Soil extracellular enzymes are the proximal drivers of decomposition. However, the relative influence of climate, soil nutrients and edaphic factors compared to microbial community composition on extracellular enzyme activities (EEA) is poorly resolved. Determining the relative effects of these factors on soil EEA is critical since changes in climate and microbial species composition may have large impacts on decomposition. We measured EEA from five sites during the growing season in March and 17 sites during the dry season in July throughout southern California and simultaneously collected data on climate, soil nutrients, soil edaphic factors and fungal community composition. The concentration of carbon and nitrogen in the soil and soil pH were most related to hydrolytic EEA. Conversely, oxidative EEA was mostly related to mean annual precipitation. Fungal community composition was not correlated with EEA at the species, genus, family or order levels. The hyphal length of fungi was correlated with EEA during the growing season while relative abundance of taxa within fungal phyla, in particular Chytridiomycota, was correlated with the EEA of beta-glucosidase, cellobiohydrolase, acid phosphatase and beta-xylosidase in the dry season. Overall, in the dry season, 35.3 % of the variation in all enzyme activities was accounted for by abiotic variables, while fungal composition accounted for 27.4 %. Because global change is expected to alter precipitation regimes and increase nitrogen deposition in soils, EEA may be affected, with consequences for decomposition.

Arbuscular mycorrhizal (AM) fungi form associations with most land plants and can control carbon, nitrogen, and phosphorus cycling between above- and belowground components of ecosystems. Current estimates of AM fungal distributions are mainly inferred from the individual distributions of plant biomes, and climatic factors. However, dispersal limitation, local environmental conditions,and interactions among AM fungal taxa may also determine local diversity and global distributions. We assessed the relative importance of these potential controls by collecting 14,961 DNA sequences from 111 published studies and testing for relationships between AM fungal community composition and geography, environment, and plant biomes. Our results indicated that the global species richness of AM fungi was up to six times higher than previously estimated, largely owing to high beta diversity among sampling sites. Geographic distance, soil temperature and moisture, and plant community type were each significantly related to AM fungal community structure, but explained only a small amount of the observed variance. AM fungal species also tended to be phylogenetically clustered within sites, further suggesting that habitat filtering or dispersal limitation is a driver of AM fungal community assembly. Therefore, predicted shifts in climate and plant species distributions under global change may alter AM fungal communities.

Evolutionary trade-offs among ecological traits are one mechanism that could determine the responses of functional groups of decomposers to global changes such as nitrogen (N) enrichment. We hypothesised that bacteria targeting recalcitrant carbon compounds require relatively high levels of N availability to support the construction costs of requisite extracellular and transport enzymes. Indeed, we found that taxa that used more recalcitrant (i.e. larger and cyclic) carbon compounds were more prevalent in ocean waters with higher nitrate concentrations. Compared to recalcitrant carbon users, labile carbon users targeted more organic N compounds, were found in relatively nitrate-poor waters, and were more common in higher latitude soils, which is consistent with the paradigm that N-limitation is stronger at higher latitudes. Altogether, evolutionary trade-offs may limit recalcitrant carbon users to habitats with higher N availability.

The relative importance of dispersal limitation versus environmental filtering for community assembly has received much attention for macroorganisms. These processes have only recently been examined in microbial communities. Instead, microbial dispersal has mostly been measured as community composition change over space (i.e., distance decay). Here we directly examined fungal composition in airborne wind currents and soil fungal communities across a 40 000 km2 regional landscape to determine if dispersal limitation or abiotic factors were structuring soil fungal communities. Over this landscape, neither airborne nor soil fungal communities exhibited compositional differences due to geographic distance. Airborne fungal communities shifted temporally while soil fungal communities were correlated with abiotic parameters. These patterns suggest that environmental filtering may have the largest influence on fungal regional community assembly in soils, especially for aerially dispersed fungal taxa. Furthermore, we found evidence that dispersal of fungal spores differs between fungal taxa and can be both a stochastic and deterministic process. The spatial range of soil fungal taxa was correlated with their average regional abundance across all sites, which may imply stochastic dispersal mechanisms. Nevertheless, spore volume was also negatively correlated with spatial range for some species. Smaller volume spores may be adapted to long-range dispersal, or establishment, suggesting that deterministic fungal traits may also influence fungal distributions. Fungal life-history traits may influence their distributions as well. Hypogeous fungal taxa exhibited high local abundance, but small spatial ranges, while epigeous fungal taxa had lower local abundance, but larger spatial ranges. This study is the first, to our knowledge, to directly sample air dispersal and soil fungal communities simultaneously across a regional landscape. We provide some of the first evidence that soil fungal communities are mostly assembled through environmental filtering and experience little dispersal limitation.

High-elevation ecosystems are expected to be particularly sensitive to climate warming because cold temperatures constrain biological processes. Deeper understanding of the consequences of climate change will come from studies that consider not only the direct effects of temperature on individual species, but also the indirect effects of altered species interactions. Here we show that 20 years of experimental warming has changed the species composition of graminoid (grass and sedge) assemblages in a subalpine meadow of the Rocky Mountains, USA, by increasing the frequency of sedges and reducing the frequency of grasses. Because sedges typically have weak interactions with mycorrhizal fungi relative to grasses, lowered abundances of arbuscular mycorrhizal (AM) fungi or other root-inhabiting fungi could underlie warming-induced shifts in plant species composition. However, warming increased root colonization by AM fungi for two grass species, possibly because AM fungi can enhance plant water uptake when soils are dried by experimental warming. Warming had no effect on AM fungal colonization of three other graminoids. Increased AM fungal colonization of the dominant shrub Artemisia tridentata provided further grounds for rejecting the hypothesis that reduced AM fungi caused the shift from grasses to sedges. Non-AM fungi (including dark septate endophytes) also showed general increases with warming. Our results demonstrate that lumping grasses and sedges when characterizing plant community responses can mask significant shifts in the responses of primary producers, and their symbiotic fungi, to climate change.