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Understanding strategies for seed dispersal by wind under contrasting atmospheric conditions
Traits associated with seed dispersal vary tremendously among sympatric wind-dispersed plants. We used two contrasting tropical tree species, seed traps, micrometeorology, and a mechanistic model to evaluate how variation in four key traits affects seed dispersal by wind. The conceptual framework of movement ecology, wherein external factors (wind) interact with internal factors (plant traits) that enable movement and determine when and where movement occurs, fully captures the variable inputs and outputs of wind dispersal models and informs their interpretation. We used model calculations to evaluate the spatial pattern of dispersed seeds for the 16 factorial combinations of four traits. The study species differed dramatically in traits related to the timing of seed release, and a strong species by season interaction affected most aspects of the spatial pattern of dispersed seeds. A rich interplay among plant traits and seasonal differences in atmo- spheric conditions caused this interaction. Several of the same plant traits are crucial for both seed dispersal and other aspects of life history variation. Observed traits that limit dispersal are likely to be constrained by their life history consequences. atmospheric turbulence 􏰚 conditional seed release 􏰚 Coupled Eulerian-Lagrangian closure (CELC) model 􏰚 long distance dispersal 􏰚 tropical forest
Movement ecology of migration in turkey vultures
We develop individual-based movement ecology models (MEM) to explore turkey vulture (Cathartes aura) migration decisions at both hourly and daily scales. Vulture movements in 10 migration events were recorded with satellite-reporting GPS sensors, and flight behavior was observed visually, aided by on-the-ground VHF radio-track- ing. We used the North American Regional Reanalysis dataset to obtain values for wind speed, turbulent kinetic energy (TKE), and cloud height and used a digital elevation model for a measure of terrain ruggedness. A turkey vulture fitted with a heart-rate logger during 124 h of flight during 38 contiguous days showed only a small increase in mean heart rate as distance traveled per day increased, which suggests that, unlike flapping, soaring flight does not lead to greatly increased metabolic costs. Data from 10 migrations for 724 hourly segments and 152 daily segments showed that vultures depended heavily upon high levels of TKE in the atmospheric bound- ary layer to increase flight distances and maintain preferred bearings at both hourly and daily scales. We suggest how the MEM can be extended to other spatial and temporal scales of avian migration. Our success in relating model-derived atmospheric variables to migration indicates the potential of using regional reanalysis data, as here, and potentially other regional, higher-resolution, atmospheric models in predicting changing movement patterns of soaring birds under var- ious scenarios of climate and land use change. energetics 􏰚 flight 􏰚 meteorology
Southern Ocean acidification: A tipping point at 450-ppm atmospheric CO2
Southern Ocean acidification via anthropogenic CO2 uptake is expected to be detrimental to multiple calcifying plankton species by lowering the concentration of carbonate ion (CO32􏱉) to levels where calcium carbonate (both aragonite and calcite) shells begin to dissolve. Natural seasonal variations in carbonate ion concentrations could either hasten or dampen the future onset of this undersaturation of calcium carbonate. We present a large-scale Southern Ocean observational analysis that examines the seasonal magnitude and variability of CO32􏱉 and pH. Our analysis shows an intense wintertime minimum in CO32􏱉 south of the Antarctic Polar Front and when combined with anthropogenic CO2 uptake is likely to induce aragonite undersaturation when atmospheric CO2 levels reach 􏰜450 ppm. Under the IPCC IS92a scenario, Southern Ocean wintertime aragonite undersaturation is projected to occur by the year 2030 and no later than 2038. Some prominent calcifying plankton, in particular the Pteropod species Limacina helicina, have important veliger larval development during winter and will have to experience detrimental carbonate conditions much earlier than previously thought, with possible deleterious flow-on impacts for the wider Southern Ocean marine ecosystem. Our results highlight the critical importance of understanding seasonal carbon dynamics within all calcifying marine ecosystems such as continental shelves and coral reefs, because natural variability may potentially hasten the onset of future ocean acidification. carbon cycle 􏰚 climate change
Multiple movement modes by large herbivores at multiple spatiotemporal scales
Recent theory suggests that animals should switch facultatively among canonical movement modes as a complex function of internal state, landscape characteristics, motion capacity, and navigational capacity. We tested the generality of this paradigm for free-ranging elk (Cervus elaphus) over 5 orders of magnitude in time (minutes to years) and space (meters to 100 km). At the coarsest spatiotemporal scale, elk shifted from a dispersive to a home-ranging phase over the course of 1–3 years after introduc- tion into a novel environment. At intermediate spatiotemporal scales, elk continued to alternate between movement modes. During the dispersive phase, elk alternated between encamped and exploratory modes, possibly linked to changes in motivational goals from foraging to social bonding. During the home-ranging phase, elk movements were characterized by a complex interplay between attraction to preferred habitat types and memory of previous movements across the home-range. At the finest tempo- ral and spatial scale, elk used area-restricted search while brows- ing, interspersed with less sinuous paths when not browsing. Encountering a patch of high-quality food plants triggered the switch from one mode to the next, creating biphasic movement dynamics that were reinforced by local resource heterogeneity. These patterns suggest that multiphasic structure is fundamental to the movement patterns of elk at all temporal and spatial scales tested. elk 􏰚 foraging 􏰚 group formation 􏰚 motivation
The movement ecology and dynamics of plant communities in fragmented landscapes
A conceptual model of movement ecology has recently been advanced to explain all movement by considering the interaction of four elements: internal state, motion capacity, navigation capacities, and external factors. We modified this framework to generate predictions for species richness dynamics of fragmented plant communities and tested them in experimental landscapes across a 7-year time series. We found that two external factors, dispersal vectors and habitat features, affected species coloniza- tion and recolonization in habitat fragments and their effects varied and depended on motion capacity. Bird-dispersed species richness showed connectivity effects that reached an asymptote over time, but no edge effects, whereas wind-dispersed species richness showed steadily accumulating edge and connectivity effects, with no indication of an asymptote. Unassisted species also showed increasing differences caused by connectivity over time, whereas edges had no effect. Our limited use of proxies for movement ecology (e.g., dispersal mode as a proxy for motion capacity) resulted in moderate predictive power for communities and, in some cases, highlighted the importance of a more complete understanding of movement ecology for predicting how landscape conservation actions affect plant community dynamics. corridors 􏰚 dispersal 􏰚 diversity 􏰚 life-history traits 􏰚 species richness
Measuring the effectiveness of protected area networks in reducing deforestation
Global efforts to reduce tropical deforestation rely heavily on the establishment of protected areas. Measuring the effectiveness of these areas is difficult because the amount of deforestation that would have occurred in the absence of legal protection cannot be directly observed. Conventional methods of evaluating the effectiveness of protected areas can be biased because protection is not randomly assigned and because protection can induce deforesta- tion spillovers (displacement) to neighboring forests. We demon- strate that estimates of effectiveness can be substantially im- proved by controlling for biases along dimensions that are observable, measuring spatial spillovers, and testing the sensitivity of estimates to potential hidden biases. We apply matching meth- ods to evaluate the impact on deforestation of Costa Rica’s re- nowned protected-area system between 1960 and 1997. We find that protection reduced deforestation: approximately 10% of the protected forests would have been deforested had they not been protected. Conventional approaches to evaluating conservation impact, which fail to control for observable covariates correlated with both protection and deforestation, substantially overesti- mate avoided deforestation (by over 65%, based on our estimates). We also find that deforestation spillovers from protected to un- protected forests are negligible. Our conclusions are robust to potential hidden bias, as well as to changes in modeling assump- tions. Our results show that, with appropriate empirical methods, conservation scientists and policy makers can better understand the relationships between human and natural systems and can use this to guide their attempts to protect critical ecosystem services. avoided deforestation 􏰚 conservation policy 􏰚 empirical evaluation 􏰚 spatial spillovers
Trends and missing parts in the study of movement ecology
Movement is important to all organisms, and accordingly it is addressed in a huge number of papers in the literature. Of nearly 26,000 papers referring to movement, an estimated 34% focused on movement by measuring it or testing hypotheses about it. This enormous amount of information is difficult to review and high- lights the need to assess the collective completeness of movement studies and identify gaps. We surveyed 1,000 randomly selected papers from 496 journals and compared the facets of movement studied with a suggested framework for movement ecology, consisting of internal state (motivation, physiology), motion and navigation capacities, and external factors (both the physical environment and living organisms), and links among these com- ponents. Most studies simply measured and described the move- ment of organisms without reference to ecological or internal factors, and the most frequently studied part of the framework was the link between external factors and motion capacity. Few studies looked at the effects on movement of navigation capacity, or internal state, and those were mainly from vertebrates. For invertebrates and plants most studies were at the population level, whereas more vertebrate studies were conducted at the individual level. Consideration of only population-level averages promul- gates neglect of between-individual variation in movement, po- tentially hindering the study of factors controlling movement. Terminology was found to be inconsistent among taxa and sub- disciplines. The gaps identified in coverage of movement studies highlight research areas that should be addressed to fully under- stand the ecology of movement. dispersal 􏰚 foraging 􏰚 migration 􏰚 navigation 􏰚 physiology
Translocation experiments with butterflies reveal limits to enhancement of poleward populations under climate change
There is a pressing need to predict how species will change their geographic ranges under climate change. Projections typically assume that temperature is a primary fitness determinant and that populations near the poleward (and upward) range boundary are preadapted to warming. Thus, poleward, peripheral populations will increase with warming, and these increases facilitate poleward range expansions. We tested the assumption that poleward, pe- ripheral populations are enhanced by warming using 2 butterflies (Erynnis propertius and Papilio zelicaon) that co-occur and have contrasting degrees of host specialization and interpopulation genetic differentiation. We performed a reciprocal translocation experiment between central and poleward, peripheral populations in the field and simulated a translocation experiment that included alternate host plants. We found that the performance of both central and peripheral populations of E. propertius were enhanced during the summer months by temperatures characteristic of the range center but that local adaptation of peripheral populations to winter conditions near the range edge could counteract that enhancement. Further, poleward range expansion in this species is prevented by a lack of host plants. In P. zelicaon, the fitness of central and peripheral populations decreased under extreme sum- mer temperatures that occurred in the field at the range center. Performance in this species also was affected by an interaction of temperature and host plant such that host species strongly medi- ated the fitness of peripheral individuals under differing simulated temperatures. Altogether we have evidence that facilitation of poleward range shifts through enhancement of peripheral populations is unlikely in either study species. Lepidoptera 􏰚 range center 􏰚 range expansion 􏰚 range periphery
The future of ice sheets and sea ice: Between reversible retreat and unstoppable loss
We discuss the existence of cryospheric “tipping points” in the Earth’s climate system. Such critical thresholds have been sug- gested to exist for the disappearance of Arctic sea ice and the retreat of ice sheets: Once these ice masses have shrunk below an anticipated critical extent, the ice–albedo feedback might lead to the irreversible and unstoppable loss of the remaining ice. We here give an overview of our current understanding of such thresh- old behavior. By using conceptual arguments, we review the recent findings that such a tipping point probably does not exist for the loss of Arctic summer sea ice. Hence, in a cooler climate, sea ice could recover rapidly from the loss it has experienced in recent years. In addition, we discuss why this recent rapid retreat of Arc- tic summer sea ice might largely be a consequence of a slow shift in ice-thickness distribution, which will lead to strongly increased year-to-year variability of the Arctic summer sea-ice extent. This variability will render seasonal forecasts of the Arctic summer sea- ice extent increasingly difficult. We also discuss why, in contrast to Arctic summer sea ice, a tipping point is more likely to exist for the loss of the Greenland ice sheet and the West Antarctic ice sheet. Greenland | West Antarctic | climate change | tipping point | Arctic
Basic mechanism for abrupt monsoon transitions
Monsoon systems influence the livelihood of hundreds of millions of people. During the Holocene and last glacial period, rainfall in India and China has undergone strong and abrupt changes. Though details of monsoon circulations are complicated, observations reveal a defining moisture-advection feedback that dominates the seasonal heat balance and might act as an internal amplifier, leading to abrupt changes in response to relatively weak external perturbations. Here we present a minimal conceptual model capturing this positive feedback. The basic equations, motivated by observed relations, yield a threshold behavior, robust with respect to addition of other physical processes. Below this threshold in net radiative influx, Rc , no conventional monsoon can develop; above Rc , two stable regimes exist. We identify a nondimensional para- meter l that defines the threshold and makes monsoon systems comparable with respect to the character of their abrupt transition. This dynamic similitude may be helpful in understanding past and future variations in monsoon circulation. Within the restrictions of the model, we compute Rc for current monsoon systems in India, China, the Bay of Bengal, West Africa, North America, and Australia, where moisture advection is the main driver of the circulation. Earth system | tipping element | abrupt climate change | atmospheric circulation | nonlinear dynamics
The potential for behavioral thermoregulation to buffer ‘‘cold-blooded’’ animals against climate warming
Increasing concern about the impacts of global warming on biodi- versity has stimulated extensive discussion, but methods to trans- late broad-scale shifts in climate into direct impacts on living animals remain simplistic. A key missing element from models of climatic change impacts on animals is the buffering influence of behavioral thermoregulation. Here, we show how behavioral and mass/energy balance models can be combined with spatial data on climate, topography, and vegetation to predict impacts of in- creased air temperature on thermoregulating ectotherms such as reptiles and insects (a large portion of global biodiversity). We show that for most ‘‘cold-blooded’’ terrestrial animals, the primary thermal challenge is not to attain high body temperatures (al- though this is important in temperate environments) but to stay cool (particularly in tropical and desert areas, where ectotherm biodiversity is greatest). The impact of climate warming on ther- moregulating ectotherms will depend critically on how changes in vegetation cover alter the availability of shade as well as the animals’ capacities to alter their seasonal timing of activity and reproduction. Warmer environments also may increase mainte- nance energy costs while simultaneously constraining activity time, putting pressure on mass and energy budgets. Energy- and mass-balance models provide a general method to integrate the complexity of these direct interactions between organisms and climate into spatial predictions of the impact of climate change on biodiversity. This methodology allows quantitative organism- and habitat-specific assessments of climate change impacts. Australia 􏰚 biophysical model 􏰚 climate change 􏰚 terrestrial ectotherm 􏰚 GIS
Climate, carbon cycling, and deep-ocean ecosystem
Climate variation affects surface ocean processes and the production of organic carbon, which ultimately comprises the primary food supply to the deep-sea ecosystems that occupy 60% of the Earth’s surface. Warming trends in atmospheric and upper ocean temperatures, attributed to anthropogenic influence, have occurred over the past four decades. Changes in upper ocean temperature influence stratification and can affect the availability of nutrients for phytoplankton production. Global warming has been predicted to intensify stratification and reduce vertical mixing. Research also suggests that such reduced mixing will enhance variability in primary production and carbon export flux to the deep sea. The dependence of deep-sea communities on surface water production has raised important questions about how climate change will affect carbon cycling and deep-ocean ecosystem function. Recently, un- precedented time-series studies conducted over the past two decades in the North Pacific and the North Atlantic at >4,000-m depth have revealed unexpectedly large changes in deep-ocean ecosystems significantly correlated to climate-driven changes in the surface ocean that can impact the global carbon cycle. Climate-driven variation affects oceanic communities from surface waters to the much-overlooked deep sea and will have impacts on the global carbon cycle. Data from these two widely separated areas of the deep ocean provide compelling evidence that changes in climate can readily influence deep-sea processes. However, the limited geographic coverage of these existing time-series studies stresses the importance of developing a more global effort to monitor deep- sea ecosystems under modern conditions of rapidly changing climate.
Re-evaluation of forest biomass carbon stocks and lessons from the world’s most carbon-dense forests
From analysis of published global site biomass data (n = 136) from primary forests, we discovered (i) the world’s highest known total biomass carbon density (living plus dead) of 1,867 tonnes carbon per ha (average value from 13 sites) occurs in Australian temperate moist Eucalyptus regnans forests, and (ii) average values of the global site biomass data were higher for sampled temperate moist forests (n 􏰀 44) than for sampled tropical (n 􏰀 36) and boreal (n 􏰀 52) forests (n is number of sites per forest biome). Spatially averaged Intergovern- mental Panel on Climate Change biome default values are lower than our average site values for temperate moist forests, because the temperate biome contains a diversity of forest ecosystem types that support a range of mature carbon stocks or have a long land-use history with reduced carbon stocks. We describe a framework for identifying forests important for carbon storage based on the factors that account for high biomass carbon densities, including (i) relatively cool temperatures and moderately high precipitation producing rates of fast growth but slow decomposition, and (ii) older forests that are often multiaged and multilayered and have experienced minimal human disturbance. Our results are relevant to negotiations under the United Nations Framework Convention on Climate Change re- garding forest conservation, management, and restoration. Conserv- ing forests with large stocks of biomass from deforestation and degradation avoids significant carbon emissions to the atmosphere, irrespective of the source country, and should be among allowable mitigation activities. Similarly, management that allows restoration of a forest’s carbon sequestration potential also should be recognized. Eucalyptus regnans 􏰂 climate mitigation 􏰂 primary forest 􏰂 deforestation and degradation 􏰂 temperate moist forest biome
Multidimensional evaluation of managed relocation
Managed relocation (MR) has rapidly emerged as a potential intervention strategy in the toolbox of biodiversity management under climate change. Previous authors have suggested that MR (also referred to as assisted colonization, assisted migration, or assisted translocation) could be a last-alternative option after interrogating a linear decision tree. We argue that numerous interacting and value-laden considerations demand a more inclu- sive strategy for evaluating MR. The pace of modern climate change demands decision making with imperfect information, and tools that elucidate this uncertainty and integrate scientific information and social values are urgently needed. We present a heuristic tool that incorporates both ecological and social criteria in a multidimensional decision-making framework. For visualization purposes, we collapse these criteria into 4 classes that can be depicted in graphical 2-D space. This framework offers a pragmatic approach for summarizing key dimensions of MR: capturing un- certainty in the evaluation criteria, creating transparency in the evaluation process, and recognizing the inherent tradeoffs that different stakeholders bring to evaluation of MR and its alternatives. assisted migration 􏰀 climate change 􏰀 conservation biology 􏰀 conservation strategy 􏰀 sustainability science
Synchronous extinction of North America’s Pleistocene mammals
The late Pleistocene witnessed the extinction of 35 genera of North American mammals. The last appearance dates of 16 of these genera securely fall between 12,000 and 10,000 radiocarbon years ago (􏰂13,800–11,400 calendar years B.P.), although whether the absence of fossil occurrences for the remaining 19 genera from this time interval is the result of sampling error or temporally staggered extinctions is unclear. Analysis of the chronology of extinctions suggests that sampling error can explain the absence of terminal Pleistocene last appearance dates for the remaining 19 genera. The extinction chronology of North American Pleistocene mammals therefore can be characterized as a synchronous event that took place 12,000–10,000 radiocarbon years B.P. Results favor an ex- tinction mechanism that is capable of wiping out up to 35 genera across a continent in a geologic instant. climate change 􏰀 extraterrestrial impact 􏰀 overkill 􏰀 Quaternary extinctions 􏰀 radiocarbon dates
The physical basis for increases in precipitation extremes in simulations of 21st-century climate change
Global warming is expected to lead to a large increase in atmospheric water vapor content and to changes in the hydrological cycle, which include an intensification of precipitation extremes. The intensity of precipitation extremes is widely held to increase proportionately to the increase in atmospheric water vapor content. Here, we show that this is not the case in 21st-century climate change scenarios simulated with climate models. In the tropics, precipitation extremes are not simulated reliably and do not change consistently among climate models; in the extratropics, they consistently increase more slowly than atmospheric water vapor content. We give a physical basis for how precipitation extremes change with climate and show that their changes depend on changes in the moist-adiabatic temperature lapse rate, in the upward velocity, and in the temperature when precipitation ex- tremes occur. For the tropics, the theory suggests that improving the simulation of upward velocities in climate models is essential for improving predictions of precipitation extremes; for the extra- tropics, agreement with theory and the consistency among climate models increase confidence in the robustness of predictions of precipitation extremes under climate change. global warming 􏰀 hydrological cycle 􏰀 rainfall 􏰀 extreme events
Overcoming systemic roadblocks to sustainability: The evolutionary redesign of worldviews, institutions, and technologies
A high and sustainable quality of life is a central goal for humanity. Our current socio-ecological regime and its set of interconnected worldviews, institutions, and technologies all support the goal of unlimited growth of material production and consumption as a proxy for quality of life. However, abundant evidence shows that, beyond a certain threshold, further material growth no longer significantly contributes to improvement in quality of life. Not only does further material growth not meet humanity’s central goal, there is mounting evidence that it creates significant roadblocks to sustainability through increasing resource constraints (i.e., peak oil, water limitations) and sink constraints (i.e., climate disruption). Overcoming these roadblocks and creating a sustainable and de- sirable future will require an integrated, systems level redesign of our socio-ecological regime focused explicitly and directly on the goal of sustainable quality of life rather than the proxy of unlimited material growth. This transition, like all cultural transitions, will occur through an evolutionary process, but one that we, to a certain extent, can control and direct. We suggest an integrated set of worldviews, institutions, and technologies to stimulate and seed this evolutionary redesign of the current socio-ecological regime to achieve global sustainability. cultural adaptation 􏰀 ecology 􏰀 societal decline
Irreversible climate change due to carbon dioxide emissions
The severity of damaging human-induced climate change depends not only on the magnitude of the change but also on the potential for irreversibility. This paper shows that the climate change that takes place due to increases in carbon dioxide concentration is largely irreversible for 1,000 years after emissions stop. Following cessation of emissions, removal of atmospheric carbon dioxide decreases radiative forcing, but is largely compensated by slower loss of heat to the ocean, so that atmospheric temperatures do not drop significantly for at least 1,000 years. Among illustrative irreversible impacts that should be expected if atmospheric carbon dioxide concentrations increase from current levels near 385 parts per million by volume (ppmv) to a peak of 450 – 600 ppmv over the coming century are irreversible dry-season rainfall reductions in several regions comparable to those of the ‘‘dust bowl’’ era and inexorable sea level rise. Thermal expansion of the warming ocean provides a conservative lower limit to irreversible global average sea level rise of at least 0.4–1.0 m if 21st century CO2 concentrations exceed 600 ppmv and 0.6 –1.9 m for peak CO2 concentrations exceeding 1,000 ppmv. Additional contributions from glaciers and ice sheet contributions to future sea level rise are uncertain but may equal or exceed several meters over the next millennium or longer. dangerous interference 􏰀 precipitation 􏰀 sea level rise 􏰀 warming
Maximum leaf conductance driven by CO2 effects on stomatal size and density over geologic time
Stomatal pores are microscopic structures on the epidermis of leaves formed by 2 specialized guard cells that control the exchange of water vapor and CO2 between plants and the atmosphere. Stomatal size (S) and density (D) determine maximum leaf diffusive (stomatal) conductance of CO2 (gcmax) to sites of assimi- lation. Although large variations in D observed in the fossil record have been correlated with atmospheric CO2, the crucial significance of similarly large variations in S has been overlooked. Here, we use physical diffusion theory to explain why large changes in S nec- essarily accompanied the changes in D and atmospheric CO2 over the last 400 million years. In particular, we show that high densities of small stomata are the only way to attain the highest gcmax values required to counter CO2‘‘starvation’’ at low atmospheric CO2 concentrations. This explains cycles of increasing D and decreasing S evident in the fossil history of stomata under the CO2 impover- ished atmospheres of the Permo-Carboniferous and Cenozoic gla- ciations. The pattern was reversed under rising atmospheric CO2 regimes. Selection for small S was crucial for attaining high gcmax under falling atmospheric CO2 and, therefore, may represent a mechanism linking CO2 and the increasing gas-exchange capacity of land plants over geologic time. Phanerozoic 􏰀 photosynthesis 􏰀 plant evolution 􏰀 transpiration 􏰀 xylem
Temperature sensitivity of drought-induced tree mortality portends increased regional die-off under global change-type drought
Large-scale biogeographical shifts in vegetation are predicted in response to the altered precipitation and temperature regimes associated with global climate change. Vegetation shifts have profound ecological impacts and are an important climate-ecosystem feedback through their alteration of carbon, water, and energy exchanges of the land surface. Of particular concern is the potential for warmer temperatures to compound the effects of increasingly severe droughts by triggering widespread vegetation shifts via woody plant mortality. The sensitivity of tree mortality to temperature is dependent on which of 2 non-mutually-exclusive mechanisms predominates—temperature-sensitive carbon starvation in response to a period of protracted water stress or temperature-insensitive sudden hydraulic failure under extreme water stress (cavitation). Here we show that experimentally induced warmer temperatures (􏰂4 °C) shortened the time to drought- induced mortality in Pinus edulis (pin ̃ on shortened pine) trees by nearly a third, with temperature-dependent differences in cumu- lative respiration costs implicating carbon starvation as the primary mechanism of mortality. Extrapolating this temperature effect to the historic frequency of water deficit in the southwestern United States predicts a 5-fold increase in the frequency of regional-scale tree die-off events for this species due to temperature alone. Projected increases in drought frequency due to changes in pre- cipitation and increases in stress from biotic agents (e.g., bark beetles) would further exacerbate mortality. Our results demon- strate the mechanism by which warmer temperatures have exac- erbated recent regional die-off events and background mortality rates. Because of pervasive projected increases in temperature, our results portend widespread increases in the extent and frequency of vegetation die-off. biosphere–atmosphere feedbacks 􏰀 drought impacts 􏰀 global-change ecology 􏰀 Pinus edulis 􏰀 carbon starvation