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One of the most reliable features of natural systems is that they change through time. Theory predicts that temporally fluctuating conditions shape community composition, species distribution patterns, and life history variation, yet features of temporal variability are rarely incorporated into studies of species–environment associations. In this study, we evaluated how two components of temporal environmental variation—variability and predictability—impact plant community composition and species distribution patterns in the alpine tundra of the Southern Rocky Mountains in Colorado (USA). Using the Sensor Network Array at the Niwot Ridge Long‐Term Ecological Research site, we used in situ, high‐resolution temporal measurements of soil moisture and temperature from 13 locations ("nodes") distributed throughout an alpine catchment to characterize the annual mean, variability, and predictability in these variables in each of four consecutive years. We combined these data with annual vegetation surveys at each node to evaluate whether variability over short (within‐day) and seasonal (2‐ to 4‐month) timescales could predict patterns in plant community composition, species distributions, and species abundances better than models that considered average annual conditions alone. We found that metrics for variability and predictability in soil moisture and soil temperature, at both daily and seasonal timescales, improved our ability to explain spatial variation in alpine plant community composition. Daily variability in soil moisture and temperature, along with seasonal predictability in soil moisture, was particularly important in predicting community composition and species occurrences. These results indicate that the magnitude and patterns of fluctuations in soil moisture and temperature are important predictors of community composition and plant distribution patterns in alpine plant communities. More broadly, these results highlight that components of temporal change provide important niche axes that can partition species with different growth and life history strategies along environmental gradients in heterogeneous landscapes. [ABSTRACT FROM AUTHOR]

Abstract: A central goal at the interface of ecology and conservation is understanding how the relationship between biodiversity and ecosystem function (B–EF) will shift with changing climate. Despite recent theoretical advances, studies which examine temporal variation in the functional traits and mechanisms (mass ratio effects and niche complementarity effects) that underpin the B–EF relationship are lacking. Here, we use 13 years of data on plant species composition, plant traits, local‐scale abiotic variables, above‐ground net primary productivity (ANPP), and climate from the alpine tundra of Colorado (USA) to investigate temporal dynamics in the B–EF relationship. To assess how changing climatic conditions may alter the B–EF relationship, we built structural equation models (SEMs) for 11 traits across 13 years and evaluated the power of different trait SEMs to predict ANPP, as well as the relative contributions of mass ratio effects (community‐weighted mean trait values; CWM), niche complementarity effects (functional dispersion; FDis) and local abiotic variables. Additionally, we coupled linear mixed effects models with Multimodel inference methods to assess how inclusion of trait–climate interactions might improve our ability to predict ANPP through time. In every year, at least one SEM exhibited good fit, explaining between 19.6% and 57.2% of the variation in ANPP. However, the identity of the trait which best explained ANPP changed depending on winter precipitation, with leaf area, plant height and foliar nitrogen isotope content (δ15N) SEMs performing best in high, middle and low precipitation years, respectively. Regardless of trait identity, CWMs exerted a stronger influence on ANPP than FDis and total biotic effects were always greater than total abiotic effects. Multimodel inference reinforced the results of SEM analysis, with the inclusion of climate–trait interactions marginally improving our ability to predict ANPP through time. Synthesis. Our results suggest that temporal variation in climatic conditions influences which traits, mechanisms and abiotic variables were most responsible for driving the B–EF relationship. Importantly, our findings suggest that future research should consider temporal variability in the B–EF relationship, particularly how the predictive power of individual functional traits and abiotic variables may fluctuate as conditions shift due to climate change.

Premise: The soil microbiome plays a role in plant trait expression and fitness, and plants may be locally adapted or maladapted to their soil microbiota. However, few studies of local adaptation in plants have incorporated a microbial treatment separate from manipulations of the abiotic environment, so our understanding of microbes in plant adaptation is limited. Methods: Here we tested microbial effects on local adaptation in four paired populations of an abundant alpine plant from two community types, dry and moist meadow. In a 5‐month greenhouse experiment, we manipulated source population, soil moisture, and soil microbiome and measured plant survival and biomass to assess treatment effects. Results: Dry meadow populations had higher biomass than moist meadow populations at low moisture, demonstrating evidence of local adaptation to soil moisture in the absence of microbes. In the presence of microbes, dry meadow populations had greater survival than moist meadow populations when grown with dry meadow microbes regardless of moisture. Moist meadow populations showed no signs of adaptation or maladaptation. Conclusions: Our research highlights the importance of microbial mutualists in local adaptation, particularly in dry environments with higher abiotic stress. Plant populations from environments with greater abiotic stress exhibit different patterns of adaptation when grown with soil microbes versus without, while plant populations from less abiotically stressful environments do not. Improving our understanding of the role microbes play in plant adaptation will require further studies incorporating microbial manipulations. [ABSTRACT FROM AUTHOR]

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains (Lewisia pygmaea, Sibbaldia procumbens, and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum, Carex rupestris, and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species' performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure.

Accurately predicting upslope shifts in subalpine tree ranges with warming requires understanding how future forest populations will be affected by climate change, as these are the seed sources for new tree line and alpine populations. Early life history stages are particularly sensitive to climate and are also influenced by genetic variation among populations. We tested the climate sensitivity of germination and initial development for two widely distributed subalpine conifers, using controlled-environment growth chambers with one temperature regime from subalpine forest in the Colorado Rocky Mountains and one 5 °C warmer, and two soil moisture levels. We tracked germination rate and timing, rate of seedling development, and seedling morphology for two seed provenances separated by ~300 m elevation. Warming advanced germination timing and initial seedling development by a total of ~2 weeks, advances comparable to mean differences between provenances. Advances were similar for both provenances and species; however, warming reduced the overall germination rate, as did low soil moisture, only for Picea engelmannii. A three-year field warming and watering experiment planted with the same species and provenances yielded responses qualitatively consistent with the lab trials. Together these experiments indicate that in a warmer, drier climate, P. engelmannii germination, and thus regeneration, could decline, which could lead to declining subalpine forest populations, while Pinus flexilis forest populations could remain robust as a seed source for upslope range shifts.

Summary: Environmental change can affect species directly by altering their physical environment and indirectly by altering the abundance of interacting species. A key challenge at the interface of community ecology and conservation biology is to predict how direct and indirect effects combine to influence response in a changing environment. In particular, little is known about how direct and indirect effects on biodiversity develop over time or their potential to influence ecosystem function. We studied how nitrogen (N), winter precipitation (snow) and warming influenced diversity and ecosystem function over 6 years in alpine tundra. We used path analyses to partition direct effects of environmental manipulations from indirect effects due to changes in the abundance of two dominant plants. We hypothesize that (i) indirect effects will develop more slowly but will become stronger than direct effects over time and (ii) after 6 years, indirect effects will more strongly influence diversity while direct effects will influence ecosystem function. Indirect effects of N on diversity were consistently stronger than direct effects and actually developed quickly, prior to direct effects. Direct effects of snow on diversity were detected in year 2 but then subsequently were reversed, while indirect effects were detected in year 4 and grew stronger over time. Overall in year 6, indirect effects were much stronger than direct effects on diversity. Direct effects predominated for three of four ecosystem functions we measured (productivity, N mineralization, winter N availability). The only indirect effects we found were that N and snow indirectly affected microbial biomass N by influencing Geum abundance. Across all four ecosystem measures, indirect effects were infrequent and weaker than direct effects. Synthesis. Increasing indirect effects on diversity over time indicate that short-term experiments or monitoring of natural systems may underestimate the full magnitude of global change effects on plant communities. Moreover, explicitly accounting for changes in dominant plant abundance may be necessary for forecasting plant community response to environmental change. Conversely, weak indirect effects for ecosystem processes suggest that predicting ecosystem function without knowledge of plant responses to global change may be possible. Increasing indirect effects on diversity over time indicate that short-term experiments or monitoring of natural systems may underestimate the full magnitude of global change effects on plant communities. Explicitly accounting for changes in dominant plant abundance may be necessary for forecasting plant community response to environmental change. Conversely, weak indirect effects for ecosystem processes suggest that predicting ecosystem function without knowledge of plant responses to global change may be possible.

Hemiparasites are known to influence community structure and ecosystem functioning, but the underlying mechanisms are not well studied. Variation in the impacts of hemiparasites on diversity and production could be due to the difference in the relative strength of two interacting pathways: direct negative effects of parasitism and positive effects on N availability via litter. Strong effects of parasitism should result in substantial changes in diversity and declines in productivity. Conversely, strong litter effects should result in minor changes in diversity and increased productivity. We conducted field-based surveys to determine the association of Castilleja occidentalis with diversity and productivity in the alpine tundra. To examine litter effects, we compared the decomposition of Castilleja litter with litter of four other abundant plant species, and examined the decomposition of those four species when mixed with Castilleja. Castilleja was associated with minor changes in diversity but almost a twofold increase in productivity and greater foliar N in co-occurring species. Our decomposition trials suggest litter effects are due to both the rapid N loss of Castilleja litter and the effects of mixing Castilleja litter with co-occurring species. Castilleja produces litter that accelerates decomposition in the alpine tundra, which could accelerate the slow N cycle and boost productivity. We speculate that these positive effects of litter outweigh the effects of parasitism in nutrient-poor systems with long-lived hemiparasites. Determining the relative importance of parasitism and litter effects of this functional group is crucial to understand the strong but variable roles hemiparasites play in affecting community structure and ecosystem processes.

Traditional methods of carbon monitoring in mountainous regions are challenged by complex terrain. Recently, solar‐induced fluorescence (SIF) has been found to be an indicator of gross primary production (GPP), and the increased availability of remotely sensed SIF provides an opportunity to estimate GPP across the Western United States. Although the empirical linkage between SIF and GPP is strong, the current mechanistic understanding of this linkage is incomplete and depends upon changes in leaf biochemical processes in which absorbed sunlight leads to photochemistry, heat (via nonphotochemical quenching [NPQ]), fluorescence, or tissue damage. An improved mechanistic understanding is necessary to leverage SIF observations to improve representation of ecosystem processes within land surface models. Here we included an improved fluorescence model within the Community Land Model, Version 4.5 (CLM 4.5), to simulate seasonal changes in SIF at a subalpine forest in Colorado. We found that when the model accounted for sustained NPQ, this provided a larger seasonal change in fluorescence yield leading to simulated SIF that more closely resembled the observed seasonal pattern (Global Ozone Monitoring Experiment‐2 [GOME‐2] satellite platform and a tower‐mounted spectrometer system). We found that an acclimation model based on mean air temperature was a useful predictor for sustained NPQ. Although light intensity was not an important factor for this analysis, it should be considered before applying the sustained NPQ and SIF to other cold climate evergreen biomes. More leaf‐level fluorescence measurements are necessary to better understand the seasonal relationship between sustained and reversible components of NPQ and to what extent that influences SIF. Key Points: Sustained nonphotochemical quenching is the major sink for excess light for a high‐elevation conifer forest in winterA temperature‐based acclimation model was able to reproduce seasonal changes in sustained NPQIncluding a representation of sustained NPQ improved the simulation of SIF for cold‐climate evergreens within a land surface model [ABSTRACT FROM AUTHOR]

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains (Lewisia pygmaea, Sibbaldia procumbens, and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum, Carex rupestris, and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species’ performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure.

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains (Lewisia pygmaea , Sibbaldia procumbens , and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum , Carex rupestris , and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species’ performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure. [ABSTRACT FROM AUTHOR]

Climate change is affecting the hydrology of high-elevation mountain ecosystems, with implications for ecosystem functioning and water availability to downstream populations. We directly and continuously measured precipitation and evapotranspiration ( ET) from both subalpine forest and alpine tundra portions of a single catchment, as well as discharge fluxes at the catchment outlet, to quantify the water balance of a mountainous, headwater catchment in Colorado, USA. Between 2008 and 2012, the water balance closure averaged 90% annually, and the catchment ET was the largest water output at 66% of precipitation. Alpine ET was greatest during the winter, in part because of sublimation from blowing snow, which contributed from 27% to 48% of the alpine, and 6% to 9% of the catchment water balance, respectively. The subalpine ET peaked in summer. Alpine areas generated the majority of the catchment discharge, despite covering only 31% of the catchment area. Although the average annual alpine runoff efficiency (discharge/precipitation; 40%) was greater than the subalpine runoff efficiency (19%), the subalpine runoff efficiency was more sensitive to changes in precipitation. Inter-annual analysis of the evaporative and dryness indices revealed persistent moisture limitations at the catchment scale, although the alpine alternated between energy-limited and water-limited states in wet and dry years. Each ecosystem generally over-generated discharge relative to that expected from a Budyko-type model. The alpine and catchment water yields were relatively unaffected by annual meteorological variability, but this interpretation was dependent on the method used to quantify potential ET. Our results indicate that correctly accounting for dissimilar hydrological cycling above and below alpine treeline is critical to quantify the water balance of high-elevation mountain catchments over periods of meteorological variability. Copyright © 2015 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

Background: Snow and ice melt provide sensitive indicators of climate change and serve as the primary source of stream flow in alpine basins. Aims: We synthesise the results of hydrological and hydrochemical studies during the period 1995–2014, building on a long history of earlier work focused on snow and water on Niwot Ridge and the adjacent Green Lakes Valley (GLV), which is part of the Niwot Ridge Long Term Ecological Research site (NWT LTER). Methods: These studies are discussed in the context of how snow, snowmelt and runoff reflect changing local climate. We review recent results of snow, snowmelt, hydrology and hydrochemistry from the plot to the basin scale, utilising new tools such as continuous global positioning system (GPS) measurements of snow depth, along with remotely-sensed measurements of snow-covered area and melt, combined with long-term measurements of snow properties, discharge and solute and isotopic content of water. Results andConclusions: Surface–groundwater interactions are important components of water quantity and quality in alpine basins. Some or most snowmelt infiltrates underlying soils and bedrock, transporting soil and bedrock products to surface waters. Infiltrating snowmelt, along with increased melt of stored ice, increases the hydrologic connectivity between the terrestrial and aquatic systems. Alpine basins are being impacted by increases in atmospheric nitrogen deposition, which has caused changes in soil microbial processes such as nitrification. Nitrate, dissolved organic carbon and dissolved organic nitrogen are thus flushed from soils and talus to streams. Our long-term results show that alpine catchments, such as Green Lake 4 at NWT LTER+, have the greatest sensitivity and least resilience to climate warming, with any warming leading to increased water yields. [ABSTRACT FROM PUBLISHER]

Background:Current understanding of treeline or forest-alpine ecotone (FAE) dynamics does not fully explain past and present FAE patterns and their underlying processes, nor allow prediction of their response to climate change. Aims:We address the overarching hypothesis that the FAE is a mosaic of distinct landscape units of vegetation and landforms that result in differential responses to climate change. We focus on climate-related, landscape and vegetation characteristics, but also consider the effect of landscape heterogeneity on biogeochemistry and overall resilience of the FAE to climate change. Results: There are three distinct FAE land units at Niwot Ridge, generated by different interactions of climate with vegetation, landforms and topography. Within these FAEs, a process of self-organisation takes place from organism to patch to landscape scales, and is modulated by positive and negative feedback loops along an elevation gradient. The underlying controls cannot be attributed solely to temperature, but to a combination of interactions along a physical/biotic gradient. Conclusions:FAE dynamics result from interactions among mechanisms and processes at the microsite, patch and landscape scales: (1) tree persistence; (2) forest patch establishment; (3) drivers of patch forest configurations and (4) resilience, increasing along a gradient of biotic control. [ABSTRACT FROM PUBLISHER]

Background:There is a paucity of information for mountain catchments, and in particular a lack of long-term data collection in alpine areas. Aims:The focus of this special issue is to synthesize alpine research undertaken in the last 20 years at high altitude research sites that comprise the Niwot Ridge Long Term Ecological Research (NWT LTER) programme. It is a timely update of the benchmark volume on alpine ecology by Bowman and Seastedt (2001) that presented a summary of work carried out over 40 years on the structure and functioning of the NWT LTER alpine terrestrial and aquatic ecosystems. Methods:We introduce and synthesize both the individual manuscripts in this monograph, along with insights from non-LTER investigators. Results and conclusions:Mountain areas are sentinels of climate change. We are seeing those effects today. Furthermore, these ecosystem changes we report are occurring in mountain areas before they occur in downstream ecosystems. Thus, the sensitivity of mountain ecosystems to changes in climate begs for enhanced and worldwide protection. Our results suggest that investing in observational and integrated social-ecological system research in mountains is key for underpinning scientifically the development and implementation of efficient mitigation and adaptation measures and sustainable development strategies. [ABSTRACT FROM PUBLISHER]

Background: Subsurface ice preserved as ice lenses and within rock glaciers as well as glacial and lake ice provides sensitive indicators of climate change and serve as a late-season source of meltwater. Aims: We synthesise the results of geomorphological, geophysical and geochemical studies during the period of 1995–2014, building on a long history of earlier work focused on ice and permafrost studies on Niwot Ridge and the adjacent Green Lakes Valley (GLV), which is part of the Niwot Ridge Long Term Ecological Research Site. Methods: These studies are discussed in the context of how bodies of ice and rock glaciers reflect changing local climate. We review recent results from geophysical investigations (resistivity, seismic refraction and ground-penetrating radar) of the shallow subsurface, ongoing monitoring of the Arikaree Glacier, three rock glaciers and lake ice in the GLV, and interpretations of how subsurface ice melt regulates the flow and chemistry of alpine surface water after seasonal snowfields melt. Results and conclusions: Permafrost conditions reported from Niwot Ridge in the 1970s are generally absent today, but ice lenses form and melt seasonally. Ice is present permanently within the Green Lakes 5 rock glacier and at nearby favourable sites. The Arikaree Glacier has shown a marked decline in cumulative mass balance during the past 12 years after a 30-year period when net mass balance was ca. 0. Duration of seasonal lake ice increases with elevation in GLV, but duration has decreased at all seven lakes that have been monitored during the last three decades. This decrease has been most marked at the lowest elevation where it amounted to a reduction of about 1 d year−1and least at Green Lake 5 where the loss has been at a rate of 0.5 d year−1. Surface temperature measurements from rock glaciers have not shown strong trends during the past 15 years. It has been suggested that almost all of the 2.5-mm year−1increase in stream discharge from the upper GLV in September and October has been derived from melting of subsurface ice. [ABSTRACT FROM PUBLISHER]

Background: Animals that hoard food to mediate seasonal deficits in resource availability might be particularly vulnerable to climate-mediated reductions in the quality and accessibility of food during the caching season. Central-place foragers might be additionally impacted by climatic constraints on their already restricted foraging range. Aims: We sought evidence for these patterns in a study of the American pika (Ochotona princeps), a territorial, central-place forager sensitive to climate. Methods: Pika food caches and available forage were re-sampled using historical methods at two long-term study sites, to quantify changes over two decades. Taxa that changed in availability or use were analysed for primary and secondary metabolites. Results: Both sites trended towards warmer summers, and snowmelt trended earlier at the lower latitude site. Graminoid cover increased at each site, and caching trends appeared to reflect available forage rather than primary metabolites. Pikas at the lower latitude site preferred species higher in secondary metabolites, known to provide higher-nutrient winter forage. However, caching of lower-nutrient graminoids increased in proportion with graminoid availability at that site. Conclusions: If our results represent trends in climate, cache quality and available forage, we predict that pikas at the lower latitude site will soon face nutritional deficiencies. [ABSTRACT FROM PUBLISHER]

Background:Surface waters are the lowest points in the landscape, and therefore serve as excellent integrators and indicators of changes taking place in the surrounding terrestrial and atmospheric environment. Aims:Here we synthesise the findings of limnological studies conducted during the past 15 years in streams and lakes in the Green Lakes Valley, which is part of the Niwot Ridge Long Term Ecological Research (LTER) Site. Methods:The importance of these studies is discussed in the context of aquatic ecosystems as indicators, integrators, and regulators of environmental change. Specifically, investigations into climatic, hydrologic, and nutrient controls on present-day phytoplankton, and historical diatom, community composition in the alpine lake, Green Lake 4, are reviewed. In addition, studies of spatial and temporal patterns in dissolved organic matter (DOM) biogeochemistry and reactive transport modelling that have taken place in the Green Lakes Valley are highlighted. Results and conclusions:The findings of these studies identify specific shifts in algal community composition and DOM biogeochemistry that are indicative of changing environmental conditions and provide a framework for detecting future environmental change in the Green Lakes Valley and in other alpine watersheds. Moreover, the studies summarised here demonstrate the importance of long-term monitoring programmes such as the LTER programme. [ABSTRACT FROM PUBLISHER]

Premise of the study: Nitrogen (N) inputs to the terrestrial environment have doubled worldwide during the past century. N negatively impacts plant diversity, but it is unknown why some species are more susceptible than others. While it is often assumed that competition drives species decline, N enrichment also strongly affects soil microbial communities. Can these changes affect plant-microbe interactions in ways that differentially influence success of plant species? Furthermore, can altered plant-microbe interactions lead to carbon (C) limitation in plants? Methods: We focused on a species that increases (Deschampsia cespitosa) and one that decreases (Geum rossii) in abundance in N-fertilized plots in alpine tundra. We measured soil microbes using phospholipid fatty acids, and C limitation and transfer using a 13C tracer experiment, C:N ratios, nonstructural carbohydrates, and leaf preformation. Key results: While N profoundly influenced microbial communities, this change occurred similarly in association with both plant species. N addition did not alter total C allocation to microbes in either species, but it changed patterns of microbial C acquisition more in Geum, specifically in gram-negative bacteria. Geum showed evidence of C limitation: it allocated less C to storage organs, had lower C : N and carbohydrate stores, and fewer preformed leaves in N plots. Conclusions: Carbon limitation may explain why some species decline with N enrichment, and the decline may be due to physiological responses of plants to N rather than to altered plant-microbe interactions. Global change will alter many processes important in structuring plant communities: noncompetitive mechanisms of species decline may be more widespread than previously thought. [ABSTRACT FROM AUTHOR]

Snow is a critical storage component in the hydrologic cycle, but current measurement networks are sparse. In addition, the heterogeneity of snow requires surveying larger areas to measure the areal average. We presented snow measurements using GPS interferometric reflectometry (GPS-IR). GPS-IR measures a large area (~100 m2), and existing GPS installations around the world have the potential to expand existing snow measurement networks. GPS-IR uses a standard, geodetic GPS installation to measure the snow surface via the reflected component of the signal. We reported GPS-IR snow depth measurements made at Niwot Ridge, Colorado, from October 2009 through June 2010. This site is in a topographic saddle at 3500 m elevation with a peak snow depth of 1.7 m near the GPS antenna. GPS-IR measurements are compared with biweekly snow surveys, a continuously operating scanning laser system and an airborne light detection and ranging (LIDAR) measurement. The GPS-IR measurement of peak snowpack (1.36-1.76 m) matches manual measurements (0.95-1.7 m) and the scanning laser (1.16 m). GPS-IR has RMS error of 13 cm (bias = 10 cm) compared with the laser, although differences between the measurement locations make comparison imprecise. Over the melt season, when the snowpack is more homogenous, the difference between the GPS-IR and the laser is reduced (RMS = 9 cm, bias = 6 cm). In other locations, the GPS and the LIDAR agree on which areas have more or less snow, but the GPS estimates more snow on the ground on tracks to the west (1.58 m) than the LIDAR (1.14 m). Copyright © 2011 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

1. Many studies of community assembly focus on two mechanisms: environmental filtering and competitive interactions. This focus ignores the importance of other assembly processes such as equalizing fitness processes and facilitation. The contribution of different processes to community assembly can be elucidated by examining functional diversity patterns of traits that differ in their contribution to different assembly processes. 2. In alpine tundra, we explored trait patterns along a stress-resource gradient that varied in productivity, nitrogen availability and soil moisture. We explore whether functional diversity is low in abiotic stressful environments and increases in more benign environments as competition becomes more important, and if equalizing fitness processes and facilitation affect functional diversity. We calculated community-weighted mean trait values and functional diversity for specific leaf area (SLA), leaf area, stomatal conductance, plant height and chlorophyll content as well as multivariate functional diversity and phylogenetic diversity. 3. At the community level, functional diversity increased at both ends of the gradient: high resource availability was associated with greater functional diversity in height and leaf area, and lower resource availability was associated with greater functional diversity in SLA, stomatal conductance, and chlorophyll content. As a result of this trade-off in functional diversity among traits, multivariate functional diversity did not change across the gradient. Phylogenetic diversity increased with increasing resource availability. 4. We find evidence for at least three assembly processes along the gradient. Abiotic filtering by wind and cold exposure may reduce functional diversity in height and leaf area at the low resource end of the gradient. Also at low resource availability, increasing functional diversity in the other three traits suggests competition for below-ground resources. At the resource-rich end of the gradient, increased functional diversity in height and leaf area suggests increased competition for light or facilitation. 5. Synthesis: Our results suggest that multiple assembly processes (abiotic filtering, above-ground competition, and below-ground competition) operate simultaneously to structure plant communities along a stress-resource gradient. These processes would be obscured by a single multivariate trait index or phylogenetic diversity and are only evident by analysing functional diversity patterns of individual traits. [ABSTRACT FROM AUTHOR]

Knowledge of import, export, and transport of nitrogen (N) in headwater catchments is essential for understanding ecosystem function and water quality in mountain ecosystems, especially as these ecosystems experience increased anthropogenic N deposition. In this study, we link spatially explicit soil and stream data at the landscape scale to investigate import, export and transport of N in a 0.89 km site at the alpine-subalpine ecotone in the Front Range of the Rocky Mountains, Colorado, U.S.A. For two of the major N inputs to our site, N deposition in the snowpack and N fixation, a complementary relationship was found across the study site, with greater abundance of N-fixing plants in areas with less snow and substantial snow inputs in areas with low N fixer abundance. During the initial phases of snowmelt, mixing model end members for oxygen isotopes in nitrate (NO) indicated that a substantial quantity of NO is transported downhill into the forested subalpine without being assimilated by soil microbes. After this initial pulse, much less NO entered the stream and most but not all of it was microbial in origin. Rising δN in stream NO indicated greater influence of fractionating processes such as denitrification later in the season. NO from both atmospheric and microbial sources was not exported from our site because it was consumed within the first several hundred meters of the stream; ultimately, N exports were in the form of dissolved organic nitrogen (DON) and particulate N (PN). The results of this study suggest that the highest elevation dry alpine meadows rely more heavily on N fixation as an N source and experience less of the effects of anthropogenic N deposition than mid and lower elevation areas that have more snow. Our data also suggest that mid-elevation krummholz, moist meadows, and talus slopes are exporting N as NO shortly after the onset of snowmelt, but that this NO is rapidly consumed as the stream flows through the subalpine forest. This consumption by assimilation and/or denitrification currently provides a buffer against increased inorganic N availability downstream. [ABSTRACT FROM AUTHOR]

Climate change is predicted to cause upward shifts in forest tree distributions, which will require seedling recruitment beyond current forest boundaries. However, predicting the likelihood of successful plant establishment beyond current species' ranges under changing climate is complicated by the interaction of genetic and environmental controls on seedling establishment. To determine how genetics and climate may interact to affect seedling establishment, we transplanted recently germinated seedlings from high- and low-elevation provenances (HI and LO, respectively) of Pinus flexilis in common gardens arrayed along an elevation and canopy gradient from subalpine forest into the alpine zone and examined differences in physiology and morphology between provenances and among sites. Plant dry mass, projected leaf area and shoot:root ratios were 12–40% greater in LO compared with HI seedlings at each elevation. There were no significant changes in these variables among sites except for decreased dry mass of LO seedlings in the alpine site. Photosynthesis, carbon balance (photosynthesis/respiration) and conductance increased >2× with elevation for both provenances, and were 35–77% greater in LO seedlings compared with HI seedlings. There were no differences in dark-adapted chlorophyll fluorescence (Fv/Fm) among sites or between provenances. Our results suggest that for P. flexilis seedlings, provenances selected for above-ground growth may outperform those selected for stress resistance in the absence of harsh climatic conditions, even well above the species' range limits in the alpine zone. This indicates that forest genetics may be important to understanding and managing species' range adjustments due to climate change. [ABSTRACT FROM AUTHOR]

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource,. and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season. and then used `15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche. (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist. [ABSTRACT FROM AUTHOR]

Eddy covariance measurements of the surface energy balance and carbon dioxide exchange above high-elevation (3,480 m above sea level) alpine tundra located near Niwot Ridge, Colorado, were compared to simultaneous measurements made over an adjacent subalpine forest over two summers and one winter, from June 9, 2007 to July 3, 2008. The surface energy balance closure at the alpine site averaged 71 and 91%, winter and summer, respectively, due to the high wind speeds, short turbulent flux footprint, and relatively flat ridge-top location of the measurement site. Throughout the year, the alpine site was cooler with higher relative humidity, and had a higher horizontal wind speed, especially in winter, compared to the forest site. Wind direction was persistently downslope at the alpine site (summer and winter, day and night), whereas upslope winds were common at the forest site during summer daytime periods. The latent and sensible heat fluxes were consistently larger in magnitude at the forest site, with the largest differences during summer. The horizontal advective flux of CO2 at the alpine site averaged 6% of the net ecosystem exchange ( NEE) during summer nights (5% during summer daytime), and was small in relation to the high wind speeds, relatively flat site, and weak sources of CO2 upwind of the site. The magnitudes and diurnal behavior of the alpine NEE calculated using three methods; eddy-covariance, friction velocity filter, and with advection and storage calculations, gave similar results. The period of net CO2 uptake (negative NEE) was 100 days at the alpine site with a net uptake of 16 g C m−2, compared to 208 days at the forest site with a net uptake of 108 g C m−2, with initiation of net uptake coinciding with air temperatures reaching +10°C. Winter respiration loss at the alpine site was 164 g C m−2 over 271 days, compared to 52 g C m−2 over 175 days at the forest site, with the initiation of net loss coinciding with air temperatures reaching −10°C at each site. [ABSTRACT FROM AUTHOR]

Abstract: Modeling how the role of forests in the carbon cycle will respond to predicted changes in water availability hinges on an understanding of the processes controlling water use in ecosystems. Recent studies in forest ecosystem modeling have employed data-assimilation techniques to generate parameter sets that conform to observations, and predict net ecosystem CO2 exchange (NEE) and its component processes. Since the carbon and water cycles are linked, there should be additional process information available from ecosystem H2O exchange. We coupled SIPNET (Simple Photosynthesis EvapoTranspiration), a simplified model of ecosystem function, with a data-assimilation system to estimate parameters leading to model predictions most closely matching the net CO2 and H2O fluxes measured by eddy covariance in a high-elevation, subalpine forest ecosystem. When optimized using measurements of CO2 exchange, the model matched observed NEE (RMSE=0.49gCm−2) but underestimated transpiration calculated independently from sap flow measurements by a factor of 4. Consequently, the carbon-only optimization was insensitive to imposed changes in water availability. Including eddy flux data from both CO2 and H2O exchange to the optimization reduced the model fit to the observed NEE fluxes only slightly (RME=0.53gCm−2), however this parameterization also reproduced transpiration calculated from independent sap flow measurements (r 2 =0.67, slope=0.6). A significant amount of information can be extracted from simultaneous analysis of CO2 and H2O exchange, which improved the accuracy of transpiration estimates from measured evapotranspiration. Conversely, failure to include both CO2 and H2O data streams can generate results that mask the responses of ecosystem carbon cycling to variation in the precipitation. In applying the model conditioned on both CO2 and H2O fluxes to the subalpine forest at the Niwot Ridge AmeriFlux site, we observed that the onset of transpiration is coincident with warm soil temperatures. However, after snow has covered the ground in the fall, we observed significant inter-annual variability in the fraction of evapotranspiration composed of transpiration; evapotranspiration was dominated by transpiration in years when late fall air temperatures were high enough to maintain photosynthesis, but by sublimation from the surface of the snowpack in years when late fall air temperatures were colder and forest photosynthetic activity had ceased. Data-assimilation techniques and simultaneous measurements of carbon and water exchange can be used to quantify the response of net carbon uptake to changes in water availability by using an ecosystem model where the carbon and water cycles are linked. [Copyright &y& Elsevier]

Repercussions of species loss on ecosystem processes depend on the effects of the lost species as well as the compensatory responses of the remaining species in the community. We experimentally removed two co-dominant plant species and added a 15N tracer in alpine tundra to compare how species’ functional differences influence community structure and N cycling. For both of the species, production compensated for the biomass removed by the second year. However, the responses of the remaining species depended on which species was removed. These differences in compensation influenced how species loss impacted ecosystem processes. After the removal of one of the co-dominant species, Acomastylis rossii, there were few changes in the relative abundance of the remaining species, and differences in functioning could be predicted based on effects associated with the removed species. In contrast, the removal of the other co-dominant, Deschampsia caespitosa, was associated with subsequent changes in community structure (species relative abundances and diversity) and impacts on ecosystem properties (microbial biomass N, dissolved organic N, and N uptake of subordinate species). Variation in compensation may contribute to the resulting effects on ecosystem functioning, with the potential to buffer or accelerate the effects of species loss. [ABSTRACT FROM AUTHOR]