rise in the Southern Scandes of Sweden , 2001, Ambio.

[44]  R. Kauth,et al.  The tasselled cap - A graphic description of the spectral-temporal development of agricultural crops as seen by Landsat , 1976 .

[45]  J. Campbell.  Introduction to remote sensing , 1987 .

[46]  Michael S. Knowles,et al.  Threatened and endangered species geography: characteristics of hot spots in the conterminous United States , 1998 .

[47]  Franco Biondi,et al.  North Pacific Decadal Climate Variability since 1661 , 2001 .

[48]  J. Wiens.  Spatial Scaling in Ecology , 1989 .

[49]  Lynn M. Resler,et al.  Ecotones in Mountain Environments: Illustrating Sensitive Biogeographical Boundaries with Remotely Sensed Imagery in the Geography Classroom , 2003 .

[50]  Charles B. Halpern,et al.  Modeling the distributions of shrub species in Pacific northwest forests , 1999 .

[51]  W. J. Carper,et al.  The use of intensity-hue-saturation transformations for merging SPOT panchromatic and multispectral image data , 1990 .

[52]  R. Lawrence,et al.  Spatial Variation in Distribution and Growth Patterns of Old Growth Strip-Bark Pines , 2003 .

[53]  C.J.F. ter Braak,et al.  A Theory of Gradient Analysis , 2004 .

[54]  Joseph K. Berry,et al.  Landscape analysis of plant diversity , 1997, Landscape Ecology.

[55]  R. Heikkinen,et al.  On the recovery of mountain birch after Epirrita damage in Finnish Lapland, with a particular emphasis on reindeer grazing , 1995 .

[56]  P. Chesson,et al.  The Roles of Harsh and Fluctuating Conditions in the Dynamics of Ecological Communities , 1997, The American Naturalist.

[57]  Edward R. Cook,et al.  Methods of Dendrochronology , 1990 .

[58]  H. Shugart,et al.  A Systems Analysis of the Global Boreal Forest: Individual-tree-based models of forest dynamics and their application in global change research , 1992 .

[59]  W. Smith,et al.  Influence of Wind Exposure on Needle Desiccation and Mortality for Timberline Conifers in Wyoming, U.S.A. , 1983, Arctic and Alpine Research.

[60]  F. L. Pérez.  Plant-induced spatial patterns of surface soil properties near caulescent Andean rosettes , 1995 .

[61]  L. Kullman.  The Changing Face of the Alpine World , 2004 .

[62]  Clinton K. Williams,et al.  Forested plant associations of the Okanogan National Forest , 1983 .

[63]  T. P. Barnett,et al.  Causes of Decadal Climate Variability over the North Pacific and North America , 1994, Science.

[64]  R. Keane,et al.  Whitebark Pine Communities: Ecology and Restoration , 2001 .

[65]  F. Berninger,et al.  Impacts of climate change on the tree line. , 2002, Annals of botany.

[66]  D. H. Ashton,et al.  Plant Succession in a Timberline Depressed by Vulcanism in South-Central Chile , 1977 .

[67]  S. Payette,et al.  Growth of Black Spruce at Its Northern Range Limit in Arctic Quebec, Canada , 1994 .

[68]  Daryl Pregibon,et al.  Tree-based models , 1992 .

[69]  A. Lugo,et al.  Climate Change and Forest Disturbances , 2001 .

[70]  Sarah C. Goslee,et al.  Extending community ecology to landscapes , 2002 .

[71]  Eric P. Crist,et al.  A Physically-Based Transformation of Thematic Mapper Data---The TM Tasseled Cap , 1984, IEEE Transactions on Geoscience and Remote Sensing.

[72]  S. Walsh,et al.  Comparison Of Landsat Thematic Mapper Digital Enhancements Of Snow-avalanche Paths: Validation Through Gis/remote Sensing Integration , 1990, 10th Annual International Symposium on Geoscience and Remote Sensing.

[73]  S. Walsh,et al.  An overview of scale, pattern, process relationships in geomorphology: a remote sensing and GIS perspective , 1998 .

[74]  S. Short,et al.  The Northern Limit of Trees in Labrador: A Discussion , 1979 .

[75]  J. Agee,et al.  FIRE FREQUENCY IN THE INTERIOR COLUMBIA RIVER BASIN: BUILDING REGIONAL MODELS FROM FIRE HISTORY DATA , 2000 .

[76]  C. Kremenetski,et al.  The Holocene vegetation history of the Khibiny Mountains: implications for the post-glacial expansion of spruce and alder on the Kola Peninsula, northwestern Russia , 1999 .

[77]  R. Holmes,et al.  Tree-Ring Chronologies of Western North America: California, Eastern Oregon and Northern Great Basin with Procedures Used in the Chronology Development Work Including Users Manuals for Computer Programs COFECHA and ARSTAN , 1986 .

[78]  F. Stuart Chapin,et al.  Response of subarctic vegetation to transient climatic change on the Seward Peninsula in north‐west Alaska , 2000 .

[79]  S. Running,et al.  Generalization of a forest ecosystem process model for other biomes, Biome-BGC, and an application for global-scale models. Scaling processes between leaf and landscape levels , 1993 .

[80]  F. N. David,et al.  Principles and procedures of statistics. , 1961 .

[81]  Antoine Guisan,et al.  Predictive habitat distribution models in ecology , 2000 .

[82]  L. Graumlich.  Subalpine Tree Growth, Climate, and Increasing CO_2: An Assessment of Recent Growth Trends , 1991 .

[83]  Silvical characteristics of subalpine fir. , 1984 .

[84]  D. Elliott-Fisk.  The Stability of the Northern Canadian Tree Limit , 1983 .

[85]  A. Bunn,et al.  Using a Simulation Model to Compare Methods of Tree-Ring Detrending and to Investigate the Detectability of Low-Frequency Signals , 2004 .

[86]  M. Germino,et al.  Conifer seedling distribution and survival in an alpine-treeline ecotone , 2002, Plant Ecology.

[87]  C. Woodcock,et al.  Autocorrelation and regularization in digital images. II. Simple image models , 1989 .

[88]  David W. Pierce,et al.  Distinguishing coupled ocean–atmosphere interactions from background noise in the North Pacific , 2001 .

[89]  N. Huntly,et al.  Pikas (Ochotona princeps : Lagomorpha) as allogenic engineers in an alpine ecosystem , 1998, Oecologia.

[90]  G. Malanson,et al.  A HISTORY OF HIGH-MAGNITUDE SNOW AVALANCHES, SOUTHERN GLACIER NATIONAL PARK, MONTANA, U.S.A. , 1985 .

[91]  de S. Bruin,et al.  Spatial uncertainty in estimates of the areal extent of land cover types , 2000 .

[92]  L. Iverson,et al.  Global Change in Forests: Responses of Species, Communities, and Biomes , 2001 .

[93]  G. Malanson,et al.  Ordinations of species and fuel arrays and their use in fire management , 1985 .

[94]  A. D. Kennedy.  Antarctic Terrestrial Ecosystem Response to Global Environmental Change , 1995 .

[95]  Sakari Tuhkanen,et al.  Climatic parameters and indices in plant geography , 1980 .

[96]  T. Yee,et al.  Generalized additive models in plant ecology , 1991 .

[97]  Charles B. Halpern,et al.  Potential response of pacific northwestern forests to climatic change, effects of stand age and initial composition , 1993 .

[98]  J. R. Jensen.  Remote Sensing of the Environment: An Earth Resource Perspective , 2000 .

[99]  D. Butler.  Snow Avalanche Path Terrain and Vegetation, Glacier National Park, Montana , 1979, Arctic and Alpine Research.

[100]  D. Tomback,et al.  Delayed seed germination in whitebark pine and regeneration patterns following the Yellowstone fires , 2001 .

[101]  D. Cairns,et al.  EXAMINATION OF THE CARBON BALANCE HYPOTHESIS OF ALPINE TREELINE LOCATION IN GLACIER NATIONAL PARK, MONTANA , 1997 .

[102]  Daniel M. Johnson,et al.  Another perspective on altitudinal limits of alpine timberlines. , 2003, Tree physiology.

[103]  Annika Hofgaard,et al.  The dynamics of the tundra-taiga boundary: an overview and suggested coordinated and integrated approach to research. , 2002, Ambio.

[104]  F. Holtmeier,et al.  Mountain Timberlines—Ecology, Patchiness, And Dynamics , 2003 .

[105]  J. B. Benedict.  Frost Creep and Gelifluction Features: A Review , 1976, Quaternary Research.

[106]  G. MacDonald,et al.  Response of the Central Canadian Treeline to Recent Climatic Changes , 1998 .

[107]  C. Körner,et al.  A world‐wide study of high altitude treeline temperatures , 2004 .

[108]  M. Stokes,et al.  An Introduction to Tree-Ring Dating , 1996 .

[109]  C. Woodcock,et al.  Autocorrelation and regularization in digital images. I. Basic theory , 1988 .

[110]  Daniel R. Cayan,et al.  Interannual Climate Variability and Snowpack in the Western United States , 1996 .

[111]  J. Franklin.  Predictive vegetation mapping: geographic modelling of biospatial patterns in relation to environmental gradients , 1995 .

[112]  R. Crawford.  Plant community responses to Scotland's changing environment , 2001 .

[113]  Helen M. Regan,et al.  Mapping epistemic uncertainties and vague concepts in predictions of species distribution , 2002 .

[114]  W. Romme,et al.  Invasion of Subalpine Meadows by Lodgepole Pine in Yellowstone National Park, Wyoming, U.S.A. , 1993 .

[115]  Thomas W. Yee,et al.  Vector generalized additive models in plant ecology , 2002 .

[116]  H. Birks,et al.  Recent increases in species richness and shifts in altitudinal distributions of Norwegian mountain plants , 2003 .

[117]  Stefan Dullinger,et al.  A regional impact assessment of climate and land‐use change on alpine vegetation , 2003 .

[118]  T. T. Veblen,et al.  Stand dynamics of Pinus flexilis-dominated subalpine forests in the Colorado Front Range , 1991 .

[119]  C. Millar,et al.  FINE‐SCALE GENETIC STRUCTURE OF WHITEBARK PINE (PINUS ALBICAULIS): ASSOCIATIONS WITH WATERSHED AND GROWTH FORM , 1999, Evolution; international journal of organic evolution.

[120]  Jacques Baudry,et al.  LANDSCAPE DYNAMICS , 2022 .

[121]  C. T. Dyrness,et al.  Natural Vegetation of Oregon and Washington , 1988 .

[122]  J. Wallace,et al.  A Pacific Interdecadal Climate Oscillation with Impacts on Salmon Production , 1997 .

[123]  J. Hadley,et al.  Wind Erosion of Leaf Surface Wax in Alpine Timberline Conifers , 1989 .

[124]  Ramakrishna R. Nemani,et al.  MTCLIM: a mountain microclimate simulation model , 1989 .

[125]  D. Fox,et al.  Mean wind patterns and snow depths in an alpine-subalpine ecosystem as measured by damage to coniferous trees , 1996 .

[126]  D. Brockway.  Forest plant diversity at local and landscape scales in the Cascade Mountains of southwestern Washington , 1998 .

[127]  Directional positive feedback and pattern at an alpine tree line , 2004 .

[128]  M. Tjus.  Summer air temperatures and tree line dynamics at Abisko , 1996 .

[129]  S. K. Jenson,et al.  Extracting topographic structure from digital elevation data for geographic information-system analysis , 1988 .

[130]  W. Bowman,et al.  The Landscape Continuum: A Model for High-Elevation Ecosystems , 2004 .

[131]  A. Taylor.  Fire regimes and forest changes in mid and upper montane forests of the southern Cascades, Lassen Volcanic National Park, California, U.S.A. , 2000 .

[132]  C. Torrence,et al.  A Practical Guide to Wavelet Analysis. , 1998 .

[133]  D. Lettenmaier,et al.  A simple hydrologically based model of land surface water and energy fluxes for general circulation models , 1994 .

[134]  J. Grace,et al.  Temperature and stature: a study of temperatures in montane vegetation , 1987 .

[135]  Crowley,et al.  Atmospheric science: Methane rises from wetlands , 2011, Nature.

[136]  Peter E. Thornton,et al.  Generating surfaces of daily meteorological variables over large regions of complex terrain , 1997 .

[137]  P. W. Dunwiddie,et al.  A 6000‐Year Record of Forest History on Mount Rainier, Washington , 1986 .

[138]  J. Major,et al.  Ecology of the Vegetation and Soils Associated with Calcareous Parent Materials in Three Alpine Regions of Montana , 1968 .

[139]  Michael H. Kutner.  Applied Linear Statistical Models , 1974 .

[140]  Craig A. Miller,et al.  Historical and current forest and range landscapes in the interior Columbia River basin and portions of the Klamath and Great Basins. Part 1: Linking vegetation patterns and landscape vulnerability to potential insect and pathogen disturbances. , 1999 .

[141]  P. W. Dunwiddie,et al.  RECENT TREE INVASION OF SUBALPINE MEADOWS IN THE WIND RIVER MOUNTAINS, WYOMING , 1977 .

[142]  Albert J. Parker.  Comparative Gradient Structure and Forest Cover Types in Lassen Volcanic and Yosemite National Parks, California , 1995 .

[143]  F. Holtmeier,et al.  WHAT DOES THE TERM "KRUMMHOLZ" REALLY MEAN? OBSERVATIONS WITH SPECIAL REFERENCE TO THE ALPS AND THE COLORADO FRONT RANGE , 1981 .

[144]  G. Campbell,et al.  On the relationship between incoming solar radiation and daily maximum and minimum temperature , 1984 .

[145]  Timothy C. Coburn,et al.  Geostatistics for Natural Resources Evaluation , 2000, Technometrics.

[146]  Robert M. Krimmel.  Glaciers of the western United States , 2002 .

[147]  J. Betancourt,et al.  Patterns and sources of multidecadal oscillations in drought‐sensitive tree‐ring records from the central and southern Rocky Mountains , 2003 .

[148]  L. Kullman.  Rapid recent range‐margin rise of tree and shrub species in the Swedish Scandes , 2002 .

[149]  A. Lloyd,et al.  Spatial and Temporal Variability in the Growth and Climate Response of Treeline Trees in Alaska , 2002 .

[150]  Range Experiment Station,et al.  Forest habitat types of central Idaho , 1981 .

[151]  D. Butler.  Snow‐avalanche hazards, Southern Glacier National Park, Montana: The nature of local knowledge and individual responses , 1987 .

[152]  Sabine Mellmann-Brown.  The regeneration of whitebark pine in the timberline ecotone of the Beartooth Plateau, Montana and Wyoming , 2002 .

[153]  A. Hofgaard.  Inter-relationships between treeline position, species diversity, land use and climate change in the Central Scandes Mountains of Norway , 1997 .

[154]  P. Guttorp,et al.  Statistical Interpretation of Species Composition , 2001 .

[155]  Ferenc Csillag,et al.  Hierarchical decomposition of variance with applications in environmental mapping based on satellite images , 1996 .

[156]  R. Neilson,et al.  Canadian vegetation sensitivity to projected climatic change at three organizational levels , 1995 .

[157]  J. B. Benedict.  Downslope Soil Movement in a Colorado Alpine Region: Rates, Processes, and Climatic Significance , 1970 .

[158]  T. Kitzberger,et al.  Climatic and human influences on fire regimes in ponderosa pine forests in the Colorado Front Range. , 2000 .

[159]  G. Ettl,et al.  Growth response of ponderosa pine (Pinus ponderosa) to climate in the eastern Cascade Mountains, Washington, U.S.A.: Implications for climatic change1 , 2002 .

[160]  Robert E. Keane,et al.  Temporal patterns of ecosystem processes on simulated landscapes in Glacier National Park, Montana, USA , 1999, Landscape Ecology.

[161]  David M. Cairns,et al.  The suitability of montane ecotones as indicators of global climatic change , 1996 .

[162]  G. Grabherr,et al.  Climate effects on mountain plants , 1994, Nature.

[163]  Stephen J. McGregor,et al.  An integrated geographic information system approach for modeling the suitability of conifer habitat in an alpine environment , 1998 .

[164]  Bernard L. Kovalchik,et al.  Forest habitat types of Montana. , 1977 .

[165]  I. Apostolova,et al.  Influence of warming on timberline rising: a case study on Pinus peuce Griseb. in Bulgaria. , 2000 .

[166]  Charles B. Halpern,et al.  Overstory influences on herb and shrub communities in mature forests of western Washington, U.S.A. , 2000 .

[167]  G. Yohe,et al.  A globally coherent fingerprint of climate change impacts across natural systems , 2003, Nature.

[168]  T. Swetnam.  Fire History and Climate Change in Giant Sequoia Groves , 1993, Science.

[169]  R. Peet.  Latitudinal variation in southern Rocky Mountain forests , 1978 .

[170]  Gian-Reto Walther,et al.  Plants in a warmer world , 2003 .

[171]  T. Seastedt,et al.  Effects of the northern pocket gopher (Thomomys talpoides) on alpine soil characteristics, Niwot Ridge, CO , 2001 .

[172]  Hughes,et al.  Biological consequences of global warming: is the signal already apparent? , 2000, Trends in ecology & evolution.

[173]  S. Cumming.  FOREST TYPE AND WILDFIRE IN THE ALBERTA BOREAL MIXEDWOOD: WHAT DO FIRES BURN? , 2001 .

[174]  I. Campbell,et al.  Forest disequilibrium caused by rapid Little Ice Age cooling , 1993, Nature.

[175]  Stephen Wolfram,et al.  Mathematica: a system for doing mathematics by computer (2nd ed.) , 1991 .

[176]  J. Hanley,et al.  The meaning and use of the area under a receiver operating characteristic (ROC) curve. , 1982, Radiology.

[177]  Patrick N. Halpin,et al.  Forest gradient response in Sierran landscapes: the physical template , 2000, Landscape Ecology.

[178]  Stephen J. Walsh,et al.  Image enhancement of landsat thematic mapper data and GIS data integration for evaluation of resource characteristics , 1990 .

[179]  V. Barnett,et al.  Applied Linear Statistical Models , 1975 .

[180]  C. Daly.  Snow distribution patterns in the alpine krummholz zone , 1984 .

[181]  J. Sowell,et al.  Cuticular Transpiration of Whitebark Pine (Pinus albicaulis) within a Sierra Nevadan Timberline Ecotone, U.S.A. , 1982 .

[182]  A. Prasad,et al.  Potential Changes in Tree Species Richness and Forest Community Types following Climate Change , 2001, Ecosystems.

[183]  Lynn M. Resler,et al.  Topographic Shelter and Conifer Establishment and Mortality in an Alpine Environment, Glacier National Park, Montana , 2005 .

[184]  B. Cade,et al.  Estimating effects of limiting factors with regression quantiles , 1999 .

[185]  D. Yocky.  Multiresolution wavelet decomposition image merger of landsat thematic mapper and SPOT panchromatic data , 1996 .

[186]  A. Schoettle,et al.  Microscale patterns of tree establishment near upper treeline, Snowy Range, Wyoming, USA , 1999 .

[187]  Tara A. Forbis,et al.  Temporal patterns in seedling establishment on pocket gopher disturbances , 2003, Oecologia.

[188]  David L. Evans,et al.  THE EFFECT OF URBAN SPRAWL ON TIMBER , 1998 .

[189]  Stéphane Mallat,et al.  A Theory for Multiresolution Signal Decomposition: The Wavelet Representation , 1989, IEEE Trans. Pattern Anal. Mach. Intell..

[190]  Stephen J. Walsh,et al.  Remote sensing and GIS techniques for spatial and biophysical analyses of alpine treeline through process and empirical models , 1994 .

[191]  R. J. Kauth,et al.  The Tasseled Cap de-mystified , 1986 .

[192]  N. Lam,et al.  On the Issues of Scale, Resolution, and Fractal Analysis in the Mapping Sciences* , 1992 .

[193]  Diane Ebert-May,et al.  EFFECTS OF INTERANNUAL CLIMATE VARIATION ON ABOVEGROUND PHYTOMASS IN ALPINE VEGETATION , 1994 .

[194]  A. Guisan,et al.  Potential Impact of Climate Change on Vegetation in the European Alps: A Review , 2001 .

[195]  H. Linderholm.  Twentieth-century Scots Pine Growth Variations in the Central Scandinavian Mountains Related to Climate Change , 2002 .

[196]  H. Fritts,et al.  Tree Rings and Climate. , 1978 .

[197]  P. Curran.  The semivariogram in remote sensing: An introduction , 1988 .

[198]  D. Butler.  Zoogeomorphology: Animals as Geomorphic Agents , 1995 .

[199]  Bruno Torrésani,et al.  Practical Time-Frequency Analysis, Volume 9: Gabor and Wavelet Transforms, with an Implementation in S , 1998 .

[200]  N. Graham,et al.  Decadal Variability of Precipitation over Western North America , 1998 .

[201]  Shoshiro Minobe,et al.  A 50–70 year climatic oscillation over the North Pacific and North America , 1997 .

[202]  Friedrich-Karl Holtmeier.  Die Verbreitung der Holzarten im Oberengadin unter dem Einfluß des Menschen und des Lokalklimas , 1967 .

[203]  W. Gaines,et al.  USE OF LANDSAT MULTISPECTRAL SCANNER IMAGERY AND GEOGRAPHIC INFORMATION SYSTEMS TO MAP VEGETATION IN THE NORTH CASCADES GRIZZLY BEAR ECOSYSTEM , 1994 .

[204]  Ling Bian,et al.  Scale Dependencies of Vegetation and Topography in a Mountainous Environment of Montana , 1993 .

[205]  O. Davis.  Pollen Analysis of a Late-Glacial and Holocene Sediment Core from Mono Lake, Mono County, California , 1999, Quaternary Research.

[206]  L. Scuderi.  Late-Holocene upper timberline variation in the southern Sierra Nevada , 1987, Nature.

[207]  John R. Jensen,et al.  Introductory Digital Image Processing: A Remote Sensing Perspective , 1986 .

[208]  R. L. Little,et al.  Changes in sub-alpine tree distribution in western North America: a review of climatic and other causal factors , 1994 .

[209]  M. Wigmosta,et al.  A distributed hydrology-vegetation model for complex terrain , 1994 .

[210]  Ross Ihaka,et al.  Gentleman R: R: A language for data analysis and graphics , 1996 .

[211]  D. Norton,et al.  rHE GROWTH FORMS AND ECOLOGY OF NOTHOFAGUS SOLANDRI AT THE ALPINE TIMBERLINE, CRAIGIEBURN RANGE, NEW ZEALAND , 1984 .

[212]  J. Camarero,et al.  Pace and Pattern of Recent Treeline Dynamics: Response of Ecotones to Climatic Variability in the Spanish Pyrenees , 2004 .

[213]  D. D. French,et al.  Recent development of high-altitude Pinus sylvestris scrub in the Northern Cairngorm mountains, Scotland , 1997 .

[214]  Gretchen G. Moisen,et al.  Comparing five modelling techniques for predicting forest characteristics , 2002 .

[215]  R. Fonda,et al.  Forest Vegetation of the Montane and Subalpine Zones, Olympic Mountains, Washington , 1969 .

[216]  L. Brubaker.  Responses of tree populations to climatic change , 1986, Vegetatio.

[217]  M. Austin.  Spatial prediction of species distribution: an interface between ecological theory and statistical modelling , 2002 .

[218]  R. Keane,et al.  Fire-BGC: A mechanistic ecological process model for simulating fire succession on coniferous forest landscapes of the northern Rocky Mountains. Forest Service research paper , 2017 .

[219]  David L. Peterson,et al.  MOUNTAIN HEMLOCK GROWTH RESPONDS TO CLIMATIC VARIABILITY AT ANNUAL AND DECADAL TIME SCALES , 2001 .

[220]  T. P. Barnett,et al.  High-resolution palaeoclimatic records for the last millennium: interpretation, integration and comparison with General Circulation Model control-run temperatures , 1998 .

[221]  Robert H. Whittaker,et al.  Vegetation of the Great Smoky Mountains , 1956 .

[222]  Felix Kienast,et al.  Comparing models for tree distributions : concept, structures, and behavior , 2000 .

[223]  G. Meehl,et al.  Climate extremes: observations, modeling, and impacts. , 2000, Science.

[224]  Ronald P. Neilson,et al.  Potentially complex biosphere responses to transient global warming , 1998 .

[225]  J. Overpeck,et al.  Abrupt Climate Change , 2003, Science.

[226]  R. Whittaker,et al.  GRADIENT ANALYSIS OF VEGETATION* , 1967, Biological reviews of the Cambridge Philosophical Society.

[227]  M. Austin,et al.  Current problems of environmental gradients and species response curves in relation to continuum theory , 1994 .

[228]  Roy W. Koch,et al.  Surface Climate and Streamflow Variability in the Western United States and Their Relationship to Large‐Scale Circulation Indices , 1991 .

[229]  M. Noguer,et al.  Climate change 2001: The scientific basis. Contribution of Working Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change , 2002 .

[230]  L. C. Bliss,et al.  Ecology of Subalpine Meadows in the Olympic Mountains, Washington , 1970 .

[231]  N. Jones,et al.  Evidence for a Neoglacial advance of the Boundary Glacier, Banff National Park, Alberta , 1985 .

[232]  G. Malanson,et al.  FLORISTIC PATTERNS ON AVALANCHE PATHS IN THE NORTHERN ROCKY MOUNTAINS, USA , 1986 .

[233]  L. Kullman.  Tree-limit landscape evolution at the southern fringe of the Swedish Scandes (Dalarna province) – Holocene and 20th century perspectives , 2004 .

[234]  S. Ferguson.  A Climate-Change Scenario for the Columbia River Basin , 1997 .

[235]  S. Walsh,et al.  Compatibility of non‐synchronous in‐situ water quality data and remotely‐sensed spectral information for assessing lake turbidity levels in complex and inaccessible terrain , 1991 .

[236]  Prof. Dr. Walter Tranquillini.  Physiological Ecology of the Alpine Timberline , 1979, Ecological Studies.

[237]  P. D. Körner.  Alpine Plant Life , 1999, Springer Berlin Heidelberg.

[238]  P. Pellikka,et al.  Remote sensing of the impact of reindeer grazing on vegetation in northern Fennoscandia using SPOT XS data , 1994 .

[239]  R. Daubenmire.  ECOLOGIC PLANT GEOGRAPHY OF THE PACIFIC NORTHWEST , 1969 .

[240]  S. Ferrier,et al.  An evaluation of alternative algorithms for fitting species distribution models using logistic regression , 2000 .

[241]  Todd R. Lookingbill,et al.  An empirical approach towards improved spatial estimates of soil moisture for vegetation analysis , 2004, Landscape Ecology.

[242]  H. Shugart,et al.  Models of forest dynamics based on roles of tree species , 1996 .

[243]  Ling Bian,et al.  Cartographic Modeling of Snow Avalanche Path Location within Glacier National Park, Montana , 1990 .

[244]  B. Smith,et al.  Field guide for forested plant associations of the Wenatchee National Forest. Forest Service general technical report , 1995 .

[245]  J A Swets,et al.  Measuring the accuracy of diagnostic systems. , 1988, Science.

[246]  Stephen J. Walsh,et al.  Characterizing multitemporal alpine snowmelt patterns for ecological inferences , 1993 .

[247]  A. O. Nicholls,et al.  Determining species response functions to an environmental gradient by means of a β‐function , 1994 .

[248]  David W. Hosmer,et al.  Applied Logistic Regression , 1991 .

[249]  J. Wallace,et al.  ENSO-like Interdecadal Variability: 1900–93 , 1997 .

[250]  Dean L. Urban,et al.  Using model analysis to design monitoring programs for landscape management and impact assessment. , 2000 .

[251]  C. Millar,et al.  Sierra Nevada Forests: Where Did They Come From? Where Are They Going? What Does It Mean? , 1999 .

[252]  Mark New,et al.  Surface air temperature and its changes over the past 150 years , 1999 .

[253]  Herman H. Shugart,et al.  A Systems Analysis of the Global Boreal Forest , 1993 .

[254]  L. Kullman,et al.  Recent tree-limit history of Piceaabies in the southern Swedish Scandes , 1986 .

[255]  Simon Ferrier,et al.  Evaluating the predictive performance of habitat models developed using logistic regression , 2000 .

[256]  C. D. Bevins,et al.  Simulating coarse-scale vegetation dynamics using the Columbia River Basin succession model-crbsum. Forest Service general technical report , 1996 .

[257]  M. Hughes,et al.  Northern hemisphere temperatures during the past millennium: Inferences, uncertainties, and limitations , 1999 .

[258]  S. Walsh,et al.  SITE CHARACTERISTICS OF DEBRIS FLOWS AND THEIR RELATIONSHIP TO ALPINE TREELINE , 1994 .

[259]  S. Schneider,et al.  Fingerprints of global warming on wild animals and plants , 2003, Nature.

[260]  S. Payette,et al.  Recent Advance of the Arctic Treeline Along the Eastern Coast of Hudson Bay , 1995 .

[261]  W. Michener,et al.  Mound Microclimate, Nutrients and Seedling Survival , 2004 .

[262]  L. Daniels,et al.  SPATIOTEMPORAL INFLUENCES OF CLIMATE ON ALTITUDINAL TREELINE IN NORTHERN PATAGONIA , 2004 .

[263]  G. Malanson,et al.  Turf-banked terrace treads and risers, turf exfoliation and possible relationships with advancing treeline , 2004 .

[264]  J. Franklin.  Predicting the distribution of shrub species in southern California from climate and terrain‐derived variables , 1998 .

[265]  G. Malanson,et al.  Avalanche paths as fuel breaks: implications for fire management , 1984 .

[266]  T. Allen,et al.  Spatial and compositional pattern of alpine treeline, Glacier National Park, Montana , 1996 .

[267]  George A. McLachlan,et al.  Threatened and Endangered Species , 2002 .

[268]  J. Houghton,et al.  Climate change 2001 : the scientific basis , 2001 .

[269]  R. Neilson,et al.  Forest Processes and Global Environmental Change: Predicting the Effects of Individual and Multiple Stressors , 2001 .

[270]  G. McCabe,et al.  DECADAL VARIATIONS IN THE STRENGTH OF ENSO TELECONNECTIONS WITH PRECIPITATION IN THE WESTERN UNITED STATES , 1999 .

[271]  Robert G. Wagner,et al.  Process versus empirical models: which approach for forest ecosystem management? , 1996 .

[272]  M. Hughes,et al.  The Climate of the Last Millennium , 2003 .

[273]  D. Lettenmaier,et al.  A Long-Term Hydrologically Based Dataset of Land Surface Fluxes and States for the Conterminous United States* , 2002 .

[274]  D. Cooper.  White Spruce above and Beyond Treeline in the Arrigetch Peaks Region, Brooks Range, Alaska , 1986 .

[275]  P. Wardle,et al.  Evidence for rising upper limits of four native New Zealand forest trees , 1992 .

[276]  N. Stephenson,et al.  Actual evapotranspiration and deficit: biologically meaningful correlates of vegetation distribution across spatial scales , 1998 .

[277]  G. C. Stevens,et al.  THE CAUSES OF TREELINE , 1991 .

[278]  F. James Rohlf,et al.  Biometry: The Principles and Practice of Statistics in Biological Research , 1969 .

[279]  Brian D. Ripley,et al.  Neural networks and flexible regression and discrimination , 1994 .

[280]  L. Kullman.  Structural change in a subalpine birch woodland in North Sweden during the past century , 1991 .

[281]  S. Levin.  The problem of pattern and scale in ecology , 1992 .

[282]  T. Swetnam,et al.  Fire-Southern Oscillation Relations in the Southwestern United States , 1990, Science.

[283]  G. Henry,et al.  Tundra plants and climate change: the International Tundra Experiment (ITEX) , 1997 .

[284]  Malcolm K. Hughes,et al.  Global-scale temperature patterns and climate forcing over the past six centuries , 1998, Nature.

[285]  Eric M. Eliason,et al.  Adaptive box filters for removal of random noise from digital images , 1990 .

[286]  A. Lloyd.  Response of tree-line populations of foxtail pine (Pinus balfouriana) to climate variation over the last 1000 years , 1997 .

[287]  Stephen J. Walsh,et al.  Influence of snow patterns and snow avalanches on the alpine treeline ecotone , 1994 .

[288]  J. Stanford,et al.  GRIZZLY BEAR DIGGING: EFFECTS ON SUBALPINE MEADOW PLANTS IN RELATION TO MINERAL NITROGEN AVAILABILITY , 1998 .

[289]  U. Gruber,et al.  Avalanche Hazard Mapping with Satellite Data and a Digital Elevation Model , 1995 .

[290]  H. Delcourt,et al.  Late-Quaternary dynamics of temperate forests: Applications of paleoecology to issues of global environmental change , 1987 .

[291]  M. Beniston,et al.  Evidence of response of vegetation to environmental change on high-elevation sites in the Swiss Alps , 2000 .

[292]  Robert S. Thompson,et al.  A strategy for assessing potential future changes in climate, hydrology, and vegetation in the Western United States , 1998 .

[293]  Stephen J. Walsh,et al.  Mapping, modeling and visualization of the influences of geomorphic processes on the alpine treeline ecotone, Glacier National Park, MT, USA , 2003 .

[294]  T. Barnett,et al.  Interdecadal modulation of ENSO teleconnections , 1998 .

[295]  J. P. Mccarty.  Ecological Consequences of Recent Climate Change , 2001 .

[296]  B. Luckman,et al.  LANDSCAPE AND CLIMATE CHANGE IN THE CENTRAL CANADIAN ROCKIES DURING THE 20TH CENTURY , 1998 .

[297]  W. Baker.  Longterm response of disturbance landscapes to human intervention and global change , 1995, Landscape Ecology.

[298]  Christian Körner,et al.  A re-assessment of high elevation treeline positions and their explanation , 1998, Oecologia.

[299]  L. Scuderi.  A 2000-Year Tree Ring Record of Annual Temperatures in the Sierra Nevada Mountains , 1993, Science.

[300]  Dean L. Urban,et al.  Spatial estimation of air temperature differences for landscape-scale studies in montane environments , 2003 .

[301]  C. Daly,et al.  A Statistical-Topographic Model for Mapping Climatological Precipitation over Mountainous Terrain , 1994 .

[302]  Bruce P. Finney,et al.  Reduced growth of Alaskan white spruce in the twentieth century from temperature-induced drought stress , 2000, Nature.

[303]  P. McCullagh,et al.  Generalized Linear Models , 1992 .

[304]  D. Butler,et al.  AVALANCHE HAZARD IN GLACIER NATIONAL PARK: AN EL NIÑO CONNECTION? , 1999 .

[305]  O. J. Reichman,et al.  The role of pocket gophers as subterranean ecosystem engineers , 2002 .

[306]  H. T. Gisborne.  Results and trends at the Northern Rocky Mountain Forest and Range Experiment Station , 1942 .

[307]  Friedrich-Karl Holtmeier.  Geoökologische Beobachtungen und Studien an der subarktischen und alpinen Waldgrenze in vergleichender Sicht (nördliches Fennoskandien/Zentralalpen) , 1974 .

[308]  Robert Baxter,et al.  How will the tundra-taiga interface respond to climate change? , 2002, Ambio.

[309]  Edward R. Cook,et al.  Low-Frequency Signals in Long Tree-Ring Chronologies for Reconstructing Past Temperature Variability , 2002, Science.

[310]  P. Carrara.  Holocene and latest Pleistocene glacial chronology, Glacier National Park, Montana , 1987 .

[311]  J. Agee,et al.  SPATIAL CONTROLS OF HISTORICAL FIRE REGIMES: A MULTISCALE EXAMPLE FROM THE INTERIOR WEST, USA , 2001 .

[312]  M. Germino,et al.  Sky exposure, crown architecture, and low‐temperature photoinhibition in conifer seedlings at alpine treeline , 1999 .

[313]  W. D. Billings,et al.  An Alpine Snowbank Environment and Its Effects on Vegetation, Plant Development, and Productivity , 1959 .

[314]  Danielle J. Marceau,et al.  Remote sensing and the measurement of geographical entities in a forested environment. 1. The scale and spatial aggregation problem , 1994 .

[315]  R. Holmes.  Computer-Assisted Quality Control in Tree-Ring Dating and Measurement , 1983 .

[316]  A. Bunn.  Temporal and spatial patterns at alpine treeline in the Sierra Nevada USA : implications for global change , 2004 .

[317]  C. Braak,et al.  Unimodal models to relate species to environment , 1988 .

[318]  W. Baker,et al.  Spatial variation in tree regeneration in the forest-tundra ecotone, Rocky Mountain National Park, Colorado , 1995 .

[319]  Hugh G. Gauch,et al.  Ordination of Vegetation Samples by Gaussian Species Distributions , 1974 .

[320]  C. Wessman,et al.  Long-term studies of snow-vegetation interactions , 1993 .

[321]  L. Kullman.  Recent reversal of Neoglacial climate cooling trend in the Swedish Scandes as evidenced by mountain birch tree-limit rise , 2003 .

[322]  G. MacDonald,et al.  Recent White Spruce Dynamics at the Subarctic Alpine Treeline of North-Western Canada , 1995 .

[323]  D. Butler.  SNOW-AVALANCHE HAZARDS IN GLACIER NATIONAL PARK, MONTANA: METEOROLOGIC AND CLIMATOLOGIC ASPECTS , 1986 .

[324]  Diana F. Tomback,et al.  Dispersal of Whitebark Pine Seeds by Clark's Nutcracker: A Mutualism Hypothesis , 1982 .

[325]  N. Stephenson.  Climatic Control of Vegetation Distribution: The Role of the Water Balance , 1990, The American Naturalist.

[326]  L. Kullman.  Early Holocene Appearance of Mountain Birch (Betula pubescens ssp. tortuosa) at Unprecedented High Elevations in the Swedish Scandes: Megafossil Evidence Exposed by Recent Snow and Ice Recession , 2004 .

[327]  J. Oksanen,et al.  Continuum theory revisited: what shape are species responses along ecological gradients? , 2002 .