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  1. Abstract

    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.

     
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    Free, publicly-accessible full text available October 26, 2025
  2. Abstract

    Terrestrial, aquatic, and marine ecosystems regulate climate at local to global scales through exchanges of energy and matter with the atmosphere and assist with climate change mitigation through nature‐based climate solutions. Climate science is no longer a study of the physics of the atmosphere and oceans, but also the ecology of the biosphere. This is the promise of Earth system science: to transcend academic disciplines to enable study of the interacting physics, chemistry, and biology of the planet. However, long‐standing tension in protecting, restoring, and managing forest ecosystems to purposely improve climate evidences the difficulties of interdisciplinary science. For four centuries, forest management for climate betterment was argued, legislated, and ultimately dismissed, when nineteenth century atmospheric scientists narrowly defined climate science to the exclusion of ecology. Today's Earth system science, with its roots in global models of climate, unfolds in similar ways to the past. With Earth system models, geoscientists are again defining the ecology of the Earth system. Here we reframe Earth system science so that the biosphere and its ecology are equally integrated with the fluid Earth to enable Earth system prediction for planetary stewardship. Central to this is the need to overcome an intellectual heritage to the models that elevates geoscience and marginalizes ecology and local land knowledge. The call for kilometer‐scale atmospheric and ocean models, without concomitant scientific and computational investment in the land and biosphere, perpetuates the geophysical view of Earth and will not fully provide the comprehensive actionable information needed for a changing climate.

     
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    Free, publicly-accessible full text available August 22, 2025
  3. Abstract Aim

    The spectral variability hypothesis (SVH) predicts that spectral diversity, defined as the variability of radiation reflected from vegetation, increases with biodiversity. While confirmation of this hypothesis would pave the path for use of remote sensing to monitor biodiversity, support in herbaceous ecosystems is mixed. Methodological aspects related to scale have been the predominant explanation for the mixed support, yet biological characteristics that vary among herbaceous systems may also affect the strength of the relationship. Therefore, we examined the influence of three biological characteristics on the relationship between spectral and taxonomic diversity: vegetation density, spatial species turnover and invasion by non‐native species. We aimed to understand when and why spectral diversity may serve as an indicator of taxonomic diversity and be useful for monitoring.

    Location

    Continental U.S.A.

    Time Period

    Peak greenness in 2017.

    Major Taxa Studied

    Grassland and herbaceous ecosystems.

    Methods

    For nine herbaceous sites in the National Ecological Observatory Network, we calculated taxonomic diversity from field surveys of 20 m × 20 m plots and derived spectral diversity for those same plots from airborne hyperspectral imagery with a spatial resolution of 1 m. The strength of the taxonomic diversity–spectral diversity relationship at each site was subsequently assessed against measurements of vegetation density, spatial species turnover and invasion.

    Results

    We found a significant relationship between taxonomic and spectral diversity at some, but not all, sites. Spectral diversity was more strongly related to taxonomic diversity in sites with high species turnover and low invasion, but vegetation density had no effect on the relationship.

    Main Conclusions

    Using spectral diversity as a proxy for taxonomic diversity in grasslands is possible in some circumstances but should not just be assumed based on the SVH. It is important to understand the biological characteristics of a community prior to considering spectral diversity to monitor taxonomic diversity.

     
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  4. ABSTRACT

    Forecasting plant responses under global change is a critical but challenging endeavour. Despite seemingly idiosyncratic responses of species to global change, greater generalisation of ‘winners’ and ‘losers’ may emerge from considering how species functional traits influence responses and how these responses scale to the community level. Here, we synthesised six long‐term global change experiments combined with locally measured functional traits. We quantified the change in abundance and probability of establishment through time for 70 alpine plant species and then assessed if leaf and stature traits were predictive of species and community responses across nitrogen addition, snow addition and warming treatments. Overall, we found that plants with more resource‐acquisitive trait strategies increased in abundance but each global change factor was related to different functional strategies. Nitrogen addition favoured species with lower leaf nitrogen, snow addition favoured species with cheaply constructed leaves and warming showed few consistent trends. Community‐weighted mean changes in trait values in response to nitrogen addition, snow addition and warming were often different from species‐specific trait effects on abundance and establishment, reflecting in part the responses and traits of dominant species. Together, these results highlight that the effects of traits can differ by scale and response of interest.

     
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  5. Abstract

    Climate change is altering interactions among plants and pollinators. In alpine ecosystems, where snowmelt timing is a key driver of phenology, earlier snowmelt may generate shifts in plant and pollinator phenology that vary across the landscape, potentially disrupting interactions. Here we ask how experimental advancement of snowmelt timing in a topographically heterogeneous alpine-subalpine landscape impacts flowering, insect pollinator visitation, and pathways connecting key predictors of plant-pollinator interaction. Snowmelt was advanced by an average of 13.5 days in three sites via the application of black sand over snow in manipulated plots, which were paired with control plots. For each forb species, we documented flowering onset and counted flowers throughout the season. We also performed pollinator observations to measure visitation rates. The majority (79.3%) of flower visits were made by dipteran insects. We found that plants flowered earlier in advanced snowmelt plots, with the largest advances in later-flowering species, but flowering duration and visitation rate did not differ between advanced snowmelt and control plots. Using piecewise structural equation models, we assessed the interactive effects of topography on snowmelt timing, flowering phenology, floral abundance, and pollinator visitation. We found that these factors interacted to predict visitation rate in control plots. However, in plots with experimentally advanced snowmelt, none of these predictors explained a significant amount of variation in visitation rate, indicating that different predictors are needed to understand the processes that directly influence pollinator visitation to flowers under future climate conditions. Our findings demonstrate that climate change-induced early snowmelt may fundamentally disrupt the predictive relationships among abiotic and biotic drivers of plant-pollinator interactions in subalpine-alpine environments.

     
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  6. Abstract

    Patterns of alpine plant productivity are extremely variable in space and time. Complex topography drives variations in temperature, wind, and solar radiation. Surface and subsurface flow paths route water between landscape patches. Redistribution of snow creates scour zones and deep drifts, which drives variation in water availability and growing season length. Hence, the distribution of snow likely plays a central role in patterns of alpine plant productivity. Given that these processes operate at sub‐1 m to sub‐10 m spatial scales and are dynamic across daily to weekly time scales, historical studies using manual survey techniques have not afforded a comprehensive assessment of the influence of snow distribution on plant productivity. To address this knowledge gap, we used weekly estimates of normalised difference vegetation index (NDVI), snow extent, and peak snow depth, acquired from drone surveys at 25 cm resolution. We derived six snowpack‐related and topographic variables that may influence vegetation productivity and analysed these with respect to the timing and magnitude of peak productivity. Peak NDVI and peak NDVI timing were most highly correlated with maximum snow depth, and snow‐off‐date. We observed up to a ~30% reduction in peak NDVI for areas with deeper and later snow cover, and a ~11‐day delay in the timing of peak NDVI in association with later snow‐off‐date. Our findings leverage a novel approach to quantify the importance of snow distribution in driving alpine vegetation productivity and provide a space for time proxy of potential changes in a warmer, lower snow future.

     
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  7. Abstract

    Nutrient limitation is widespread in terrestrial ecosystems. Accordingly, representations of nitrogen (N) limitation in land models typically dampen rates of terrestrial carbon (C) accrual, compared with C‐only simulations. These previous findings, however, rely on soil biogeochemical models that implicitly represent microbial activity and physiology. Here we present results from a biogeochemical model testbed that allows us to investigate how an explicit versus implicit representation of soil microbial activity, as represented in the MIcrobial‐MIneral Carbon Stabilization (MIMICS) and Carnegie‐Ames‐Stanford Approach (CASA) soil biogeochemical models, respectively, influence plant productivity, and terrestrial C and N fluxes at initialization and over the historical period. When forced with common boundary conditions, larger soil C pools simulated by the MIMICS model reflect longer inferred soil organic matter (SOM) turnover times than those simulated by CASA. At steady state, terrestrial ecosystems experience greater N limitation when using the MIMICS‐CN model, which also increases the inferred SOM turnover time. Over the historical period, however, warming‐induced acceleration of SOM decomposition over high latitude ecosystems increases rates of N mineralization in MIMICS‐CN. This reduces N limitation and results in faster rates of vegetation C accrual. Moreover, as SOM stoichiometry is an emergent property of MIMICS‐CN, we highlight opportunities to deepen understanding of sources of persistent SOM and explore its potential sensitivity to environmental change. Our findings underscore the need to improve understanding and representation of plant and microbial resource allocation and competition in land models that represent coupled biogeochemical cycles under global change scenarios.

     
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  8. Abstract

    Increased plant growth under elevated carbon dioxide (CO2) slows the pace of climate warming and underlies projections of terrestrial carbon (C) and climate dynamics. However, this important ecosystem service may be diminished by concurrent changes to vegetation carbon‐to‐nitrogen (C:N) ratios. Despite clear observational evidence of increasing foliar C:N under elevated CO2, our understanding of potential ecological consequences of foliar stoichiometric flexibility is incomplete. Here, we illustrate that when we incorporated CO2‐driven increases in foliar stoichiometry into the Community Land Model the projected land C sink decreased two‐fold by the end of the century compared to simulations with fixed foliar chemistry. Further, CO2‐driven increases in foliar C:N profoundly altered Earth's hydrologic cycle, reducing evapotranspiration and increasing runoff, and reduced belowground N cycling rates. These findings underscore the urgency of further research to examine both the direct and indirect effects of changing foliar stoichiometry on soil N cycling and plant productivity.

     
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  9. Abstract

    Alpine tundra ecosystems are highly vulnerable to climate warming but are governed by local‐scale abiotic heterogeneity, which makes it difficult to predict tundra responses to environmental change. Although land models are typically implemented at global scales, they can be applied at local scales to address process‐based ecological questions. In this study, we ran ecosystem‐scale Community Land Model (CLM) simulations with a novel hillslope hydrology configuration to represent topographically heterogeneous alpine tundra vegetation across a moisture gradient at Niwot Ridge, Colorado, USA. We used local observations to evaluate our simulations and investigated the role of topography and aspect in mediating patterns of snow, productivity, soil moisture, and soil temperature, as well as the potential exposure to climate change across an alpine tundra hillslope. Overall, our simulations captured observed gradients in abiotic conditions and productivity among heterogeneous, hydrologically connected vegetation communities (moist, wet, and dry). We found that south facing aspects were characterized by reduced snowpack and drier and warmer soils in all communities. When we extended our simulations to the year 2100, we found that earlier snowmelt altered the timing of runoff, with cascading effects on soil moisture, productivity, and growing season length. However, these effects were not distributed equally across the tundra, highlighting potential vulnerabilities of alpine vegetation in dry, wind‐scoured, and south facing areas. Overall, our results demonstrate how land model outputs can be applied to advance process‐based understanding of climate change impacts on ecosystem function.

     
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  10. Abstract

    Fine‐scale microclimate variation due to complex topography can shape both current vegetation distributional patterns and how vegetation responds to changing climate. Topographic heterogeneity in mountains is hypothesized to mediate responses to regional climate change at the scale of metres. For alpine vegetation especially, the interplay between changing temperatures and topographically mediated variation in snow accumulation will determine the overall impact of climate change on vegetation dynamics.

    We combined 30 years of co‐located measurements of temperature, snow and alpine plant community composition in Colorado, USA, to investigate vegetation community trajectories across a snow depth gradient.

    Our analysis of long‐term trends in plant community composition revealed notable directional change in the alpine vegetation with warming temperatures. Furthermore, community trajectories are divergent across the snow depth gradient, with exposed parts of the landscape that experience little snow accumulation shifting towards stress‐tolerant, cold‐ and drought‐adapted communities, while snowier areas shifted towards more warm‐adapted communities.

    Synthesis: Our findings demonstrate that fine‐scale topography can mediate both the magnitude and direction of vegetation responses to climate change. We documented notable shifts in plant community composition over a 30‐year period even though alpine vegetation is known for slow dynamics that often lag behind environmental change. These results suggest that the processes driving alpine plant population and community dynamics at this site are strong and highly heterogeneous across the complex topography that is characteristic of high‐elevation mountain systems.

     
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