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  1. 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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  2. 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.

     
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  3. Abstract

    Global change is altering patterns of community assembly, with net outcomes dependent on species' responses to the abiotic environment, both directly and mediated through biotic interactions. Here, we assess alpine plant community responses in a 15‐year factorial nitrogen addition, warming and snow manipulation experiment. We used a dynamic competition model to estimate the density‐dependent and ‐independent processes underlying changes in species‐group abundances over time. Density‐dependent shifts in competitive interactions drove long‐term changes in abundance of species‐groups under global change while counteracting environmental drivers limited the growth response of the dominant species through density‐independent mechanisms. Furthermore, competitive interactions shifted with the environment, primarily with nitrogen and drove non‐linear abundance responses across environmental gradients. Our results highlight that global change can either reshuffle species hierarchies or further favour already‐dominant species; predicting which outcome will occur requires incorporating both density‐dependent and ‐independent mechanisms and how they interact across multiple global change factors.

     
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  4. Abstract

    Global change alters ecosystems and their functioning, and biotic interactions can either buffer or amplify such changes. We utilized a long‐term nitrogen (N) addition and species removal experiment in the Front Range of Colorado, USA to determine whether a codominant forb and a codominant grass, with different effects on nutrient cycling and plant community structure, would buffer or amplify the effects of simulated N deposition on soil bacterial and fungal communities. While the plant community was strongly shaped by both the presence of dominant species and N addition, we did not find a mediating effect of the plant community on soil microbial response to N. In contrast to our hypothesis, we found a decoupling of the plant and microbial communities such that the soil microbial community shifted under N independently of directional shifts in the plant community. These findings suggest there are not strong cascading effects of N deposition across the plant–soil interface in our system.

     
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  5. Abstract

    As organisms shift their geographic distributions in response to climate change, biotic interactions have emerged as an important factor driving the rate and success of range expansions. Plant–microbe interactions are an understudied but potentially important factor governing plant range shifts. We studied the distribution and function of microbes present in high‐elevation unvegetated soils, areas that plants are colonizing as climate warms, snow melts earlier, and the summer growing season lengthens. Using a manipulative snowpack and microbial inoculation transplant experiment, we tested the hypothesis that growing‐season length and microbial community composition interact to control plant elevational range shifts. We predicted that a lengthening growing season combined with dispersal to patches of soils with more mutualistic microbes and fewer pathogenic microbes would facilitate plant survival and growth in previously unvegetated areas. We identified negative effects on survival of the common alpine bunchgrassDeschampsia cespitosain both short and long growing seasons, suggesting an optimal growing‐season length for plant survival in this system that balances time for growth with soil moisture levels. Importantly, growing‐season length and microbes interacted to affect plant survival and growth, such that microbial community composition increased in importance in suboptimal growing‐season lengths. Further, plants grown with microbes from unvegetated soils grew as well or better than plants grown with microbes from vegetated soils. These results suggest that the rate and spatial extent of plant colonization of unvegetated soils in mountainous areas experiencing climate change could depend on both growing‐season length and soil microbial community composition, with microbes potentially playing more important roles as growing seasons lengthen.

     
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  6. Abstract

    Fluctuations in population abundances are often correlated through time across multiple locations, a phenomenon known as spatial synchrony. Spatial synchrony can exhibit complex spatial structures, termed ‘geographies of synchrony’, that can reveal mechanisms underlying population fluctuations. However, most studies have focused on spatial extents of 10s to 100s of kilometres, making it unclear how synchrony concepts and approaches should apply to dynamics at finer spatial scales.

    We used network analyses, multiple regression on similarity matrices, and wavelet coherence analyses to examine micro‐scale synchrony and geographies of synchrony, over distances up to 30 m, in a serpentine grassland plant community.

    We found that species' populations exhibited a geography of synchrony even over such short distances. Often, well‐synchronized populations were geographically separate, a spatial structure that was shaped mainly by gopher disturbance and dispersal limitation, and to a lesser extent by relationships with other plant species. Precipitation was a significant driver of site‐ and community‐wide temporal dynamics. Gopher disturbance appeared to drive synchrony on 2‐ to 6‐year timescales, and we detected coherent fluctuations among pairs of focal plant taxa.

    Synthesis. Micro‐geographies of synchrony are an intriguing phenomenon that may also help us better understand community dynamics. Additionally, the related geographies of synchrony and coherent temporal dynamics among some species pairs indicate that incorporating interspecific interactions can improve understanding of population spatial synchrony.

     
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  7. Abstract

    Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments—species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

     
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