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Anthropogenic climate change is altering interactions among numerous species, including plants and pollinators. Plant-pollinator interactions, crucial for the persistence of most plant and many insect species, are threatened by climate change-driven phenological shifts. Phenological mismatches between plants and their pollinators may affect pollination services, and simulations indicated that these mismatches may reduce floral resources available to up to 50% of insect pollinator species. Although alpine plants rely heavily on vegetative reproduction, seedling recruitment and seed dispersal are likely to be important drivers of alpine community structure. Similarly, advanced flowering may expose plants to increased risk of frost damage and shifted soil moisture regimes; phenologically advanced plants will experience these environmental factors differently, which may alter their floral resource production. These effects may be dependent upon topography. Some species of alpine plants on the Niwot Ridge have displayed advanced phenology under treatments of advanced snowmelt (Forrester, 2021). However, little is understood about how these differences in distribution and phenology affect pollinator community composition and plant fecundity. Here we strive to examine how experimentally-induced changes in the timing of flowering and number of flowers produced by plants impact plant-pollinator interactions and seed set. We also ask how topography and the number of flowers interact with early snowmelt to affect pollination rates and the diversity of pollinating insects. Finally, we ask how seed set of Geum rossii is affected by pollinator visitation at different times of the season, under experimentally advanced snowmelt versus unmanipulated snowmelt, and with visitation by different insect taxa. In summer 2020, we found that plots with advanced phenology experienced peaks in pollinator visitation rates and pollinator diversity earlier than plots with unmanipulated snowmelt. We expect this to be because of the advanced floral phenology of certain key species in these plots. References: Forrester, C.C. (2021). Advancing, Using, and Teaching Climate Change Ecology Research. [Doctoral dissertation, University of Colorado, Boulder]. ProQuest Dissertations and Theses. 

As a result of climate change, the Rocky Mountain Front Range is experiencing warmer summers and earlier snowmelt. Due to the importance of snow for regulating soil temperature, growing season length, and available moisture in alpine ecosystems, even small shifts in the snow-free period could have large impacts. The focus of the Black Sand Extended Growing Season Length Experiment is to examine how terrain-related differences in climate exposure influence the way alpine habitats respond to climate change via earlier snowmelt. To simulate how climate exposure may affect plant communities, NWT LTER researchers established 5 experimental sites each containing a pair 10 x 40m rectangular plots. These sites include north and south facing aspects, subalpine and alpine tundra meadows in a range of hydrological conditions (e.g. dry meadows, moist meadows, wet meadows). We accelerated snowmelt in one plot of each block by adding chemically inert black sand, while keeping the second plot as an unmanipulated control; black sand was added to these plots after snow had naturally melted. This dataset includes geolocations of individual subplots and sensors within the experiment, measured in summer 2023. 

In 2006 we established a global change experiment in the Front Range of the Rocky Mountains to investigate how manipulations of warmer summer temperature, N deposition, and increased snowpack would affect the growth of alpine plants. The experiment was implemented on Niwot Ridge, where shrub cover has expanded by over 400% since 1946 (Formica et al. 2014). We established experimental plots north of the Niwot Ridge saddle, in an area of moist meadow tundra where willow shrub (Salix sp.) patches are present. Within experimental plots, Salix glauca seedlings were transplanted in 2006 and 2007 to test whether changing environmental conditions facilitated shrub survival and growth. In 2007 and 2008, phenological observations were recorded for all (2007) or abundant (2008) species in experimental plots. Measurements of plant species composition and aboveground net primary productivity (ANPP) are also made annually or biennially (ANPP, 2017-onward). In 2021, canopy height and NDVI began being measured annually. In July of 2016, a community transplant experiment was implemented to test whether changing environmental conditions support changes in alpine tundra plant communities. Two species characteristic of (1) dry meadow tundra (Tetraneuris acaulis, Erigeron pinnatisectus), (2) snowbed tundra (Ranunculus adoneus, Saxifraga rhomboidea) and (3) subalpine meadow (Trollius albiflorus, Polemonium pulcherrimum) were transplanted into experimental plots. Survival and growth of transplants was documented annually through 2021. 

Abstract While most studies of species coexistence focus on the mechanisms that maintain coexistence, it is equally important to understand the mechanisms that structure failed coexistence. For example, California annual grasslands are heavily invaded ecosystems, where non‐native annuals have largely dominated and replaced native communities. These systems are also highly variable, with a high degree of rainfall seasonality and interannual rainfall variability—a quality implicated in the coexistence of functionally distinct species. Yet, despite the apparent strength of this variation, coexistence between native and non‐native annuals in this system has faltered.To test how variation‐dependent coexistence mechanisms modulate failed coexistence, we implemented a competition experiment between two previously common native forbs and three now‐dominant non‐native annual grasses spanning a conservative‐acquisitive range of traits. We grew individuals from each species under varying densities of all other species as competitors, under either wetter or drier early season rainfall treatments. Using subsequent seed production, we parameterized competition models, assessed the potential for coexistence among species pairs and quantified the relative influence of variation‐dependent coexistence mechanisms.As expected, we found little potential for coexistence. Competition was dominated by the non‐native grassAvena fatua, while native forbs were unable to invade non‐native grasses. Mutual competitive exclusion was common across almost all species and often contingent on rainfall, suggesting rainfall‐mediated priority effects. Among variation‐dependent mechanisms, the temporal storage effect had a moderate stabilizing effect for four of five species when averaged across competitors, while relative nonlinearity in competition was largely destabilizing, except for the most conservative non‐native grass, which benefited from a competitive release under dry conditions.Synthesis: Our findings suggest that rainfall variability does little to mitigate the fitness differences that underlie widespread annual grass invasion in California, but that it influences coexistence dynamics among the now‐dominant non‐native grasses. 

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. 

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. 

Abstract Accompanying the climate crisis is the more enigmatic biodiversity crisis. Rapid reorganization of biodiversity due to global environmental change has defied prediction and tested the basic tenets of conservation and restoration. Conceptual and practical innovation is needed to support decision making in the face of these unprecedented shifts. Critical questions include: How can we generalize biodiversity change at the community level? When are systems able to reorganize and maintain integrity, and when does abiotic change result in collapse or restructuring? How does this understanding provide a template to guide when and how to intervene in conservation and restoration? To this end, we frame changes in community organization as the modulation of external abiotic drivers on the internal topology of species interactions, using plant–plant interactions in terrestrial communities as a starting point. We then explore how this framing can help translate available data on species abundance and trait distributions to corresponding decisions in management. Given the expectation that community response and reorganization are highly complex, the external‐driver internal‐topology (EDIT) framework offers a way to capture general patterns of biodiversity that can help guide resilience and adaptation in changing environments. 

Summary Predicting shifts in species composition with global change remains challenging, but plant functional traits provide a key link to scale from plant to community and ecosystem levels. The extent to which functional trait shifts may mediate ecosystem response to climate change remains a critical question.We ran point‐scale Community Land Model (CLM) simulations with site‐specific functional trait and phenology observations to represent alpine tundra growth strategies. We validated our results with site observations and compared parameterized results to those using the default parameterization. We then quantified the relative contribution of plant functional trait shifts vs climate change scenarios (and the resulting phenological shifts) to uncertainty in future tundra ecosystem productivity outcomes.We found that using community‐specific functional traits and phenology observations significantly improved productivity estimates compared with overestimates in a default simulation. Uncertainty in potential plant trait shifts often had a larger effect on ecosystem productivity responses than uncertainty in the forced response from different climate change scenarios.These findings highlight the key role of functional traits in shaping vegetation responses to climate change and the value of incorporating site‐level measurements into land models to more accurately forecast climate change impacts on ecosystem function. 

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.