Climate warming threatens the persistence of a community of disturbance-adapted native annual plants.

With ongoing climate change, populations are expected to exhibit shifts in demographic performance that will alter where a species can persist. This presents unique challenges for managing plant populations and may require ongoing interventions, including in situ management or introduction into new locations. However, few studies have examined how climate change may affect plant demographic performance for a suite of species, or how effective management actions could be in mitigating climate change effects. Over the course of two experiments spanning 6 yr and four sites across a latitudinal gradient in the Pacific Northwest, United States, we manipulated temperature, precipitation, and disturbance intensity, and quantified effects on the demography of eight native annual prairie species. Each year we planted seeds and monitored germination, survival, and reproduction. We found that disturbance strongly influenced demographic performance and that seven of the eight species had increasingly poor performance with warmer conditions. Across species and sites, we observed 11% recruitment (the proportion of seeds planted that survived to reproduction) following high disturbance, but just 3.9% and 2.3% under intermediate and low disturbance, respectively. Moreover, mean seed production following high disturbance was often more than tenfold greater than under intermediate and low disturbance. Importantly, most species exhibited precipitous declines in their population growth rates (λ) under warmer-than-ambient experimental conditions and may require more frequent disturbance intervention to sustain populations. Aristida oligantha, a C4 grass, was the only species to have λ increase with warmer conditions. These results suggest that rising temperatures may cause many native annual plant species to decline, highlighting the urgency for adaptive management practices that facilitate their restoration or introduction to newly suitable locations. Frequent and intense disturbances are critical to reduce competitors and promote native annuals' persistence, but even such efforts may prove futile under future climate regimes.

Latitudinal gradients in population growth do not reflect demographic responses to climate.

Spatial gradients in population growth, such as across latitudinal or elevational gradients, are often assumed to primarily be driven by variation in climate, and are frequently used to infer species' responses to climate change. Here, we use a novel demographic, mixed-model approach to dissect the contributions of climate variables vs. other latitudinal or local site effects on spatiotemporal variation in population performance in three perennial bunchgrasses. For all three species, we find that performance of local populations decreases with warmer and drier conditions, despite latitudinal trends of decreasing population growth toward the cooler and wetter northern portion of each species' range. Thus, latitudinal gradients in performance are not predictive of either local or species-wide responses to climate. This pattern could be common, as many environmental drivers, such as habitat quality or species' interactions, are likely to vary with latitude or elevation, and thus influence or oppose climate responses.

Prairie plant phenology driven more by temperature than moisture in climate manipulations across a latitudinal gradient in the Pacific Northwest, USA.

Plant phenology will likely shift with climate change, but how temperature and/or moisture regimes will control phenological responses is not well understood. This is particularly true in Mediterranean climate ecosystems where the warmest temperatures and greatest moisture availability are seasonally asynchronous. We examined plant phenological responses at both the population and community levels to four climate treatments (control, warming, drought, and warming plus additional precipitation) embedded within three prairies across a 520 km latitudinal Mediterranean climate gradient within the Pacific Northwest, USA. At the population level, we monitored flowering and abundances in spring 2017 of eight range-restricted focal species planted both within and north of their current ranges. At the community level, we used normalized difference vegetation index (NDVI) measured from fall 2016 to summer 2018 to estimate peak live biomass, senescence, seasonal patterns, and growing season length. We found that warming exerted a stronger control than our moisture manipulations on phenology at both the population and community levels. Warming advanced flowering regardless of whether a species was within or beyond its current range. Importantly, many of our focal species had low abundances, particularly in the south, suggesting that establishment, in addition to phenological shifts, may be a strong constraint on their future viability. At the community level, warming advanced the date of peak biomass regardless of site or year. The date of senescence advanced regardless of year for the southern and central sites but only in 2018 for the northern site. Growing season length contracted due to warming at the southern and central sites (~3 weeks) but was unaffected at the northern site. Our results emphasize that future temperature changes may exert strong influence on the timing of a variety of plant phenological events, especially those events that occur when temperature is most limiting, even in seasonally water-limited Mediterranean ecosystems.

Greenhouse gas emissions limited by low nitrogen and carbon availability in natural, restored, and agricultural Oregon seasonal wetlands.

Wetlands are the major natural source of the greenhouse gas methane (CH4) and are also potentially an important source of nitrous oxide (N2O), though there is considerable variability among wetland types with some of the greatest uncertainty in freshwater mineral-soil wetlands. In particular, trace gas emissions from seasonal wetlands have been very poorly studied. We measured fluxes of CH4, N2O, and CO2(carbon dioxide), soil nutrients, and net primary productivity over one year in natural, restored, and agricultural seasonal wetland prairies in the Willamette Valley, Oregon, USA. We found zero fluxes for CH4 and N2O, even during periods of extended waterlogging of the soil. To explore this lack of emissions, we performed a laboratory experiment to examine the controls over these gases. In a fully-factorial design, we amended anaerobic soils from all wetlands with nitrate, glucose, and NaOH (to neutralize pH) and measured production potentials of N2, N2O, CH4, and CO2. We found that denitrification and N2O production were co-limited by nitrate and carbon, with little difference between the three wetland types. This co-limitation suggests that low soil carbon availability will continue to constrain N2O emissions and denitrification in these systems even when receiving relatively high levels of nitrogen inputs. Contrary to the results for N2O, the amended wetland soils never produced significant amounts of CH4 under any treatment. We hypothesize that high concentrations of alternative electron acceptors exist in these soils so that methanogens are noncompetitive with other microbial groups. As a result, these wetlands do not appear to be a significant source or sink of soil carbon and thus have a near zero climate forcing effect. Future research should focus on determining if this is a generalizable result in other seasonal wetlands.

Temperature response of soil respiration largely unaltered with experimental warming.

The respiratory release of carbon dioxide (CO2) from soil is a major yet poorly understood flux in the global carbon cycle. Climatic warming is hypothesized to increase rates of soil respiration, potentially fueling further increases in global temperatures. However, despite considerable scientific attention in recent decades, the overall response of soil respiration to anticipated climatic warming remains unclear. We synthesize the largest global dataset to date of soil respiration, moisture, and temperature measurements, totaling >3,800 observations representing 27 temperature manipulation studies, spanning nine biomes and over 2 decades of warming. Our analysis reveals no significant differences in the temperature sensitivity of soil respiration between control and warmed plots in all biomes, with the exception of deserts and boreal forests. Thus, our data provide limited evidence of acclimation of soil respiration to experimental warming in several major biome types, contrary to the results from multiple single-site studies. Moreover, across all nondesert biomes, respiration rates with and without experimental warming follow a Gaussian response, increasing with soil temperature up to a threshold of ∼25 °C, above which respiration rates decrease with further increases in temperature. This consistent decrease in temperature sensitivity at higher temperatures demonstrates that rising global temperatures may result in regionally variable responses in soil respiration, with colder climates being considerably more responsive to increased ambient temperatures compared with warmer regions. Our analysis adds a unique cross-biome perspective on the temperature response of soil respiration, information critical to improving our mechanistic understanding of how soil carbon dynamics change with climatic warming.

Experimental warming decreases arbuscular mycorrhizal fungal colonization in prairie plants along a Mediterranean climate gradient.

BACKGROUND: Arbuscular mycorrhizal fungi (AMF) provide numerous services to their plant symbionts. Understanding climate change effects on AMF, and the resulting plant responses, is crucial for predicting ecosystem responses at regional and global scales. We investigated how the effects of climate change on AMF-plant symbioses are mediated by soil water availability, soil nutrient availability, and vegetation dynamics. METHODS: We used a combination of a greenhouse experiment and a manipulative climate change experiment embedded within a Mediterranean climate gradient in the Pacific Northwest, USA to examine this question. Structural equation modeling (SEM) was used to determine the direct and indirect effects of experimental warming on AMF colonization. RESULTS: Warming directly decreased AMF colonization across plant species and across the climate gradient of the study region. Other positive and negative indirect effects of warming, mediated by soil water availability, soil nutrient availability, and vegetation dynamics, canceled each other out. DISCUSSION: A warming-induced decrease in AMF colonization would likely have substantial consequences for plant communities and ecosystem function. Moreover, predicted increases in more intense droughts and heavier rains for this region could shift the balance among indirect causal pathways, and either exacerbate or mitigate the negative, direct effect of increased temperature on AMF colonization.

Climate change alters plant biogeography in Mediterranean prairies along the West Coast, USA.

Projected changes in climate are expected to have widespread effects on plant community composition and diversity in coming decades. However, multisite, multifactor climate manipulation studies that have examined whether observed responses are regionally consistent and whether multiple climate perturbations are interdependent are rare. Using such an experiment, we quantified how warming and increased precipitation intensity affect the relative dominance of plant functional groups and diversity across a broad climate gradient of Mediterranean prairies. We implemented a fully factorial climate manipulation of warming (+2.5-3.0 °C) and increased wet-season precipitation (+20%) at three sites across a 520-km latitudinal gradient in the Pacific Northwest, USA. After seeding with a nearly identical mix of native species at all sites, we measured plant community composition (i.e., cover, richness, and diversity), temperature, and soil moisture for 3 years. Warming and the resultant drying of soils altered plant community composition, decreased native diversity, and increased total cover, with warmed northern communities becoming more similar to communities further south. In particular, after two full years of warming, annual cover increased and forb cover decreased at all sites mirroring the natural biogeographic pattern. This suggests that the extant climate gradient of increasing heat and drought severity is responsible for a large part of the observed biogeographic pattern of increasing annual invasion in US West Coast prairies as one moves further south. Additional precipitation during the rainy season did little to relieve drought stress and had minimal effects on plant community composition. Our results suggest that the projected increase in drought severity (i.e., hotter, drier summers) in Pacific Northwest prairies may lead to increased invasion by annuals and a loss of forbs, similar to what has been observed in central and southern California, resulting in novel species assemblages and shifts in functional composition, which in turn may alter ecosystem functions.

The herbaceous landlord: integrating the effects of symbiont consortia within a single host.

Plants are typically infected by a consortium of internal fungal associates, including endophytes in their leaves, as well as arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) in their roots. It is logical that these organisms will interact with each other and the abiotic environment in addition to their host, but there has been little work to date examining the interactions of multiple symbionts within single plant hosts, or how the relationships among symbionts and their host change across environmental conditions. We examined the grass Agrostis capillaris in the context of a climate manipulation experiment in prairies in the Pacific Northwest, USA. Each plant was tested for presence of foliar endophytes in the genus Epichloë, and we measured percent root length colonized (PRLC) by AMF and DSE. We hypothesized that the symbionts in our system would be in competition for host resources, that the outcome of that competition could be driven by the benefit to the host, and that the host plants would be able to allocate carbon to the symbionts in such a way as to maximize fitness benefit within a particular environmental context. We found a correlation between DSE and AMF PRLC across climatic conditions; we also found a fitness cost to increasing DSE colonization, which was negated by presence of Epichloë endophytes. These results suggest that selective pressure on the host is likely to favor host/symbiont relationships that structure the community of symbionts in the most beneficial way possible for the host, not necessarily favoring the individual symbiont that is most beneficial to the host in isolation. These results highlight the need for a more integrative, systems approach to the study of host/symbiont consortia.

An exploration of hypotheses that explain herbivore and pathogen attack in restored plant communities.

Many hypotheses address the associations of plant community composition with natural enemies, including: (i) plant species diversity may reduce enemy attack, (ii) attack may increase as host abundance increases, (iii) enemy spillover may lead to increased attack on one host species due to transmission from another host species, or enemy dilution may lead to reduced attack on a host that would otherwise have more attack, (iv) physical characteristics of the plant community may influence attack, and (v) plant vigor may affect attack. Restoration experiments with replicated plant communities provide an exceptional opportunity to explore these hypotheses. To explore the relative predictive strengths of these related hypotheses and to investigate the potential effect of several restoration site preparation techniques, we surveyed arthropod herbivore and fungal pathogen attack on the six most common native plant species in a restoration experiment. Multi-model inference revealed a weak but consistent negative correlation with pathogen attack and host diversity across the plant community, and no correlation between herbivory and host diversity. Our analyses also revealed host species-specific relationships between attack and abundance of the target host species, other native plant species, introduced plant species, and physical community characteristics. We found no relationship between enemy attack and plant vigor. We found minimal differences in plant community composition among several diverse site preparation techniques, and limited effects of site preparation techniques on attack. The strongest associations of community characteristics with attack varied among plant species with no community-wide patterns, suggesting that no single hypothesis successfully predicts the dominant community-wide trends in enemy attack.

Soil respiration response to climate change in Pacific Northwest prairies is mediated by a regional Mediterranean climate gradient.

Soil respiration is expected to increase with rising global temperatures but the degree of response may depend on soil moisture and other local factors. Experimental climate change studies from single sites cannot discern whether an observed response is site-dependent or generalizable. To deconvolve site-specific vs. regional climatic controls, we examined soil respiration for 18 months along a 520 km climate gradient in three Pacific Northwest, USA prairies that represents increasingly severe Mediterranean conditions from north to south. At each site we implemented a fully factorial combination of 2.5-3 °C warming and 20% added precipitation intensity. The response of soil respiration to warming was driven primarily by the latitudinal climate gradient and not site-specific factors. Warming increased respiration at all sites during months when soil moisture was not limiting. However, these gains were offset by reductions in respiration during seasonal transitions and summer drought due to lengthened periods of soil moisture limitation. The degree of this offset varied along the north-south climate gradient such that in 2011 warming increased cumulative annual soil respiration 28.6% in the northern site, 13.5% in the central site, and not at all in the southern site. Precipitation also stimulated soil respiration more frequently in the south, consistent with an increased duration of moisture limitation. The best predictors of soil respiration in nonlinear models were the Normalized Difference Vegetation Index (NDVI), antecedent soil moisture, and temperature but these models provided biased results at high and low soil respiration. NDVI was an effective integrator of climate and site differences in plant productivity in terms of their combined effects on soil respiration. Our results suggest that soil moisture limitation can offset the effect of warming on soil respiration, and that greater growing-season moisture limitation would constrain cumulative annual responses to warming.

Pushing the limit: experimental evidence of climate effects on plant range distributions.

Whether species will be extirpated in their current geographic ranges due to rapidly changing climate, and if so, whether they can avoid extinction by shifting their distributions are pressing questions for biodiversity conservation. However, forecasts of climate change impacts on species' geographic distributions rarely incorporate a demographic understanding of species' responses to climate. Because many biotic and abiotic factors at multiple scales control species' range limits, experimentation is essential to establish underlying mechanisms. We used a manipulative climate change experiment embedded within a natural climate gradient to examine demographic responses of 12 prairie species with northern range limits within the Pacific Northwest, USA. During the first year, warming decreased recruitment of species even at the coolest edge of their current ranges, but this effect disappeared when they were moved poleward beyond their current ranges. This response was largely driven by differences in germination rates. Other vital rates responded in unique and sometimes opposing ways (survivorship vs. fitness) to species' current ranges and climate change, and were mediated by indirect effects of climate on competition and nutrient availability. Our results demonstrate the importance of using regional-scale climate manipulations and the need for longer-term experiments on the demographic responses that control species' distributions.

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie.

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments.