Understanding the interactive effects of species invasions and climate change is essential for predicting future shifts in biodiversity. Because multiple stressors can interact in synergistic or antagonistic ways, it is notoriously difficult to anticipate their combined effects on species assemblages. However, some hypotheses predict that plant invasions will become increasingly problematic as climate change improves conditions for invaders or lowers the biotic resistance of native communities. In a 4‐yr field experiment, we quantified the individual and interactive effects of invasion by a globally problematic C4 grass, Imperata cylindrica, and chronic simulated drought imposed by rainout shelters on the whole plant communities of regenerating longleaf pine forest. Invasion both inhibited plant colonization and enhanced plot‐level extinctions, resulting in a severe (60%) loss of plant diversity across all functional groups, including perennial grasses and forbs, annual forbs, and woody species and dramatic shifts in community composition. Experimental drought reduced diversity by 20%, and caused a shift in the dominant functional groups, but had no significant effect on cover of the invader. The invader partially ameliorated water stress in the drought treatment such that invaded plots had higher soil moisture than uninvaded plots. Consequently, the combined effects of invasion and drought were lower than expected from an additive model of multiple stressors. These findings, which may have broader implications for how other C4 grass invaders will interact with drought to shift native community dynamics, challenge the perception that climate change will exacerbate invasions. In revealing that invasive species pose a major threat to the diversity and structure of native communities despite their moderating effects on abiotic stress, this work also highlights that management of aggressive invaders may be critical to preserving biodiversity regardless of future climate.

Abiotic global change drivers affect ecosystem structure and function, but how they interact with biotic factors such as invasive plants is understudied. Such interactions may be additive, synergistic, or offsetting, and difficult to predict. We present methods to test the individual and interactive effects of drought and plant invasion on native ecosystems. We coupled a factorial common garden experiment containing resident communities exposed to drought (imposed with rainout shelters) and invasion with a field experiment where the invader was removed from sites spanning a natural soil moisture gradient. We detail treatments and their effects on abiotic conditions, including soil moisture, light, temperature, and humidity, which shape community and ecosystem responses. Ambient precipitation during the garden experiment exceeded historic norms despite severe drought in prior years. Soil moisture was 48% lower in drought than ambient plots, but the invader largely offset drought effects. Additionally, temperature and light were lower and humidity higher in invaded plots. Field sites spanned up to a 10-fold range in soil moisture and up to a 2.5-fold range in light availability.Invaded and resident vegetation did not differentially mediate soil moisture, unlike in the garden experiment. Herbicide effectively removed invaded and resident vegetation, with removal having site-specific effects on soil moisture and light availability. However, light was generally higher in invader-removal than control plots, whereas resident removal had less effect on light, similar to the garden experiment. Invasion mitigated a constellation of abiotic conditions associated with drought stress in the garden experiment. In the field, where other factors co-varied, these patterns did not emerge. Still, neither experiment suggested that drought and invasion will have synergistic negative effects on ecosystems, although invasion can limit light availability. Coupling factorial garden experiments with field experiments across environmental gradients will be effective for predicting how multiple stressors interact in natural systems.

Global change stressors such as drought and plant invasion can affect ecosystem structure and function via mediation of resource availability and plant competition outcomes. Yet, it remains uncertain how native plants respond to drought stress that co-occurs with potentially novel resource conditions created by a nonnative invader. Further, there is likely to be temporal variation in competition outcomes between native and nonnative plant species depending on which resources are most limiting at a given time. Interacting stressors coupled with temporal variation make it difficult to predict how global change will impact native plant communities. To address this knowledge gap, we conducted a 5-yr factorial field experiment to quantify how simulated drought, plant invasion (by cogongrass, Imperata cylindrica), and these stressors combined, affected resource availability (soil moisture and light) and competition dynamics between the invader and native longleaf pine (Pinus palustris), a foundation species in southeast U.S. forests. Drought and invasion mediated the survival and performance of pine seedlings in temporally dynamic and unexpected ways. Drought and invasion alone each significantly reduced pine seedling survival. However, when the stressors occurred together, the invader offset drought stress for pine seedlings by maintaining high levels of soil moisture, humidity, and shade compared to uninvaded vegetation. This facilitative effect was pronounced for 2 yr, yet shifted to strong competitive exclusion as the invasion progressed and the limiting resource switched from soil moisture to light. After 3 yr, pine tree survival was low except for pines growing with uninvaded vegetation under ambient precipitation conditions. After 5 yr, pines experiencing a single stressor were taller and had greater height to diameter ratios than pines under no stress or both stressors. This outcome revealed a filtering effect where poorly performing trees were culled under stressful conditions, especially when pines were growing with the invader. Together, these results demonstrate that although drought and invasion suppressed a foundation tree species, the invader temporarily moderated stressful drought conditions, and at least some trees were able to survive despite increasingly strong competition. Such unpredictable effects of interacting global change stressors on native plant species highlight the need for additional long-term studies.

Soil microbiomes could play a major role in ecosystem responses to escalating anthropogenic global change. However, we currently have a poor understanding of how soil microbes will respond to interacting global change factors and if responses will be mediated by changes in plant community structure. We used a field experiment to assess changes in soil fungal and bacterial communities in response to plant invasion, experimental drought, and their combination. In addition, we evaluated the relative importance of direct versus indirect pathways of invasion and drought through changes in associated plant communities with structural equation models. We found that fungal communities were interactively structured by invasion and drought, where fungal richness was lowest with invasion under ambient conditions but highest with invasion under drought conditions. Bacterial richness was lower under drought but unaffected by invasion. Changes in the plant community, including lower plant richness and higher root biomass, moderated the direct effects of invasion on microbial richness. Fungal and bacterial functional groups, including pathogens, mutualists, and nitrogen metabolizers, were also influenced by plant community changes. In sum, plant communities mediated the effects of interacting global change drivers on soil microbial community structure, with significant potential consequences for community dynamics and ecosystem functions.

828 I. What is codependency and why should we care? 829 II. Should we expect codependency among AM communities? 830 III. Requirements for codependency 830 IV. When and where has codependency been observed? 831 V. Is codependency scale-and resolution-dependent? 831 VI. Unexplained variation: if not codependency then what? 834 VII Recommendations for future studies 834 VIII. Conclusion 835 Acknowledgements 835 References 835

Symbiotic arbuscular mycorrhizal fungi (AMF) are ubiquitous in tropical forests. AMF play a role in the forest carbon cycle because they can increase nutrient acquisition and biomass of host plants, but also incur a carbon cost to the plant. Through their interactions with their host plants they have the potential to affect how plants respond to environmental perturbation such as global warming. Our objective was to experimentally determine how plant respiration rates and responses to warmer environment are affected by AMF colonization in seedlings of five tropical tree species at the whole plant level. We evaluated the interaction between AMF colonization and temperature on plant respiration against four possible outcomes; acclimation does or does not occur regardless of AMF, or AMF can increase or decrease respiratory acclimation. Seedlings were inoculated with AMF spores or sterilized inoculum and grown at ambient or elevated nighttime temperature. We measured whole plant and belowground respiration rates, as well as plant growth and biomass allocation. There was an overall increase in whole plant, root, and shoot respiration rate with AMF colonization, whereas temperature acclimation varied among species, showing support for three of the four possible responses. The influence of AMF colonization on growth and allocation also varied among plant species. This study shows that the effect of AMF colonization on acclimation differs among plant species. Given the cosmopolitan nature of AMF and the importance of plant acclimation for predicting climate feedbacks a better understanding of the patterns and mechanisms of acclimation is essential for improving predictions of how climate warming may influence vegetation feedbacks.

1. Invasive plants can alter soil microbial communities and generate positive plantsoil feedbacks that facilitate their performance, but the magnitude and direction of feedbacks may change with novel conditions under climate change. We assessed how potential soil legacy effects of plant invasion and simulated drought influenced plant performance and competition in the longleaf pine ecosystem.2. We collected soil from a 4-year factorial invasion (cogongrass Imperata cylindrica) by drought (simulated with rainout shelters) field experiment and used it as live or sterilized soil inoculum in a greenhouse experiment that included two native foundation species, longleaf pine Pinus palustris and wiregrass Aristida stricta, and cogongrass, grown individually or in competition.3. There was no evidence of biotic soil legacy effects of invasion or drought for any plant species, but microbes played a significant role in competition. When plants were grown alone, the invader had 12% greater biomass in live soil than sterile soil but both native species had 25% less biomass in live soil. When grown in competition, these effects were reversed for cogongrass (37% smaller in live soil) and pine (17% larger in live soil). In competition, the three species grown in sterile soil produced similar amounts of biomass, whereas live soil created a competitive hierarchy where pine was more competitive than wiregrass and cogongrass. 4.Synthesis. These results emphasize the importance of soil biota in native plant restoration because, although the invader was highly successful when grown alone, plant-microbe interactions influenced the outcome of competition between native and invasive species by promoting native longleaf pine. There was little evidence that invasive cogongrass inhibited native plants via biotic soil legacies, instead, results suggested that plant-soil interactions can be highly resilient to global change such that the biotic legacy of invasion and drought may not promote or inhibit invasion.

Interactions with soil microbiota determine the success of restoring plants to their native habitats. The goal of our study was to understand the effects of restoration practices on interactions of giant sequoia Sequoiadendron giganteum with arbuscular mycorrhizal (AM) fungi (Glomeromycota). Natural regeneration of Sequoiadendron is threatened by the absence of severe fires that create forest canopy gaps. Generating artificial canopy gaps offers an alternative tool for giant sequoia restoration. We investigated the effect of regeneration practices, including (i) sapling location within gaps, (ii) gap size and (iii) soil substrate, on AM fungal colonization of giant sequoia sapling roots in a native giant sequoia grove of the Sierra Nevada, California. We found that the extent of AM fungal root colonization was positively correlated with sapling height and light availability, which were related to the location of the sapling within the gap and the gap size. While colonization frequency by arbuscules in saplings on ash substrate was higher relative to saplings in mineral soil, the total AM fungal root colonization was similar between the substrates. A negative correlation between root colonization by Glomeromycota and non-AM fungal species indicated antagonistic interactions between different classes of root-associated fungi. Using DNA genotyping, we identified six AM fungal taxa representing genera Glomus and Ambispora present in Sequoiadendron roots. Overall, we found that AM fungal colonization of giant sequoia roots was associated with availability of plant-assimilated carbon to the fungus rather than with the AM fungal supply of mineral nutrients to the roots. We conclude that restoration practices affecting light availability and carbon assimilation alter feedbacks between sapling growth and activity of AM fungi in the roots.