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Recent reports of net ecosysytem production >100 g C m -2 yr -1 in deserts are incompatible with existing measurements of net primary production and carbon pools in deserts. The comparisions suggest that gas exchange measurements should be used with caution and better validation if they are expected to indicate the magnitude of carbon sink in these ecosysytems. ?? 2009 Blackwell Publishing.
Marshes worldwide are actively degrading in response to increased sea level rise rates and reduced sediment delivery, though the growth rate of vegetation plays a critical role in determining their stability. We have compiled 56 measurements of above-ground annual productivity for Spartina alterniflora, the dominant macrophyte in North American coastal wetlands. Our compilation indicates a significant latitudinal gradient in productivity, which we interpret to be determined primarily by temperature and/or the length of growing season. Simple linear regression yields a 27 g m-2 yr -1 increase in productivity with an increase of mean annual temperature by one degree C. If temperatures warm 2?4 C over the next century, then marsh productivity may increase by 10?40%, though physiological research suggests that increases in the north could potentially be offset by some decreases in the south. This increase in productivity is roughly equivalent to estimates of marsh lost due to future sea level change. If a warming-induced stimulation of vegetation growth will enhance vertical accretion and limit erosion, then the combined effects of global change may be to increase the total productivity and ecosystem services of tidal wetlands, at least in Northern latitudes.
Mountain terrestrial and aquatic ecosystems are responsive to external drivers of change, especially climate change and atmospheric deposition of nitrogen (N). We explored the consequences of a temperature-warming trend on stream nitrate in an alpine and subalpine watershed in the Colorado Front Range that has long been the recipient of elevated atmospheric N deposition. Mean annual stream nitrate concentrations since 2000 are higher by 50% than an earlier monitoring period of 1991-1999. Mean annual N export increased by 28% from 2.03 kg N ha -1 yr -1 before 2000 to 2.84 kg N ha -1 yr -1 in Loch Vale watershed since 2000. The substantial increase in N export comes as a surprise, since mean wet atmospheric N deposition from 1991 to 2006 (3.06 kg N ha -1 yr -1 ) did not increase. There has been a period of below average precipitation from 2000 to 2006 and a steady increase in summer and fall temperatures of 0.12??C yr -1 in both seasons since 1991. Nitrate concentrations, as well as the weathering products calcium and sulfate, were higher for the period 2000-2006 in rock glacier meltwater at the top of the watershed above the influence of alpine and subalpine vegetation and soils. We conclude the observed recent N increases in Loch Vale are the result of warmer summer and fall mean temperatures that are melting ice in glaciers and rock glaciers. This, in turn, has exposed sediments from which N produced by nitrification can be flushed. We suggest a water quality threshold may have been crossed around 2000. The phenomenon observed in Loch Vale may be indicative of N release from ice features such as rock glaciers worldwide as mountain glaciers retreat. ?? 2009 Blackwell Publishing Ltd.
I examined the potential influence of climate change on the dynamics of a previously studied hybrid zone between a pair of terrestrial salamanders at the Coweeta Hydrologic Laboratory, U.S. Forest Service, in the Nantahala Mountains of North Carolina, USA. A 16-year study led by Nelson G. Hairston, Sr. revealed that Plethodon teyahalee and Plethodon shermani hybridized at intermediate elevations, forming a cline between 'pure' parental P. teyahalee at lower elevations and 'pure' parental P. shermani at higher elevations. From 1974 to 1990 the proportion of salamanders at the higher elevation scored as 'pure' P. shermani declined significantly, indicating that the hybrid zone was spreading upward. To date there have been no rigorous tests of hypotheses for the movement of this hybrid zone. Using temperature and precipitation data from Coweeta, I re-analyzed Hairston's data to examine whether the observed elevational shift was correlated with variation in either air temperature or precipitation from the same time period. For temperature, my analysis tracked the results of the original study: the proportion of 'pure' P. shermani at the higher elevation declined significantly with increasing mean annual temperature, whereas the proportion of 'pure' P. teyahalee at lower elevations did not. There was no discernable relationship between proportions of 'pure' individuals of either species with variation in precipitation. From 1974 to 1990, low-elevation air temperatures at the Coweeta Laboratory ranged from annual means of 11.8 to 14.2 °C, compared with a 55-year average (1936-1990) of 12.6 °C. My re-analyses indicate that the upward spread of the hybrid zone is correlated with increasing air temperatures, but not precipitation, and provide an empirical test of a hypothesis for one factor that may have influenced this movement. My results aid in understanding the potential impact that climate change may have on the ecology and evolution of terrestrial salamanders in montane regions.
The boreal forest contains large reserves of carbon. Across this region, wildfires influence the temporal and spatial dynamics of carbon storage. In this study, we estimate fire emissions and changes in carbon storage for boreal North America over the 21st century. We use a gridded data set developed with a multivariate adaptive regression spline approach to determine how area burned varies each year with changing climatic and fuel moisture conditions. We apply the process‐based Terrestrial Ecosystem Model to evaluate the role of future fire on the carbon dynamics of boreal North America in the context of changing atmospheric carbon dioxide (CO 2 ) concentration and climate in the A2 and B2 emissions scenarios of the CGCM2 global climate model. Relative to the last decade of the 20th century, decadal total carbon emissions from fire increase by 2.5–4.4 times by 2091–2100, depending on the climate scenario and assumptions about CO 2 fertilization. Larger fire emissions occur with warmer climates or if CO 2 fertilization is assumed to occur. Despite the increases in fire emissions, our simulations indicate that boreal North America will be a carbon sink over the 21st century if CO 2 fertilization is assumed to occur in the future. In contrast, simulations excluding CO 2 fertilization over the same period indicate that the region will change to a carbon source to the atmosphere, with the source being 2.1 times greater under the warmer A2 scenario than the B2 scenario. To improve estimates of wildfire on terrestrial carbon dynamics in boreal North America, future studies should incorporate the role of dynamic vegetation to represent more accurately post‐fire successional processes, incorporate fire severity parameters that change in time and space, account for human influences through increased fire suppression, and integrate the role of other disturbances and their interactions with future fire regime.
Shifts in the timing of spring phenology are a central feature of global change research. Long-term observations of plant phenology have been used to track vegetation responses to climate variability but are often limited to particular species and locations and may not represent synoptic patterns. Satellite remote sensing is instead used for continental to global monitoring. Although numerous methods exist to extract phenological timing, in particular start-of-spring (SOS), from time series of reflectance data, a comprehensive intercomparison and interpretation of SOS methods has not been conducted. Here, we assess 10 SOS methods for North America between 1982 and 2006. The techniques include consistent inputs from the 8 km Global Inventory Modeling and Mapping Studies Advanced Very High Resolution Radiometer NDVIg dataset, independent data for snow cover, soil thaw, lake ice dynamics, spring streamflow timing, over 16 000 individual measurements of ground-based phenology, and two temperature-driven models of spring phenology. Compared with an ensemble of the 10 SOS methods, we found that individual methods differed in average day-of-year estimates by ±60 days and in standard deviation by ±20 days. The ability of the satellite methods to retrieve SOS estimates was highest in northern latitudes and lowest in arid, tropical, and Mediterranean ecoregions. The ordinal rank of SOS methods varied geographically, as did the relationships between SOS estimates and the cryospheric/hydrologic metrics. Compared with ground observations, SOS estimates were more related to the first leaf and first flowers expanding phenological stages. We found no evidence for time trends in spring arrival from ground- or model-based data; using an ensemble estimate from two methods that were more closely related to ground observations than other methods, SOS trends could be detected for only 12% of North America and were divided between trends towards both earlier and later spring.
Drought- and insect-associated tree mortality at low-elevation ecotones is a widespread phenomenon but the underlying mechanisms are uncertain. Enhanced growth sensitivity to climate is widely observed among trees that die, indicating that a predisposing physiological mechanism(s) underlies tree mortality. We tested three, linked hypotheses regarding mortality using a ponderosa pine ( Pinus ponderosa ) elevation transect that experienced low-elevation mortality following prolonged drought. The hypotheses were: (1) mortality was associated with greater growth sensitivity to climate, (2) mortality was associated with greater sensitivity of gas exchange to climate, and (3) growth and gas exchange were correlated. Support for all three hypotheses would indicate that mortality results at least in part from gas exchange constraints. We assessed growth using basal area increment normalized by tree basal area [basal area increment (BAI)/basal area (BA)] to account for differences in tree size. Whole-crown gas exchange was indexed via estimates of the CO 2 partial pressure difference between leaf and atmosphere ( p a − p c ) derived from tree ring carbon isotope ratios ( δ 13 C), corrected for temporal trends in atmospheric CO 2 and δ 13 C and elevation trends in pressure. Trees that survived the drought exhibited strong correlations among and between BAI, BAI/BA, p a − p c , and climate. In contrast, trees that died exhibited greater growth sensitivity to climate than trees that survived, no sensitivity of p a − p c to climate, and a steep relationship between p a − p c and BAI/BA. The p a − p c results are consistent with predictions from a theoretical hydraulic model, suggesting trees that died had a limited buffer between mean water availability during their lifespan and water availability during drought – i.e., chronic water stress. It appears that chronic water stress predisposed low-elevation trees to mortality during drought via constrained gas exchange. Continued intensification of drought in mid-latitude regions may drive increased mortality and ecotone shifts in temperate forests and woodlands.
Revitalization of degraded landscapes may provide sinks for rising atmospheric CO2, especially in reconstructed prairies where substantial belowground productivity is coupled with large soil organic carbon (SOC) deficits after many decades of cultivation. The restoration process also provides opportunities to study the often-elusive factors that regulate soil processes. Although the precise mechanisms that govern the rate of SOC accrual are unclear, factors such as soil moisture or vegetation type may influence the net accrual rate by affecting the balance between organic matter inputs and decomposition. A resampling approach was used to assess the control that soil moisture and plant community type each exert on SOC and total nitrogen (TN) accumulation in restored grasslands. Five plots that varied in drainage were sampled at least four times over two decades to assess SOC, TN, and C4- and C3-derived C. We found that higher long-term soil moisture, characterized by low soil magnetic susceptibility, promoted SOC and TN accrual, with twice the SOC and three times the TN gain in seasonally saturated prairies compared with mesic prairies. Vegetation also influenced SOC and TN recovery, as accrual was faster in the prairies compared with C3-only grassland, and C4-derived C accrual correlated strongly to total SOC accrual but C3-C did not. High SOC accumulation at the surface (0-10 cm) combined with losses at depth (10-20 cm) suggested these soils are recovering the highly stratified profiles typical of remnant prairies. Our results suggest that local hydrology and plant community are critical drivers of SOC and TN recovery in restored grasslands. Because these factors and the way they affect SOC are susceptible to modification by climate change, we contend that predictions of the C-sequestration performance of restored grasslands must account for projected climatic changes on both soil moisture and the seasonal productivity of C4 and C3 plants. ?? 2009 Blackwell Publishing Ltd.
The fate of carbon (C) contained within permafrost in boreal forest environments is an important consideration for the current and future carbon cycle as soils warm in northern latitudes. Currently, little is known about the microbiology or chemistry of permafrost soils that may affect its decomposition once soils thaw. We tested the hypothesis that low microbial abundances and activities in permafrost soils limit decomposition rates compared with active layer soils. We examined active layer and permafrost soils near Fairbanks, AK, the Yukon River, and the Arctic Circle. Soils were incubated in the lab under aerobic and anaerobic conditions. Gas fluxes at -5 and 5 ??C were measured to calculate temperature response quotients (Q10). The Q10 was lower in permafrost soils (average 2.7) compared with active layer soils (average 7.5). Soil nutrients, leachable dissolved organic C (DOC) quality and quantity, and nuclear magnetic resonance spectroscopy of the soils revealed that the organic matter within permafrost soils is as labile, or even more so, than surface soils. Microbial abundances (fungi, bacteria, and subgroups: methanogens and Basidiomycetes) and exoenzyme activities involved in decomposition were lower in permafrost soils compared with active layer soils, which, together with the chemical data, supports the reduced Q10 values. CH4 fluxes were correlated with methanogen abundance and the highest CH4 production came from active layer soils. These results suggest that permafrost soils have high inherent decomposability, but low microbial abundances and activities reduce the temperature sensitivity of C fluxes. Despite these inherent limitations, however, respiration per unit soil C was higher in permafrost soils compared with active layer soils, suggesting that decomposition and heterotrophic respiration may contribute to a positive feedback to warming of this eco region. Published 2010. This article is a US Government work and is in the public domain in the USA.
We combined year‐round eddy covariance with biometry and biomass harvests along a chronosequence of boreal forest stands that were 1, 6, 15, 23, 40, ∼74, and ∼154 years old to understand how ecosystem production and carbon stocks change during recovery from stand‐replacing crown fire. Live biomass (C live ) was low in the 1‐ and 6‐year‐old stands, and increased following a logistic pattern to high levels in the 74‐ and 154‐year‐old stands. Carbon stocks in the forest floor (C forest floor ) and coarse woody debris (C CWD ) were comparatively high in the 1‐year‐old stand, reduced in the 6‐ through 40‐year‐old stands, and highest in the 74‐ and 154‐year‐old stands. Total net primary production (TNPP) was reduced in the 1‐ and 6‐year‐old stands, highest in the 23‐ through 74‐year‐old stands and somewhat reduced in the 154‐year‐old stand. The NPP decline at the 154‐year‐old stand was related to increased autotrophic respiration rather than decreased gross primary production (GPP). Net ecosystem production (NEP), calculated by integrated eddy covariance, indicated the 1‐ and 6‐year‐old stands were losing carbon, the 15‐year‐old stand was gaining a small amount of carbon, the 23‐ and 74‐year‐old stands were gaining considerable carbon, and the 40‐ and 154‐year‐old stands were gaining modest amounts of carbon. The recovery from fire was rapid; a linear fit through the NEP observations at the 6‐ and 15‐year‐old stands indicated the transition from carbon source to sink occurred within 11–12 years. The NEP decline at the 154‐year‐old stand appears related to increased losses from C live by tree mortality and possibly from C forest floor by decomposition. Our findings support the idea that NPP, carbon production efficiency (NPP/GPP), NEP, and carbon storage efficiency (NEP/TNPP) all decrease in old boreal stands.
High‐latitude regions store large amounts of organic carbon (OC) in active‐layer soils and permafrost, accounting for nearly half of the global belowground OC pool. In the boreal region, recent warming has promoted changes in the fire regime, which may exacerbate rates of permafrost thaw and alter soil OC dynamics in both organic and mineral soil. We examined how interactions between fire and permafrost govern rates of soil OC accumulation in organic horizons, mineral soil of the active layer, and near‐surface permafrost in a black spruce ecosystem of interior Alaska. To estimate OC accumulation rates, we used chronosequence, radiocarbon, and modeling approaches. We also developed a simple model to track long‐term changes in soil OC stocks over past fire cycles and to evaluate the response of OC stocks to future changes in the fire regime. Our chronosequence and radiocarbon data indicate that OC turnover varies with soil depth, with fastest turnover occurring in shallow organic horizons (∼60 years) and slowest turnover in near‐surface permafrost (>3000 years). Modeling analysis indicates that OC accumulation in organic horizons was strongly governed by carbon losses via combustion and burial of charred remains in deep organic horizons. OC accumulation in mineral soil was influenced by active layer depth, which determined the proportion of mineral OC in a thawed or frozen state and thus, determined loss rates via decomposition. Our model results suggest that future changes in fire regime will result in substantial reductions in OC stocks, largely from the deep organic horizon. Additional OC losses will result from fire‐induced thawing of near‐surface permafrost. From these findings, we conclude that the vulnerability of deep OC stocks to future warming is closely linked to the sensitivity of permafrost to wildfire disturbance.
Given the threatened status of many primate species, the impacts of global warming on primate reproduction and, consequently, population growth should be of concern. We examined relations between climatic variability and birth seasonality, offspring production, and infant sex ratios in two ateline primates, northern muriquis, and woolly monkeys. In both species, the annual birth season was delayed by dry conditions and El Niño years, and delayed birth seasons were linked to lower birth rates. Additionally, increased mean annual temperatures were associated with lower birth rates for northern muriquis. Offspring sex ratios varied with climatic conditions in both species, but in different ways: directly in woolly monkeys and indirectly in northern muriquis. Woolly monkeys displayed an increase in the proportion of males among offspring in association with El Niño events, whereas in northern muriquis, increases in the proportion of males among offspring were associated with delayed onset of the birth season, which itself was related, although weakly, to warm, dry conditions. These results illustrate that global warming, increased drought frequency, and changes in the frequency of El Niño events could limit primate reproductive output, threatening the persistence and recovery of ateline primate populations.
During the past ∼50 years, the number and area of lakes have declined in several regions in boreal forests. However, there has been substantial finer-scale heterogeneity; some lakes decreased in area, some showed no trend, and others increased. The objective of this study was to identify the primary mechanisms underlying heterogeneous trends in closed-basin lake area. Eight lake characteristics ( δ 18 O, electrical conductivity, surface : volume index, bank slope, floating mat width, peat depth, thaw depth at shoreline, and thaw depth at the forest boundary) were compared for 15 lake pairs in Alaskan boreal forest where one lake had decreased in area since ∼1950, and the other had not. Mean differences in characteristics between paired lakes were used to identify the most likely of nine mechanistic scenarios that combined three potential mechanisms for decreasing lake area (talik drainage, surface water evaporation, and terrestrialization) with three potential mechanisms for nondecreasing lake area (subpermafrost groundwater recharge through an open talik, stable permafrost, and thermokarst). A priori expectations of the direction of mean differences between decreasing and nondecreasing paired lakes were generated for each scenario. Decreasing lakes had significantly greater electrical conductivity, greater surface : volume indices, shallower bank slopes, wider floating mats, greater peat depths, and shallower thaw depths at the forest boundary. These results indicated that the most likely scenario was terrestrialization as the mechanism for lake area reduction combined with thermokarst as the mechanism for nondecreasing lake area. Terrestrialization and thermokarst may have been enhanced by recent warming which has both accelerated permafrost thawing and lengthened the growing season, thereby increasing plant growth, floating mat encroachment, transpiration rates, and the accumulation of organic matter in lake basins. The transition to peatlands associated with terrestrialization may provide a transient increase in carbon storage enhancing the role of northern ecosystems as major stores of global carbon.
The timing of spring leaf development, trajectories of summer leaf area, and the timing of autumn senescence have profound impacts to the water, carbon, and energy balance of ecosystems, and are likely influenced by global climate change. Limited field-based and remote-sensing observations have suggested complex spatial patterns related to geographic features that influence climate. However, much of this variability occurs at spatial scales that inhibit a detailed understanding of even the dominant drivers. Recognizing these limitations, we used nonlinear inverse modeling of medium-resolution remote sensing data, organized by day of year, to explore the influence of climate-related landscape factors on the timing of spring and autumn leaf-area trajectories in mid-Atlantic, USA forests. We also examined the extent to which declining summer greenness (greendown) degrades the precision and accuracy of observations of autumn offset of greenness. Of the dominant drivers of landscape phenology, elevation was the strongest, explaining up to 70% of the spatial variation in the onset of greenness. Urban land cover was second in importance, influencing spring onset and autumn offset to a distance of 32 km from large cities. Distance to tidal water also influenced phenological timing, but only within ~5 km of shorelines. Additionally, we observed that (i) growing season length unexpectedly increases with increasing elevation at elevations below 275 m; (ii) along gradients in urban land cover, timing of autumn offset has a stronger effect on growing season length than does timing of spring onset; and (iii) summer greendown introduces bias and uncertainty into observations of the autumn offset of greenness. These results demonstrate the power of medium grain analyses of landscape-scale phenology for understanding environmental controls on growing season length, and predicting how these might be affected by climate change.
Forest insects and pathogens are major disturbance agents that have affected millions of hectares in North America in recent decades, implying significant impacts to the carbon (C) cycle. Here, we review and synthesize published studies of the effects of biotic disturbances on forest C cycling in the United States and Canada. Primary productivity in stands was reduced, sometimes considerably, immediately following insect or pathogen attack. After repeated growth reductions caused by some insects or pathogens or a single infestation by some bark beetle species, tree mortality occurred, altering productivity and decomposition. In the years following disturbance, primary productivity in some cases increased rapidly as a result of enhanced growth by surviving vegetation, and in other cases increased slowly because of lower forest regrowth. In the decades following tree mortality, decomposition increased as a result of the large amount of dead organic matter. Net ecosystem productivity decreased immediately following attack, with some studies reporting a switch to a C source to the atmosphere, and increased afterward as the forest regrew and dead organic matter decomposed. Large variability in C cycle responses arose from several factors, including type of insect or pathogen, time since disturbance, number of trees affected, and capacity of remaining vegetation to increase growth rates following outbreak. We identified significant knowledge gaps, including limited understanding of carbon cycle impacts among different biotic disturbance types (particularly pathogens), their impacts at landscape and regional scales, and limited capacity to predict disturbance events and their consequences for carbon cycling. We conclude that biotic disturbances can have major impacts on forest C stocks and fluxes and can be large enough to affect regional C cycling. However, additional research is needed to reduce the uncertainties associated with quantifying biotic disturbance effects on the North American C budget.
Hotter and drier conditions projected for the southwestern United States can have a large impact on the abundance and composition of long-lived desert plant species. We used long-term vegetation monitoring results from 39 large plots across four protected sites in the Sonoran Desert region to determine how plant species have responded to past climate variability. This cross-site analysis identified the plant species and functional types susceptible to climate change, the magnitude of their responses, and potential climate thresholds. In the relatively mesic mesquite savanna communities, perennial grasses declined with a decrease in annual precipitation, cacti increased, and there was a reversal of the Prosopis velutina expansion experienced in the 20th century in response to increasing mean annual temperature (MAT). In the more xeric Arizona Upland communities, the dominant leguminous tree, Cercidium microphyllum, declined on hillslopes, and the shrub Fouquieria splendens decreased, especially on south- and west-facing slopes in response to increasing MAT. In the most xeric shrublands, the codominant species Larrea tridentata and its hemiparasite Krameria grayi decreased with a decrease in cool season precipitation and increased aridity, respectively. This regional-scale assessment of plant species response to recent climate variability is critical for forecasting future shifts in plant community composition, structure, and productivity.
We develop an approach for estimating net ecosystem exchange (NEE) using inventory-based information over North America (NA) for a recent 7-year period (ca. 2000–2006). The approach notably retains information on the spatial distribution of NEE, or the vertical exchange between land and atmosphere of all non-fossil fuel sources and sinks of CO 2 , while accounting for lateral transfers of forest and crop products as well as their eventual emissions. The total NEE estimate of a -327 ± 252 TgC yr -1 sink for NA was driven primarily by CO 2 uptake in the Forest Lands sector (-248 TgC yr -1 ), largely in the Northwest and Southeast regions of the US, and in the Crop Lands sector (-297 TgC yr -1 ), predominantly in the Midwest US states. These sinks are counteracted by the carbon source estimated for the Other Lands sector (+218 TgC yr -1 ), where much of the forest and crop products are assumed to be returned to the atmosphere (through livestock and human consumption). The ecosystems of Mexico are estimated to be a small net source (+18 TgC yr -1 ) due to land use change between 1993 and 2002. We compare these inventory-based estimates with results from a suite of terrestrial biosphere and atmospheric inversion models, where the mean continental-scale NEE estimate for each ensemble is -511 TgC yr -1 and -931 TgC yr -1 , respectively. In the modeling approaches, all sectors, including Other Lands, were generally estimated to be a carbon sink, driven in part by assumed CO 2 fertilization and/or lack of consideration of carbon sources from disturbances and product emissions. Additional fluxes not measured by the inventories, although highly uncertain, could add an additional -239 TgC yr -1 to the inventory-based NA sink estimate, thus suggesting some convergence with the modeling approaches.
Widespread documentation of positive winter temperature anomalies, declining snowpack and earlier snow melt in the Northern Hemisphere have raised concerns about the consequences for regional water resources as well as wildfire. A topic that has not been addressed with respect to declining snowpack is effects on ecosystem water balance. Changes in water balance dynamics will be particularly pronounced at low elevations of mid-latitude dry regions because these areas will be the first to be affected by declining snow as a result of rising temperatures. As a model system, we used simulation experiments to investigate big sagebrush ecosystems that dominate a large fraction of the semiarid western United States. Our results suggest that effects on future ecosystem water balance will increase along a climatic gradient from dry, warm and snow-poor to wet, cold and snow-rich. Beyond a threshold within this climatic gradient, predicted consequences for vegetation switched from no change to increasing transpiration. Responses were sensitive to uncertainties in climatic prediction; particularly, a shift of precipitation to the colder season could reduce impacts of a warmer and snow-poorer future, depending on the degree to which ecosystem phenology tracks precipitation changes. Our results suggest that big sagebrush and other similar semiarid ecosystems could decrease in viability or disappear in dry to medium areas and likely increase only in the snow-richest areas, i.e. higher elevations and higher latitudes. Unlike cold locations at high elevations or in the arctic, ecosystems at low elevations respond in a different and complex way to future conditions because of opposing effects of increasing water-limitation and a longer snow-free season. Outcomes of such nonlinear interactions for future ecosystems will likely include changes in plant composition and productivity, dynamics of water balance, and availability of water resources.