Tamara J. Zelikova

Long-term exposure to elevated CO2 enhances plant community stability by suppressing dominant plant species in a mixed-grass prairie

Significance Evaluating ecological responses to climate change is essential to predict ecosystem function under future climate scenarios. In a mixed-grass prairie, we use a multifactor field experiment to show that the effects of elevated CO2 and warming on plant community structure and productivity depend on interannual variation in precipitation. We also show that shifts in plant dominance patterns driven by elevated CO2 in a mixed-grass prairie ecosystem promoted biomass and compositional stability and resistance to interannual variation in precipitation. The economic value of grasslands is largely dependent on the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate. Climate controls vegetation distribution across the globe, and some vegetation types are more vulnerable to climate change, whereas others are more resistant. Because resistance and resilience can influence ecosystem stability and determine how communities and ecosystems respond to climate change, we need to evaluate the potential for resistance as we predict future ecosystem function. In a mixed-grass prairie in the northern Great Plains, we used a large field experiment to test the effects of elevated CO2, warming, and summer irrigation on plant community structure and productivity, linking changes in both to stability in plant community composition and biomass production. We show that the independent effects of CO2 and warming on community composition and productivity depend on interannual variation in precipitation and that the effects of elevated CO2 are not limited to water saving because they differ from those of irrigation. We also show that production in this mixed-grass prairie ecosystem is not only relatively resistant to interannual variation in precipitation, but also rendered more stable under elevated CO2 conditions. This increase in production stability is the result of altered community dominance patterns: Community evenness increases as dominant species decrease in biomass under elevated CO2. In many grasslands that serve as rangelands, the economic value of the ecosystem is largely dependent on plant community composition and the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate.

Seasonality of soil moisture mediates responses of ecosystem phenology to elevated CO2 and warming in a semi‐arid grassland

Summary Vegetation greenness, detected using digital photography, is useful for monitoring phenology of plant growth, carbon uptake and water loss at the ecosystem level. Assessing ecosystem phenology by greenness is especially useful in spatially extensive, water-limited ecosystems such as the grasslands of the western United States, where productivity is moisture dependent and may become increasingly vulnerable to future climate change. We used repeat photography and a novel means of quantifying greenness in digital photographs to assess how the individual and combined effects of warming and elevated CO2 impact ecosystem phenology (greenness and plant cover) in a semi-arid grassland over an 8-year period. Climate variability within and among years was the proximate driver of ecosystem phenology. Individual and combined effects of warming and elevated CO2 were significant at times, but mediated by variation in both intra- and interannual precipitation. Specifically, warming generally enhanced plant cover and greenness early in the growing season but often had a negative effect during the middle of the summer, offsetting the early season positive effects. The individual effects of elevated CO2 on plant cover and greenness were generally neutral. Opposing seasonal variations in the effects of warming and less so elevated CO2 cancelled each other out over an entire growing season, leading to no net effect of treatments on annual accumulation of greenness. The main effect of elevated CO2 dampened quickly, but warming continued to affect plant cover and plot greenness throughout the experiment. The combination of warming and elevated CO2 had a generally positive effect on greenness, especially early in the growing season and in later years of the experiment, enhanced annual greenness accumulation. However, interannual precipitation variation had larger effect on greenness, with two to three times greater greenness in wet years than in dry years. Synthesis. Seasonal variation in timing and amount of precipitation governs grassland phenology, greenness and the potential for carbon uptake. Our results indicate that concurrent changes in precipitation regimes mediate vegetation responses to warming and elevated atmospheric CO2 in semi-arid grasslands. Even small changes in vegetation phenology and greenness in response to warming and rising atmospheric CO2 concentrations, such as those we report here, can have large consequences for the future of grasslands.

Eco-evolutionary responses of Bromus tectorum to climate change: implications for biological invasions.

How plant populations, communities, and ecosystems respond to climate change is a critical focus in ecology today. The responses of introduced species may be especially rapid. Current models that incorporate temperature and precipitation suggest that future Bromus tectorum invasion risk is low for the Colorado Plateau. With a field warming experiment at two sites in southeastern Utah, we tested this prediction over 4 years, measuring B. tectorum phenology, biomass, and reproduction. In a complimentary greenhouse study, we assessed whether changes in field B. tectorum biomass and reproductive output influence offspring performance. We found that following a wet winter and early spring, the timing of spring growth initiation, flowering, and summer senescence all advanced in warmed plots at both field sites and the shift in phenology was progressively larger with greater warming. Earlier green-up and development was associated with increases in B. tectorum biomass and reproductive output, likely due early spring growth, when soil moisture was not limiting, and a lengthened growing season. Seeds collected from plants grown in warmed plots had higher biomass and germination rates and lower mortality than seeds from ambient plots. However, in the following two dry years, we observed no differences in phenology between warmed and ambient plots. In addition, warming had a generally negative effect on B. tectorum biomass and reproduction in dry years and this negative effect was significant in the plots that received the highest warming treatment. In contrast to models that predict negative responses of B. tectorum to warmer climate on the Colorado Plateau, the effects of warming were more nuanced, relied on background climate, and differed between the two field sites. Our results highlight the importance of considering the interacting effects of temperature, precipitation, and site-specific characteristics such as soil texture, on plant demography and have direct implications for B. tectorum invasion dynamics on the Colorado Plateau.

Challenging terrestrial biosphere models with data from the long-term multifactor Prairie Heating and CO2 Enrichment experiment

Abstract Multifactor experiments are often advocated as important for advancing terrestrial biosphere models (TBMs), yet to date, such models have only been tested against single‐factor experiments. We applied 10 TBMs to the multifactor Prairie Heating and CO2 Enrichment (PHACE) experiment in Wyoming, USA. Our goals were to investigate how multifactor experiments can be used to constrain models and to identify a road map for model improvement. We found models performed poorly in ambient conditions; there was a wide spread in simulated above‐ground net primary productivity (range: 31–390 g C m−2 yr−1). Comparison with data highlighted model failures particularly with respect to carbon allocation, phenology, and the impact of water stress on phenology. Performance against the observations from single‐factors treatments was also relatively poor. In addition, similar responses were predicted for different reasons across models: there were large differences among models in sensitivity to water stress and, among the N cycle models, N availability during the experiment. Models were also unable to capture observed treatment effects on phenology: they overestimated the effect of warming on leaf onset and did not allow CO2‐induced water savings to extend the growing season length. Observed interactive (CO2 × warming) treatment effects were subtle and contingent on water stress, phenology, and species composition. As the models did not correctly represent these processes under ambient and single‐factor conditions, little extra information was gained by comparing model predictions against interactive responses. We outline a series of key areas in which this and future experiments could be used to improve model predictions of grassland responses to global change.

Ecological Genetics, Local Adaptation, and Phenotypic Plasticity in Bromus tectorum in the Context of a Changing Climate

Effective management of invasive species spread requires understanding the potential of exotic species to colonize different habitat types. In the case of Bromus tectorum, colonization potential includes persisting in variable environments via phenotypic plasticity or via genetic variation. Bromus tectorum L. (cheatgrass or downy brome) is a highly invasive, self-pollinating, winter annual grass that was introduced to the intermountain region of North America around 1890 and expanded to its modern range within 40 years. Common garden studies have helped shed light on outcrossing frequency, microsite effects on establishment and growth, traits that could confer invasiveness, and variation in germination, morphology, and physiology. Here, we review the evidence for existing local adaptation and phenotypic plasticity in B. tectorum in its invaded range along with the potential for responses to climate change and discuss implications of both for its success as an invader and future management. All of these studies show that B. tectorum can tolerate a wide range of habitats as the result of genetic variation among populations, a range of locally adapted ecotypes, and phenotypic plasticity. The success of B. tectorum could be due to its ability to maintain fitness in both high-quality and marginal environments.

Seasonally contrasting responses of evapotranspiration to warming and elevated CO 2 in a semiarid grassland

Global climate change is expected to alter seasonal patterns and rates of evapotranspiration in dry regions. While climate change will involve elevated CO2 and increased temperatures, independently these factors may have different impacts on actual evapotranspiration (AET) due to their opposing effects on transpiration. We used canopy gas exchange chambers to quantify AET in a semi-arid grassland experimentally altered by elevated CO2 and warming over three years with contrasting ambient precipitation. Seasonal and interannual variations in AET due to background climate variability were larger than the effects of climate manipulation treatments. However, in a year with average precipitation, cumulative growing season AET was suppressed by warming by 23%. Across years, warming increased AET early in the growing season and suppressed it later in the growing season. By contrast, elevated CO2 suppressed AET early in the growing season and enhanced it later, but only in years with average or above-average precipitation. Vegetation greenness (a proxy for photosynthetically active leaf area) was consistently the strongest predictor of AET, while soil moisture and vapor pressure deficit were secondary drivers. Our research demonstrates that effects of increased atmospheric CO2 and temperature on AET will be mediated by plant phenological development and seasonal climatic conditions.

Elevated CO2 and water addition enhance nitrogen turnover in grassland plants with implications for temporal stability

Temporal variation in soil nitrogen (N) availability affects growth of grassland communities that differ in their use and reuse of N. In a 7-year-long climate change experiment in a semi-arid grassland, the temporal stability of plant biomass production varied with plant N turnover (reliance on externally acquired N relative to internally recycled N). Species with high N turnover were less stable in time compared to species with low N turnover. In contrast, N turnover at the community level was positively associated with asynchrony in biomass production, which in turn increased community temporal stability. Elevated CO2 and summer irrigation, but not warming, enhanced community N turnover and stability, possibly because treatments promoted greater abundance of species with high N turnover. Our study highlights the importance of plant N turnover for determining the temporal stability of individual species and plant communities affected by climate change.