The impacts of increasing drought on forest dynamics, structure, and biodiversity in the United States

We synthesize insights from current understanding of drought impacts at stand-to-biogeographic scales, including management options, and we identify challenges to be addressed with new research. Large stand-level shifts underway in western forests already are showing the importance of interactions involving drought, insects, and fire. Diebacks, changes in composition and structure, and shifting range limits are widely observed. In the eastern US, the effects of increasing drought are becoming better understood at the level of individual trees, but this knowledge cannot yet be confidently translated to predictions of changing structure and diversity of forest stands. While eastern forests have not experienced the types of changes seen in western forests in recent decades, they too are vulnerable to drought and could experience significant changes with increased severity, frequency, or duration in drought. Throughout the continental United States, the combination of projected large climate-induced shifts in suitable habitat from modeling studies and limited potential for the rapid migration of tree populations suggests that changing tree and forest biogeography could substantially lag habitat shifts already underway. Forest management practices can partially ameliorate drought impacts through reductions in stand density, selection of drought-tolerant species and genotypes, artificial regeneration, and the development of multistructured stands. However, silvicultural treatments also could exacerbate drought impacts unless implemented with careful attention to site and stand characteristics. Gaps in our understanding should motivate new research on the effects of interactions involving climate and other species at the stand scale and how interactions and multiple responses are represented in models. This assessment indicates that, without a stronger empirical basis for drought impacts at the stand scale, more complex models may provide limited guidance.

[1]  R. Keane,et al.  Modeling climate changes and wildfire interactions: Effects on whitebark pine (Pinus albicaulis) and implications for restoration, Glacier National Park, Montana, USA , 2011 .

[2]  R. Norby,et al.  Elevated CO₂ increases tree-level intrinsic water use efficiency: insights from carbon and oxygen isotope analyses in tree rings across three forest FACE sites. , 2013, The New phytologist.

[3]  David J. Parsons,et al.  Impact of fire suppression on a mixed-conifer forest , 1979 .

[4]  N. McDowell,et al.  Homeostatic maintenance of ponderosa pine gas exchange in response to stand density changes. , 2006, Ecological applications : a publication of the Ecological Society of America.

[5]  James S. Clark,et al.  Competition‐interaction landscapes for the joint response of forests to climate change , 2014, Global change biology.

[6]  M. Hutchinson,et al.  Potential Impacts of Climate Change on the Distribution of North American Trees , 2007 .

[7]  J. Vose,et al.  Climate change adaptation and mitigation management options: a guide for natural resource managers in southern forest ecosystems. , 2013 .

[8]  D. Haase,et al.  Soil Moisture Stress Induces Transplant Shock in Stored and Unstored 2 + 0 Douglas-Fir Seedlings of Varying Root Volumes , 1993 .

[9]  D. Lawrence,et al.  Projected Future Changes in Vegetation in Western North America in the Twenty-First Century , 2013 .

[10]  M. M. Moore,et al.  REFERENCE CONDITIONS AND ECOLOGICAL RESTORATION: A SOUTHWESTERN PONDEROSA PINE PERSPECTIVE , 1999 .

[11]  S. LaDeau,et al.  Rising CO2 Levels and the Fecundity of Forest Trees , 2001, Science.

[12]  J. Clark Effects of long-term water balances on fire regime, north-western Minnesota , 1989 .

[13]  B. Clinton,et al.  Does Rhododendron maximum L. (Ericaceae) Reduce the Availability of Resources Above and Belowground for Canopy Tree Seedlings? , 2001 .

[14]  J. Hicke,et al.  Cross-scale Drivers of Natural Disturbances Prone to Anthropogenic Amplification: The Dynamics of Bark Beetle Eruptions , 2008 .

[15]  M. Goulden,et al.  Rapid shifts in plant distribution with recent climate change , 2008, Proceedings of the National Academy of Sciences.

[16]  A. Prasad,et al.  Estimating potential habitat for 134 eastern US tree species under six climate scenarios , 2008 .

[17]  Drew W. Purves,et al.  Climate‐related variation in mortality and recruitment determine regional forest‐type distributions , 2013 .

[18]  S. Fei,et al.  Change in oak abundance in the eastern United States from 1980 to 2008 , 2011 .

[19]  W. Koenig,et al.  Large-scale spatial synchrony and cross-synchrony in acorn production by two California oaks. , 2013, Ecology.

[20]  T. C. Hennessey,et al.  Effects of Planting Density and Seed Source on Loblolly Pine Stands in Southeastern Oklahoma , 2010 .

[21]  F. Schurr,et al.  Estimating demographic models for the range dynamics of plant species , 2010 .

[22]  Joel D. McMillin,et al.  Bark beetle-caused mortality in a drought-affected ponderosa pine landscape in Arizona, USA , 2009 .

[23]  David J. Mladenoff,et al.  Simulated effects of climate change, fragmentation, and inter-specific competition on tree species migration in northern Wisconsin, USA , 2008 .

[24]  Arjan J. H. Meddens,et al.  Carbon stocks of trees killed by bark beetles and wildfire in the western United States , 2013 .

[25]  A. Hessl,et al.  Climatic Controls on Historical Wildfires in West Virginia, 1939-2008 , 2010 .

[26]  Andrew M. Barton Intense wildfire in southeastern Arizona: transformation of a Madrean oak-pine forest to oak woodland , 2002 .

[27]  T. Hutchinson,et al.  Fire history and the establishment of oaks and maples in second-growth forests , 2008 .

[28]  James S. Clark,et al.  Plant migration and climate change , 1997 .

[29]  J. N. Long,et al.  Utah State University From the SelectedWorks of James Long 2012 Drought-driven disturbance history characterizes a southern Rocky Mountain subalpine forest , 2017 .

[30]  Joy Nystrom Mast,et al.  How resilient are southwestern ponderosa pine forests after crown fires , 2005 .

[31]  L. Anderegg,et al.  Consequences of widespread tree mortality triggered by drought and temperature stress , 2013 .

[32]  L. Iverson,et al.  Competition and climate affects US hardwood-forest tree mortality , 2013 .

[33]  C. Augspurger,et al.  Process-based modeling of species' distributions: what limits temperate tree species' range boundaries? , 2007, Ecology.

[34]  F. J. Barnes,et al.  Tree die-off in response to global change-type drought: mortality insights from a decade of plant water potential measurements. , 2009 .

[35]  M. Clifford,et al.  Multi-decadal drought and amplified moisture variability drove rapid forest community change in a humid region. , 2012, Ecology.

[36]  Philip N. Omi,et al.  Fuel treatments and fire severity: A meta-analysis , 2013 .

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

[38]  T. Fahey RECENT CHANGES IN AN UPLAND FOREST IN SOUTH-CENTRAL NEW YORK , 1998 .

[39]  J. Abatzoglou,et al.  Forest structure and species traits mediate projected recruitment declines in western US tree species , 2015 .

[40]  David R. Foster,et al.  FOREST RESPONSE TO CATASTROPHIC WIND: RESULTS FROM AN EXPERIMENTAL HURRICANE , 1999 .

[41]  Y. Malhi,et al.  Upslope migration of Andean trees , 2011 .

[42]  R. Minnich,et al.  Densification, stand-replacement wildfire, and extirpation of mixed conifer forest in Cuyamaca Rancho State Park, southern California , 2008 .

[43]  Peter A. Troch,et al.  Temperature sensitivity of drought-induced tree mortality portends increased regional die-off under global-change-type drought , 2009, Proceedings of the National Academy of Sciences.

[44]  D. Spittlehouse,et al.  GENETIC RESPONSES TO CLIMATE IN PINUS CONTORTA: NICHE BREADTH, CLIMATE CHANGE, AND REFORESTATION , 1999 .

[45]  Sarah Eichmann,et al.  Tree Rings And Climate , 2016 .

[46]  Jonathan M Adams,et al.  Masting behaviour in beech: linking reproduction and climatic variation , 2001 .

[47]  D. Bell,et al.  Early indicators of change: divergent climate envelopes between tree life stages imply range shifts in the western United States , 2014 .

[48]  T. Whitham,et al.  Deadly combination of genes and drought: increased mortality of herbivore‐resistant trees in a foundation species , 2009 .

[49]  J. Feddema,et al.  The role of climate in a pine forest regeneration pulse in the southwestern United States , 1996 .

[50]  J. Kane,et al.  Drought-induced mortality of a foundation species (Juniperus monosperma) promotes positive afterlife effects in understory vegetation , 2011, Plant Ecology.

[51]  W. Thuiller,et al.  Predicting species distribution: offering more than simple habitat models. , 2005, Ecology letters.

[52]  D. A. King,et al.  Climate change and fire effects on a prairie–woodland ecotone: projecting species range shifts with a dynamic global vegetation model , 2013, Ecology and evolution.

[53]  M. G. Ryan,et al.  Evaluating theories of drought-induced vegetation mortality using a multimodel-experiment framework. , 2013, The New phytologist.

[54]  J. Owens Constraints to seed production: temperate and tropical forest trees. , 1995, Tree physiology.

[55]  Urban Nilsson,et al.  Effects of regeneration methods on drought damage to newly planted Norway spruce seedlings , 1995 .

[56]  W. Bauerle,et al.  Drought impact on forest growth and mortality in the southeast USA: an analysis using Forest Health and Monitoring data. , 2009, Ecological applications : a publication of the Ecological Society of America.

[57]  R. Snell,et al.  Simulating long‐distance seed dispersal in a dynamic vegetation model , 2014 .

[58]  K. Elliott,et al.  Impacts of drought on tree mortality and growth in a mixed hardwood forest , 1994 .

[59]  M. G. Messina,et al.  Intensive Forest Management Affects Loblolly Pine (Pinus taeda L.) Growth and Survival on Poorly Drained Sites in Southern Arkansas , 2006 .

[60]  Andrea Vannini,et al.  Interactive effects of drought and pathogens in forest trees , 2006 .

[61]  Grant M. Domke,et al.  A Framework for Assessing Global Change Risks to Forest Carbon Stocks in the United States , 2013, PloS one.

[62]  P. Fulé,et al.  Woody debris and tree regeneration dynamics following severe wildfires in Arizona ponderosa pine forests , 2012 .

[63]  James S. Clark,et al.  The relationship between growth and mortality for seven co‐occurring tree species in the southern Appalachian Mountains , 2002 .

[64]  C. Allen,et al.  Drought-induced shift of a forest-woodland ecotone: rapid landscape response to climate variation. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[65]  M. DANIELW. Revisiting projected shifts in the climate envelopes of North American trees using updated general circulation models , 2011 .

[66]  E. Hogg,et al.  Factors affecting interannual variation in growth of western Canadian aspen forests during 1951-2000 , 2005 .

[67]  J. Linares,et al.  Interacting effects of changes in climate and forest cover on mortality and growth of the southernmost European fir forests , 2009 .

[68]  K. Waring,et al.  Old Pinus ponderosa growth responses to restoration treatments, climate and drought in a southwestern US landscape , 2014 .

[69]  Frank W Davis,et al.  Gene movement and genetic association with regional climate gradients in California valley oak (Quercus lobata Née) in the face of climate change , 2010, Molecular ecology.

[70]  Julio L. Betancourt,et al.  INFLUENCE OF LANDSCAPE STRUCTURE AND CLIMATE VARIABILITY ON A LATE HOLOCENE PLANT MIGRATION , 2003 .

[71]  G. Nowacki,et al.  The Demise of Fire and “Mesophication” of Forests in the Eastern United States , 2008 .

[72]  M. Abrams The Red Maple Paradox What explains the widespread expansion of red maple in eastern forests , 1998 .

[73]  J. Koricheva,et al.  Drought effects on damage by forest insects and pathogens: a meta‐analysis , 2012 .

[74]  J. Webster,et al.  Reforestation after the Fountain Fire in Northern California: An Untold Success Story , 2008 .

[75]  G. Parker,et al.  Evidence for a recent increase in forest growth , 2010, Proceedings of the National Academy of Sciences.

[76]  Antoine Guisan,et al.  Climatic extremes improve predictions of spatial patterns of tree species , 2009, Proceedings of the National Academy of Sciences.

[77]  S. Schiavon,et al.  Climate Change 2007: Impacts, Adaptation and Vulnerability. , 2007 .

[78]  E. Quarterman,et al.  Southern Mixed Hardwood Forest: Climax in the Southeastern Coastal Plain, U.S.A. , 1962 .

[79]  R. Shaw,et al.  Range shifts and adaptive responses to Quaternary climate change. , 2001, Science.

[80]  P. A. Mason,et al.  The influence of spatial patterns of damping‐off disease and arbuscular mycorrhizal colonization on tree seedling establishment in Ghanaian tropical forest soil , 2004 .

[81]  J. Clark,et al.  Interpreting recruitment limitation in forests. , 1999, American journal of botany.

[82]  Erin M. Schliep,et al.  Stochastic Modeling for Velocity of Climate Change , 2015, Journal of Agricultural, Biological, and Environmental Statistics.

[83]  N. Breda,et al.  Temperate forest trees and stands under severe drought: a review of ecophysiological responses, adaptation processes and long-term consequences , 2006 .

[84]  J. Franklin Moving beyond static species distribution models in support of conservation biogeography , 2010 .

[85]  Jason A. Clark,et al.  Modeling effects of climate change and fire management on western white pine (Pinus monticola) in the northern Rocky Mountains, USA , 2011 .

[86]  M. Abrams Adaptations and responses to drought in Quercus species of North America. , 1990, Tree physiology.

[87]  S. Roberts,et al.  Harvest residue and competing vegetation affect soil moisture, soil temperature, N availability, and Douglas-fir seedling growth , 2005 .

[88]  James S. Clark,et al.  Integration of ecological levels: individual plant growth, population mortality and ecosystem processes. , 1990 .

[89]  Marc D. Abrams,et al.  Fire and the Development of Oak Forests , 1992 .

[90]  F. Vuilleumier An atlas of past and Present Pollen Maps for Europe: 0-13000 Years Ago , 1987 .

[91]  Margaret M. Moore,et al.  Southwestern Ponderosa Forest Structure: Changes Since Euro-American Settlement , 1994, Journal of Forestry.

[92]  B. Day,et al.  The Influence of Precipitation upon the Width of Annual Rings of Certain Timber Trees , 1939 .

[93]  M. Stambaugh,et al.  Historic fire regime dynamics and forcing factors in the Boston Mountains, Arkansas, USA , 2006 .

[94]  David Frank,et al.  Climate signal age effects—Evidence from young and old trees in the Swiss Engadin , 2008 .

[95]  V. Rozas Dendrochronology of pedunculate oak (Quercus robur L.) in an old-growth pollarded woodland in northern Spain: tree-ring growth responses to climate , 2005 .

[96]  E. Assmann,et al.  The Principles of Forest Yield Study: Studies in the Organic Production, Structure, Increment and Yield of Forest Stands , 2013 .

[97]  M. Goulden,et al.  Rapid vegetation redistribution in Southern California during the early 2000s drought , 2012 .

[98]  C. Nowak,et al.  Changes in tree sapling composition within powerline corridors appear to be consistent with climatic changes in New York State , 2011 .

[99]  P. Grubb THE MAINTENANCE OF SPECIES‐RICHNESS IN PLANT COMMUNITIES: THE IMPORTANCE OF THE REGENERATION NICHE , 1977 .

[100]  Upmanu Lall,et al.  Is an Epic Pluvial Masking the Water Insecurity of the Greater New York City Region , 2013 .

[101]  Neil Pederson,et al.  A long-term perspective on a modern drought in the American Southeast , 2012 .

[102]  Edward R. Cook,et al.  IDENTIFYING FUNCTIONAL GROUPS OF TREES IN WEST GULF COAST FORESTS (USA): A TREE-RING APPROACH , 2001 .

[103]  S S I T C H,et al.  Evaluation of Ecosystem Dynamics, Plant Geography and Terrestrial Carbon Cycling in the Lpj Dynamic Global Vegetation Model , 2022 .

[104]  M. Walzer The long-term perspective. , 1986, Bulletin of the New York Academy of Medicine.

[105]  Nate G. McDowell,et al.  On underestimation of global vulnerability to tree mortality and forest die-off from hotter drought in the Anthropocene , 2015 .

[106]  D. Ackerly,et al.  Climate change impacts on California vegetation: physiology, life history, and ecosystem change. , 2012 .

[107]  Kevin McGarigal,et al.  Inhabitants of Landscape Scars: Succession of Woody Plants After Large, Severe Forest Fires in Arizona and New Mexico , 2008 .

[108]  H. Spiecker,et al.  Five decades of growth in a genetic field trial of Douglas-fir reveal trade-offs between productivity and drought tolerance , 2015, Tree Genetics & Genomes.

[109]  S. Grossnickle,et al.  Importance of root growth in overcoming planting stress , 2005, New Forests.

[110]  P. Reich,et al.  Geographic range predicts photosynthetic and growth response to warming in co-occurring tree species , 2015 .

[111]  J. Varner,et al.  Canopy disturbance and tree recruitment over two centuries in a managed longleaf pine landscape , 2008 .

[112]  R. Vilgalys,et al.  Evaluating the impacts of multiple generalist fungal pathogens on temperate tree seedling survival. , 2012, Ecology.

[113]  Marianne E. Porter,et al.  Differential tree mortality in response to severe drought: evidence for long‐term vegetation shifts , 2005 .

[114]  A. F. Hough,et al.  The Ecology and Silvics of Forests in the High Plateau of Pennsylvania , 1943 .

[115]  Joy Nystrom Mast,et al.  Spatial patch patterns and altered forest structure in middle elevation versus upper ecotonal mixed-conifer forests, Grand Canyon National Park, Arizona, USA , 2006 .

[116]  N. Barger,et al.  Tree regeneration following drought- and insect-induced mortality in piñon-juniper woodlands. , 2013, The New phytologist.

[117]  M. Sykes,et al.  Climate change threats to plant diversity in Europe. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[118]  D. Peterson,et al.  Climate and wildfire area burned in western U.S. ecoprovinces, 1916-2003. , 2009, Ecological applications : a publication of the Ecological Society of America.

[119]  M. Abrams,et al.  Leaf structural characteristics of 31 hardwood and conifer tree species in central Wisconsin: influence of light regime and shade-tolerance rank. , 1990 .

[120]  James S. Clark,et al.  MOLECULAR INDICATORS OF TREE MIGRATION CAPACITY UNDER RAPID CLIMATE CHANGE , 2005 .

[121]  N. Zimmermann,et al.  Field Evidence of Colonisation by Holm Oak, at the Northern Margin of Its Distribution Range, during the Anthropocene Period , 2013, PloS one.

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

[123]  David Saah,et al.  Aboveground live carbon stock changes of California wildland ecosystems, 2001–2010 , 2015 .

[124]  S. Running,et al.  Forests [Chapter 7] , 2014 .

[125]  D. Whitehead,et al.  Changes in composition, structure and aboveground biomass over seventy-six years (1930-2006) in the Black Rock Forest, Hudson Highlands, southeastern New York State. , 2008, Tree physiology.

[126]  J. Feddema,et al.  Modeling high-severity fire, drought and climate change impacts on ponderosa pine regeneration , 2013 .

[127]  J. Franklin,et al.  Modeling plant species distributions under future climates: how fine scale do climate projections need to be? , 2013, Global change biology.

[128]  S. Running,et al.  Ch. 7: Forests. Climate Change Impacts in the United States: The Third National Climate Assessment , 2014 .

[129]  H. Erkert Der Einfluß des Mondlichtes auf die Aktivitätsperiodik nachtaktiver Säugetiere , 1974, Oecologia.

[130]  T. Kolb,et al.  Establishment and growth of piñon pine regeneration vary by nurse type along a soil substrate age gradient in northern Arizona , 2015 .

[131]  J. Clark,et al.  LOCAL AND REGIONAL SEDIMENT CHARCOAL EVIDENCE FOR FIRE REGIMES IN PRESETTLEMENT NORTH-EASTERN NORTH AMERICA , 1996 .

[132]  T. Swetnam,et al.  Mesoscale Disturbance and Ecological Response to Decadal Climatic Variability in the American Southwest , 1998 .

[133]  James S. Clark,et al.  Predicting biodiversity change: outside the climate envelope, beyond the species-area curve. , 2006, Ecology.

[134]  H. Grissino-Mayer,et al.  Climate-tree growth relationships of longleaf pine (Pinus palustris Mill.) in the Southeastern Coastal Plain, USA. , 2009 .

[135]  M. Lesser,et al.  Making a stand: five centuries of population growth in colonizing populations of Pinus ponderosa. , 2012, Ecology.

[136]  A. Prasad,et al.  Thinning, fire, and oak regeneration across a heterogeneous landscape in the eastern U.S.: 7-year results , 2008 .

[137]  James S. Clark,et al.  Climate change vulnerability of forest biodiversity: climate and competition tracking of demographic rates , 2011 .

[138]  P. Marquet,et al.  A Significant Upward Shift in Plant Species Optimum Elevation During the 20th Century , 2008, Science.

[139]  M. Loik,et al.  A multi-scale perspective of water pulses in dryland ecosystems: climatology and ecohydrology of the western USA , 2004, Oecologia.

[140]  J. Clair,et al.  Comparative genetic responses to climate in the varieties of Pinus ponderosa and Pseudotsuga menziesii: Clines in growth potential , 2014 .

[141]  L. Graumlich Response of tree growth to climatic variation in the mixed conifer and deciduous forests of the upper Great Lakes region , 1993 .

[142]  Dirk R. Schmatz,et al.  Climate, competition and connectivity affect future migration and ranges of European trees , 2012 .

[143]  A. Cooper,et al.  Long-term drought sensitivity of trees in second-growth forests in a humid region , 2012 .

[144]  Ottar Michelsen,et al.  Continent-wide response of mountain vegetation to climate change , 2012 .

[145]  Christopher B. Field,et al.  Tree mortality predicted from drought-induced vascular damage , 2015 .

[146]  N. McDowell,et al.  The interdependence of mechanisms underlying climate-driven vegetation mortality. , 2011, Trends in ecology & evolution.

[147]  Antoine Guisan,et al.  Tree line shifts in the Swiss Alps: Climate change or land abandonment? , 2007 .

[148]  D. Twidwell,et al.  Drought‐induced woody plant mortality in an encroached semi‐arid savanna depends on topoedaphic factors and land management , 2014 .

[149]  A. Cescatti,et al.  Silvicultural alternatives, competition regime and sensitivity to climate in a European beech forest , 1998 .

[150]  Kai Zhu,et al.  Dual impacts of climate change: forest migration and turnover through life history , 2014, Global change biology.

[151]  Kristen A. Pelz,et al.  How will aspen respond to mountain pine beetle? A review of literature and discussion of knowledge gaps , 2013 .

[152]  D. Foster,et al.  Fire on the New England landscape: regional and temporal variation, cultural and environmental controls , 2002 .

[153]  Dominique Bachelet,et al.  DYNAMIC SIMULATION OF TREE–GRASS INTERACTIONS FOR GLOBAL CHANGE STUDIES , 2000 .

[154]  Peter Z. Fulé,et al.  Changes in forest structure of a mixed conifer forest, southwestern Colorado, USA. , 2009 .

[155]  B. McGlynn,et al.  Ecohydrology of an outbreak: mountain pine beetle impacts trees in drier landscape positions first , 2013 .

[156]  R. Guyette,et al.  Individual Tree and Stand Level Influences on the Growth, Vigor, and Decline of Red Oaks in the Ozarks , 2008 .

[157]  R. Seager,et al.  Megadroughts in North America: placing IPCC projections of hydroclimatic change in a long‐term palaeoclimate context , 2010 .

[158]  K. Price,et al.  Regional vegetation die-off in response to global-change-type drought. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[159]  R. Ohlemüller,et al.  Rapid Range Shifts of Species Associated with High Levels of Climate Warming , 2011, Science.

[160]  Michael F. Wehner,et al.  Projections of Future Drought in the Continental United States and Mexico , 2011 .

[161]  Christopher D. O’Connor Spatial and temporal dynamics of disturbance interactions along an ecological gradient , 2013 .

[162]  R. Neilson,et al.  Response of vegetation distribution, ecosystem productivity, and fire to climate change scenarios for California , 2008 .

[163]  A. Hamann,et al.  Recent declines of Populus tremuloides in North America linked to climate , 2013 .

[164]  H. Safford,et al.  Widespread shifts in the demographic structure of subalpine forests in the Sierra Nevada, California, 1934 to 2007 , 2013 .

[165]  J. Overpeck,et al.  Dramatic response to climate change in the Southwest: Robert Whittaker's 1963 Arizona Mountain plant transect revisited , 2013, Ecology and evolution.

[166]  N. Coops,et al.  Modeling the occurrence of 15 coniferous tree species throughout the Pacific Northwest of North America using a hybrid approach of a generic process‐based growth model and decision tree analysis , 2011 .

[167]  C. Allen Interactions Across Spatial Scales among Forest Dieback, Fire, and Erosion in Northern New Mexico Landscapes , 2007, Ecosystems.

[168]  Olivier Bouriaud,et al.  Tree diversity does not always improve resistance of forest ecosystems to drought , 2014, Proceedings of the National Academy of Sciences.

[169]  J. Miller,et al.  Effect of vegetative competition on the moisture and nutrient status of loblolly pine , 1984 .

[170]  Ben A. Minteer,et al.  Managed Relocation: Integrating the Scientific, Regulatory, and Ethical Challenges , 2012 .

[171]  J. Kutzbach,et al.  Widespread drought episodes in the western Great Lakes region during the past 2000 years: Geographic extent and potential mechanisms , 2006 .

[172]  Steven P. Brumby,et al.  Quantifying tree mortality in a mixed species woodland using multitemporal high spatial resolution satellite imagery , 2013 .

[173]  K. O’Hara,et al.  The Stand: Revisiting a Central Concept in Forestry , 2013 .

[174]  B. Pedersen,et al.  THE ROLE OF STRESS IN THE MORTALITY OF MIDWESTERN OAKS AS INDICATED BY GROWTH PRIOR TO DEATH , 1998 .

[175]  S. Goeking Disentangling Forest Change from Forest Inventory Change: A Case Study from the US Interior West , 2015 .

[176]  Han Y. H. Chen,et al.  Observations from old forests underestimate climate change effects on tree mortality , 2013, Nature Communications.

[177]  Hans Peter Schmid,et al.  Chronic water stress reduces tree growth and the carbon sink of deciduous hardwood forests , 2014, Global change biology.

[178]  D. Dey,et al.  A Meta-Analysis of the Fire-Oak Hypothesis: Does Prescribed Burning Promote Oak Reproduction in Eastern North America? , 2013 .

[179]  D. Breshears,et al.  Vegetation Responses to Extreme Hydrological Events: Sequence Matters , 2008, The American Naturalist.

[180]  H. Kowarzyk Structure and Function. , 1910, Nature.

[181]  R. Robberecht,et al.  High temperature and drought stress effects on survival of Pinus ponderosa seedlings. , 1996, Tree physiology.

[182]  D. Ackerly,et al.  Beyond a warming fingerprint: individualistic biogeographic responses to heterogeneous climate change in California , 2014, Global change biology.

[183]  Andrew O. Finley,et al.  An indicator of tree migration in forests of the eastern United States , 2009 .

[184]  John L. Innes,et al.  The occurrence of flowering and fruiting on individual trees over 3 years and their effects on subsequent crown condition , 1994, Trees.

[185]  Todd E. Dawson,et al.  Determining water use by trees and forests from isotopic, energy balance and transpiration analyses: the roles of tree size and hydraulic lift. , 1996, Tree physiology.

[186]  J. Betancourt,et al.  Role of multidecadal climate variability in a range extension of pinyon pine. , 2006, Ecology.

[187]  Jerry F. Franklin,et al.  Twentieth-century decline of large-diameter trees in Yosemite National Park, California, USA , 2009 .

[188]  J. Feddema,et al.  Double whammy: high-severity fire and drought in ponderosa pine forests of the Southwest , 2013 .

[189]  T. Parchman,et al.  Development of genetic diversity, differentiation and structure over 500 years in four ponderosa pine populations , 2013, Molecular ecology.

[190]  B. Beckage,et al.  A rapid upward shift of a forest ecotone during 40 years of warming in the Green Mountains of Vermont , 2008, Proceedings of the National Academy of Sciences.

[191]  C. Messier,et al.  Amounts of logging residues affect planting microsites: A manipulative study across northern forest ecosystems , 2014 .

[192]  Julio L. Betancourt,et al.  Ecology and the ratchet of events: Climate variability, niche dimensions, and species distributions , 2009, Proceedings of the National Academy of Sciences.

[193]  E. Engber,et al.  Patterns of flammability of the California oaks: the role of leaf traits , 2012 .

[194]  B. Sturtevant,et al.  Modeling Forest Mortality Caused by Drought Stress: Implications for Climate Change , 2012, Ecosystems.

[195]  J. Martínez‐Vilalta,et al.  The effect of fungal pathogens on the water and carbon economy of trees: implications for drought-induced mortality. , 2014, The New phytologist.

[196]  J. Guldin Adapting silviculture to a changing climate in the southern United States , 2014 .

[197]  N. Cobb,et al.  Relationship of stand characteristics to drought-induced mortality in three southwestern piñion-juniper woodlands. , 2009, Ecological applications : a publication of the Ecological Society of America.

[198]  Peter B. Reich,et al.  Frontiers inEcology and the Environment Will environmental changes reinforce the impact of global warming on the prairie – forest border of central North America ? , 2009 .

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

[200]  F. A. Bazzaz,et al.  Changes in drought response strategies with ontogeny in Quercus rubra: implications for scaling from seedlings to mature trees , 2000, Oecologia.

[201]  N. McDowell,et al.  Perpetuating old ponderosa pine , 2007 .

[202]  D. Board,et al.  The role of germination microsite in the establishment of sugar pine and Jeffrey pine seedlings , 2010 .

[203]  G. Aussenac Interactions between forest stands and microclimate: Ecophysiological aspects and consequences for silviculture , 2000 .

[204]  R. B. Jackson,et al.  Quantifying surface albedo and other direct biogeophysical climate forcings of forestry activities , 2015, Global change biology.

[205]  C. Fastie,et al.  Causes and Ecosystem Consequences of Multiple Pathways of Primary Succession at Glacier Bay, Alaska , 1995 .

[206]  David Medvigy,et al.  Predicting ecosystem dynamics at regional scales: an evaluation of a terrestrial biosphere model for the forests of northeastern North America , 2012, Philosophical Transactions of the Royal Society B: Biological Sciences.

[207]  M. Sarasola,et al.  Stand density and drought interaction on water relations of Nothofagus antarctica: contribution of forest management to climate change adaptability , 2011, Trees.

[208]  A. Taylor,et al.  Widespread Increase of Tree Mortality Rates in the Western United States , 2009, Science.

[209]  R. Seager,et al.  Temperature as a potent driver of regional forest drought stress and tree mortality , 2013 .

[210]  Heike Lischke,et al.  Interannual climate variability and population density thresholds can have a substantial impact on simulated tree species’ migration , 2013 .

[211]  P. Hanson,et al.  Drought disturbance from climate change: response of United States forests. , 2000, The Science of the total environment.

[212]  M. Payton,et al.  Productivity, crown architecture, and gas exchange of North Carolina and Oklahoma/Arkansas loblolly pine families growing on a droughty site in southeastern Oklahoma ☆ , 2004 .

[213]  Paul D. Henne,et al.  The past ecology of Abies alba provides new perspectives on future responses of silver fir forests to global warming , 2013 .

[214]  Mark W. Schwartz,et al.  How fast and far might tree species migrate in the eastern United States due to climate change , 2004 .

[215]  T. Kolb,et al.  Post-fire ponderosa pine regeneration with and without planting in Arizona and New Mexico , 2015 .

[216]  J. Marshall,et al.  Photosynthetic response, carbon isotopic composition, survival, and growth of three stock types under water stress enhanced by vegetative competition , 2012 .

[217]  J. Hicke,et al.  Climate, weather, and recent mountain pine beetle outbreaks in the western United States , 2014 .

[218]  S. Fraver,et al.  Effects of thinning on drought vulnerability and climate response in north temperate forest ecosystems. , 2013, Ecological applications : a publication of the Ecological Society of America.

[219]  S. Yeaman,et al.  Adaptation, migration or extirpation: climate change outcomes for tree populations , 2008, Evolutionary applications.

[220]  N. Barger,et al.  Declines in pinyon pine cone production associated with regional warming , 2012 .

[221]  P. Brown,et al.  CLIMATE AND DISTURBANCE FORCING OF EPISODIC TREE RECRUITMENT IN A SOUTHWESTERN PONDEROSA PINE LANDSCAPE , 2005 .

[222]  I. C. Prentice,et al.  Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model , 2003 .

[223]  A. Dai Increasing drought under global warming in observations and models , 2013 .

[224]  T. Whitham,et al.  Mortality Gradients within and among Dominant Plant Populations as Barometers of Ecosystem Change During Extreme Drought , 2006, Conservation biology : the journal of the Society for Conservation Biology.

[225]  T. Webb, Eastern North America , 2003, Riesling Rediscovered.

[226]  S. Rambal,et al.  Mast seeding under increasing drought: results from a long-term data set and from a rainfall exclusion experiment. , 2010, Ecology.

[227]  K. Waring,et al.  Enhancing resiliency and restoring ecological attributes in second-growth ponderosa pine stands in Northern New Mexico, USA , 2015 .

[228]  J. Ganey,et al.  Tree mortality in drought-stressed mixed-conifer and ponderosa pine forests, Arizona, USA , 2011 .

[229]  W. Bauerle,et al.  Effect of natural atmospheric CO2 fertilization suggested by open‐grown white spruce in a dry environment , 2006 .

[230]  Peter Z. Fulé,et al.  Restoring Ecosystem Health in Ponderosa Pine Forests of the Southwest , 1997, Journal of Forestry.

[231]  Benjamin Smith,et al.  Simulating past and future dynamics of natural ecosystems in the United States , 2003 .

[232]  Kai Zhu,et al.  Dynamic Inverse Prediction and Sensitivity Analysis With High-Dimensional Responses: Application to Climate-Change Vulnerability of Biodiversity , 2013 .

[233]  Jessica Gurevitch,et al.  Responses of insect pests, pathogens, and invasive plant species to climate change in the forests of northeastern North America: what can we predict? , 2009 .

[234]  D. Coomes,et al.  Influences of Forest Structure, Climate and Species Composition on Tree Mortality across the Eastern US , 2010, PloS one.

[235]  A. S. Meador,et al.  Effectiveness of fuel reduction treatments: Assessing metrics of forest resiliency and wildfire severity after the Wallow Fire, AZ , 2014 .

[236]  Henry J. Oosting,et al.  An Ecological Analysis of the Plant Communities of Piedmont, North Carolina , 1942 .

[237]  CH' , 2018, Dictionary of Upriver Halkomelem.

[238]  Xiaodong Zeng,et al.  Investigation of uncertainties of establishment schemes in dynamic global vegetation models , 2014, Advances in Atmospheric Sciences.

[239]  Antoine Guisan,et al.  Tree line shifts in the Swiss Alps: Climate change or land abandonment? , 2007 .

[240]  C. Allen,et al.  Watering the forest for the trees: an emerging priority for managing water in forest landscapes , 2011 .

[241]  Deborah Charlesworth,et al.  Introduction to plant population ecology , 1983, Vegetatio.

[242]  T. Knürr,et al.  Gene Flow and Local Adaptation in Trees , 2007 .

[243]  Andrei Rycoff Genetic Resource Management and Climate Change: Genetic Options for Adapting National Forests to Climate Change , 2012 .

[244]  James S. Clark,et al.  FECUNDITY OF TREES AND THE COLONIZATION–COMPETITION HYPOTHESIS , 2004 .

[245]  Kai Zhu,et al.  More than the sum of the parts: forest climate response from joint species distribution models. , 2014, Ecological applications : a publication of the Ecological Society of America.

[246]  D. Peterson,et al.  Forest responses to climate change in the northwestern United States: Ecophysiological foundations for adaptive management , 2011 .

[247]  V. Monleon,et al.  Evidence of Tree Species’ Range Shifts in a Complex Landscape , 2015, PloS one.

[248]  T. Brown,et al.  Spatial Distribution of Water Supply in the Coterminous United States 1 , 2008 .

[249]  J. Hicke,et al.  Spatiotemporal patterns of observed bark beetle-caused tree mortality in British Columbia and the western United States. , 2012, Ecological applications : a publication of the Ecological Society of America.

[250]  G. Houle Mast seeding in Abies balsamea, Acer saccharum and Betula alleghaniensis in an old growth, cold temperate forest of north‐eastern North America , 1999 .

[251]  James S. Clark,et al.  Impacts of Increasing Drought on Forest Dynamics, Structure, Diversity, and Management , 2016 .

[252]  Janneke HilleRisLambers,et al.  ESTIMATING POPULATION SPREAD: WHAT CAN WE FORECAST AND HOW WELL? , 2003 .

[253]  C. Field,et al.  Drought's legacy: multiyear hydraulic deterioration underlies widespread aspen forest die‐off and portends increased future risk , 2013, Global change biology.

[254]  M. Abrams Where Has All the White Oak Gone? , 2003 .

[255]  N. Zimmermann,et al.  TreeMig: A forest-landscape model for simulating spatio-temporal patterns from stand to landscape scale , 2006 .

[256]  L. Iverson,et al.  Long-term droughtiness and drought tolerance of eastern US forests over five decades , 2015 .

[257]  R. Q. Thomas,et al.  Frequency, not relative abundance, of temperate tree species varies along climate gradients in eastern North America. , 2010, Ecology.

[258]  David A. Orwig,et al.  The Legacy of Episodic Climatic Events in Shaping Temperate, Broadleaf Forests , 2014 .

[259]  S. Saatchi,et al.  Compositional shifts in Costa Rican forests due to climate‐driven species migrations , 2013, Global change biology.

[260]  Michael Dorman,et al.  Forest performance during two consecutive drought periods: Diverging long-term trends and short-term responses along a climatic gradient , 2013 .

[261]  Effect of Light Conditions on the Resistance of Current-year Fagus Crenata Seedlings Against Fungal Pathogens Causing Damping-off in a Natural Beech Forest: Fungus Isolation and Histological and Chemical Resistance , 2009, Journal of Chemical Ecology.

[262]  M. Lesser,et al.  Contributions of long-distance dispersal to population growth in colonising Pinus ponderosa populations. , 2013, Ecology letters.

[263]  B. Muys,et al.  Growth responses of West-Mediterranean Pinus nigra to climate change are modulated by competition and productivity: Past trends and future perspectives , 2011 .

[264]  T. Swetnam,et al.  Warming and Earlier Spring Increase Western U.S. Forest Wildfire Activity , 2006, Science.

[265]  L. C. Wensel,et al.  The relationship between tree diameter growth and climate for coniferous species in northern California , 2000 .

[266]  Tim Parshall,et al.  Canopy mortality and stand-scale change in a northern hemlock–hardwood forest , 1995 .

[267]  B. Shuman,et al.  Woodland-to-forest transition during prolonged drought in Minnesota after ca. AD 1300. , 2009, Ecology.

[268]  A. Burdett Physiological processes in plantation establishment and the development of specifications for forest planting stock , 1990 .

[269]  A. Prasad,et al.  Modifying climate change habitat models using tree species-specific assessments of model uncertainty and life history-factors , 2011 .

[270]  S. Schwinning,et al.  Hydraulic responses to extreme drought conditions in three co-dominant tree species in shallow soil over bedrock , 2013, Oecologia.

[271]  S. Pallardy,et al.  Successional trends and apparent Acer saccharum regeneration failure in an oak-hickory forest in central Missouri, USA , 2009, Plant Ecology.

[272]  Jesse K. Kreye,et al.  Toward a mechanism for eastern North American forest mesophication: differential litter drying across 17 species. , 2013, Ecological applications : a publication of the Ecological Society of America.

[273]  H. Bugmann,et al.  Growth-Mortality Relationships in Piñon Pine (Pinus edulis) during Severe Droughts of the Past Century: Shifting Processes in Space and Time , 2014, PloS one.

[274]  J. Foster,et al.  Montane forest ecotones moved downslope in northeastern USA in spite of warming between 1984 and 2011 , 2015, Global change biology.

[275]  M. K. Cleaveland,et al.  Reconstruction and Analysis of Spring Rainfall over the Southeastern U.S. for the Past 1000 Years , 1992 .

[276]  D. Kulakowski,et al.  Fire regimes of quaking aspen in the Mountain West , 2013 .

[277]  D. Stahle,et al.  North Carolina Climate Changes Reconstructed from Tree Rings: A.D. 372 to 1985 , 1988, Science.

[278]  A. Hamann,et al.  Geographic variation in growth response of Douglas‐fir to interannual climate variability and projected climate change , 2010 .

[279]  Christopher J. Still,et al.  Forest responses to increasing aridity and warmth in the southwestern United States , 2010, Proceedings of the National Academy of Sciences.

[280]  C. Lafon,et al.  Relationships of Fire and Precipitation Regimes in Temperate Forests of the Eastern United States , 2012 .

[281]  P. Moorcroft,et al.  Tree mortality in the eastern and central United States: patterns and drivers , 2011 .

[282]  D. Faber-Langendoen,et al.  Oak mortality in sand savannas following drought in east-central Minnesota. , 1993 .

[283]  B. Jędrzejewska,et al.  Rodent population dynamics in a primeval deciduous forest (Białowieża National Park) in relation to weather, seed crop, and predation , 1993 .

[284]  G. Rehfeldt,et al.  Climatic niche, ecological genetics, and impact of climate change on eastern white pine (Pinus strobus L.): Guidelines for land managers , 2013 .

[285]  J. Peñuelas,et al.  The altitude-for-latitude disparity in the range retractions of woody species. , 2009, Trends in ecology & evolution.

[286]  I. Ibáñez,et al.  EXPLOITING TEMPORAL VARIABILITY TO UNDERSTAND TREE RECRUITMENT RESPONSE TO CLIMATE CHANGE , 2007 .

[287]  M. G. Ryan,et al.  Continued warming could transform Greater Yellowstone fire regimes by mid-21st century , 2011, Proceedings of the National Academy of Sciences.

[288]  J. Régnière,et al.  Climate Change and Bark Beetles of the Western United States and Canada: Direct and Indirect Effects , 2010 .

[289]  D. Tinker,et al.  Forest structure and regeneration following a mountain pine beetle epidemic in southeastern Wyoming , 2012 .

[290]  N. Coops,et al.  A process-based approach to estimate lodgepole pine (Pinus contorta Dougl.) distribution in the Pacific Northwest under climate change , 2011 .

[291]  M. Palmer,et al.  Stand dynamics of an Appalachian old-growth forest during a severe drought episode , 2003 .

[292]  C. Field,et al.  The velocity of climate change , 2009, Nature.

[293]  T. C. Hennessey,et al.  Water use in thinned loblolly pine plantations , 1992 .

[294]  E. Cook,et al.  Drought Reconstructions for the Continental United States , 1999 .

[295]  A. Flint,et al.  Forest mortality in high-elevation whitebark pine (Pinus albicaulis) forests of eastern California, USA; influence of environmental context, bark beetles, climatic water deficit, and warming , 2012 .

[296]  S. Arndt,et al.  Modelling the potential impact of climate variability and change on species regeneration potential in the temperate forests of South‐Eastern Australia , 2012 .

[297]  James S. Clark,et al.  Failure to migrate: lack of tree range expansion in response to climate change , 2012 .

[298]  Ralph D. Nyland,et al.  Silviculture: Concepts and Applications , 1996 .

[299]  Laura Boisvert-Marsh,et al.  Shifting with climate? Evidence for recent changes in tree species distribution at high latitudes , 2014 .

[300]  Judith D. Gardiner,et al.  Exploring tree species colonization potentials using a spatially explicit simulation model: implications for four oaks under climate change , 2013, Global change biology.

[301]  James M. Dyer,et al.  Multiple interacting ecosystem drivers: toward an encompassing hypothesis of oak forest dynamics across eastern North America , 2011 .

[302]  Aaron S. Weed,et al.  Consequences of climate change for biotic disturbances in North American forests , 2013 .

[303]  N. McDowell,et al.  A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests , 2010 .

[304]  K. Shine,et al.  Intergovernmental panel on Climate change (IPCC),in encyclopedia of Enviroment and society,Vol.3 , 2007 .