Abstract. Global change is altering species distributions and thus interactions among organisms.Organisms live in concert with thousands of other species, some beneficial, some pathogenic, some which have little to no effect in complex communities. Since natural communities are composed of organisms with very different life history traits and dispersal ability it is unlikely they will all respond to climatic change in a similar way. Disjuncts in plant-pollinator and plant-herbivore interactions under global change have been relatively well described, but plant-soil microorganism and soil microbe-microbe relationships have received less attention. Since soil microorganisms regulate nutrient transformations, provide plants with nutrients, allow co-existence among neighbors, and control plant populations, changes in soil microorganism-plant interactions could have significant ramifications for plant community composition and ecosystem function. In this paper we explore how climatic change affects soil microbes and soil microbe-plant interactions directly and indirectly, discuss what we see as emerging and exciting questions and areas for future research, and discuss what ramifications changes in these interactions may have on the composition and function of ecosystems.

Summary Plant roots, their associated microbial community and free‐living soil microbes interact to regulate the movement of carbon from the soil to the atmosphere, one of the most important and least understood fluxes of terrestrial carbon. Our inadequate understanding of how plant–microbial interactions alter soil carbon decomposition may lead to poor model predictions of terrestrial carbon feedbacks to the atmosphere. Roots, mycorrhizal fungi and free‐living soil microbes can alter soil carbon decomposition through exudation of carbon into soil. Exudates of simple carbon compounds can increase microbial activity because microbes are typically carbon limited. When both roots and mycorrhizal fungi are present in the soil, they may additively increase carbon decomposition. However, when mycorrhizas are isolated from roots, they may limit soil carbon decomposition by competing with free‐living decomposers for resources. We manipulated the access of roots and mycorrhizal fungi to soil in situ in a temperate mixed deciduous forest. We added 13C‐labelled substrate to trace metabolized carbon in respiration and measured carbon‐degrading microbial extracellular enzyme activity and soil carbon pools. We used our data in a mechanistic soil carbon decomposition model to simulate and compare the effects of root and mycorrhizal fungal presence on soil carbon dynamics over longer time periods. Contrary to what we predicted, root and mycorrhizal biomass did not interact to additively increase microbial activity and soil carbon degradation. The metabolism of 13C‐labelled starch was highest when root biomass was high and mycorrhizal biomass was low. These results suggest that mycorrhizas may negatively interact with the free‐living microbial community to influence soil carbon dynamics, a hypothesis supported by our enzyme results. Our steady‐state model simulations suggested that root presence increased mineral‐associated and particulate organic carbon pools, while mycorrhizal fungal presence had a greater influence on particulate than mineral‐associated organic carbon pools. Synthesis. Our results suggest that the activity of enzymes involved in organic matter decomposition was contingent upon root–mycorrhizal–microbial interactions. Using our experimental data in a decomposition simulation model, we show that root–mycorrhizal–microbial interactions may have longer‐term legacy effects on soil carbon sequestration. Overall, our study suggests that roots stimulate microbial activity in the short term, but contribute to soil carbon storage over longer periods of time.

Climate change affects communities both directly and indirectly via changes in interspecific interactions. One such interaction that may be altered under climate change is the ant-plant seed dispersal mutualism common in deciduous forests of eastern North America. As climatic warming alters the abundance and activity levels of ants, the potential exists for shifts in rates of ant-mediated seed dispersal. We used an experimental temperature manipulation at two sites in the eastern US (Harvard Forest in Massachusetts and Duke Forest in North Carolina) to examine the potential impacts of climatic warming on overall rates of seed dispersal (using Asarum canadense seeds) as well as species-specific rates of seed dispersal at the Duke Forest site. We also examined the relationship between ant critical thermal maxima (CTmax) and the mean seed removal temperature for each ant species. We found that seed removal rates did not change as a result of experimental warming at either study site, nor were there any changes in species-specific rates of seed dispersal. There was, however, a positive relationship between CTmax and mean seed removal temperature, whereby species with higher CTmax removed more seeds at hotter temperatures. The temperature at which seeds were removed was influenced by experimental warming as well as diurnal and day-to-day fluctuations in temperature. Taken together, our results suggest that while temperature may play a role in regulating seed removal by ants, ant plant seed-dispersal mutualisms may be more robust to climate change than currently assumed.

Roots release carbon into soil and can alleviate energy limitation of microbial organic matter decomposition. We know little about the effects of roots on microbial decomposition of different organic matter substrates, despite the importance for soil carbon stocks and turnover. Through implementing root–microbe interactions, the Carbon, Organisms, Rhizosphere and Protection in the Soil Environment (CORPSE) model was previously shown to represent dynamics of total soil carbon in temperate forest field experiments. However, the model permits alternative hypotheses concerning microbial‐substrate affinity. We investigated how root inputs affect decomposition of soil organic carbon (SOC) with variable decomposability. We simulated SOC stocks in CORPSE and compared microbial degradation of two substrate types with varying root–microbe interactions under two alternative hypotheses that varied in microbial‐substrate affinity. We compared our modelled hypotheses to a forest field experiment where we quantified decomposition of isotopically labelled starch and leaf tissues in soils with manipulated root access to microbes. We tested the hypothesis that decomposition of leaves would be more sensitive to root inputs than decomposition of starch, corresponding to the alternative model hypotheses. In the field study, leaf decomposition increased with root density, whereas starch decomposition was unchanged by root density. Microbial biomass and enzyme activity consistently increased with root inputs in CORPSE and the field study. Our field experiment supported the CORPSE simulations with high microbial‐substrate affinity. Roots stimulated microbial growth and enzyme production, which increased the degradation of more complex substrates such as leaf tissues. Substrates that were easily decomposed, such as starch, may already be degrading at a maximum rate in the absence of rhizosphere influence because their decomposition rate was unchanged by root inputs. We found that the degree to which roots stimulate microbial decomposition depends on the substrate being decomposed, and that root–microbe interactions influenced SOC stocks in both our model and field experiment. Environmental changes that alter root–microbe interactions could, therefore, alter soil C stocks and biogeochemical cycling, and models of these interactions should incorporate differential influence of rhizosphere inputs on different substrates. A free Plain Language Summary can be found within the Supporting Information of this article.

To maximize limited conservation funds and prioritize management projects that are likely to succeed, accurate assessment of invasive nonnative species impacts is essential. A common challenge to prioritization is a limited knowledge of the difference between the impacts of a single nonnative species compared to the impacts of nonnative species when they co-occur, and in particular predicting when impacts of co-occurring nonnative species will be non-additive. Understanding non-additivity is important for management decisions because the management of only one co-occurring invader will not necessarily lead to a predictable reduction in the impact or growth of the other nonnative plant. Nonnative plants are frequently associated with changes in soil biotic and abiotic characteristics, which lead to plant-soil interactions that influence the performance of other species grown in those soils. Whether co-occurring nonnative plants alter soil properties additively or non-additively relative to their effects on soils when they grow in monoculture is rarely addressed. We use a greenhouse plant-soil feedback experiment to test for non-additive soil impacts of two common invasive nonnative woody shrubs, Lonicera maackii and Ligustrum sinense, in deciduous forests of the southeastern United States. We measured the performance of each nonnative shrub, a native herbaceous community, and a nonnative woody vine in soils conditioned by each shrub singly or together in polyculture. Soils conditioned by both nonnative shrubs had non-additive impacts on native and nonnative performance. Root mass of the native herbaceous community was 1.5 times lower and the root mass of the nonnative L. sinense was 1.8 times higher in soils conditioned by both L. maackii and L. sinense than expected based upon growth in soils conditioned by either shrub singly. This result indicates that when these two nonnative shrubs co-occur, their influence on soils disproportionally favors persistence of the nonnative L. sinense relative to this native herbaceous community, and could provide an explanation of why native species abundance is frequently depressed in these communities. Additionally, the difference between native and nonnative performance demonstrates that invasive impact studies focusing on the impact only of single species can be insufficient for determining the impact of co-occurring invasive plant species.

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19Climate change affects communities both directly and indirectly via changes in interspecific 20 interactions. One such interaction that may be altered under climate change is the ant-plant seed 21 dispersal mutualism common in deciduous forests of the eastern US. As climatic warming alters 22 the abundance and activity levels of ants, the potential exists for shifts in rates of ant-mediated 23 seed removal. We used an experimental temperature manipulation at two sites in the eastern US as well as species-specific rates of seed dispersal at the Duke Forest site. We also examined the 27 relationship between ant critical thermal maxima (CT max ) and the mean seed removal 28 temperature for each ant species. We found that seed removal rates did not change as a result of 29 experimental warming at either study site, nor were there any changes in species-specific rates of 30 seed dispersal. There was, however, a positive relationship between CT max and mean seed 31 removal temperature, whereby species with higher CT max removed more seeds at hotter 32 temperatures. The temperature at which seeds were removed was influenced by experimental 33 warming as well as diurnal and day-to-day fluctuations in temperature. Taken together, our 34 results suggest that while temperature may play a role in regulating seed removal by ants, ant 35 plant seed-dispersal mutualisms may be more robust to climate change than currently assumed.

Heatwaves are becoming more frequent and intense with climate change, but the demographic and evolutionary consequences of heatwaves are rarely investigated in herbaceous plant species. We examine the consequences of a short but extreme heatwave in Oregon populations of the common yellow monkeyflower (Mimulus guttatus) by leveraging a common garden experiment planted with range wide populations and observational studies of eleven local populations. In the common garden, 89% of seedlings died during the heatwave including >96% of seedlings from geographically local populations. Some populations from hotter and drier environments had higher fitness, however others from comparable environments performed poorly. Observational studies of local natural populations drastically differed in the consequences of the heatwave: one population was completely extirpated and nearly half had a >50% decrease in fitness. However, a few populations had greater fitness during the heatwave year. Differences in mortality corresponded to the impact of the heatwave on soil moisture where retention of soil moisture throughout the heatwave led to greater survivorship. Our results suggest that not all populations experience the same intensity or degree of mortality during extreme events and such heterogeneity could be important for genetic rescue or to facilitate the distribution of adaptive variants throughout the region.