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Self‐reinforcing differences in fire frequency help closed‐canopy forests, which resist fire, and open woodlands, which naturally burn often, to co‐occur stably at landscape scales. Forest tree seedlings, which could otherwise encroach and overgrow woodlands, are killed by regular fire, yet fire has other effects that may also influence these feedbacks. In particular, many forest trees require symbiotic ectomycorrhizal fungi in order to establish. By restructuring soil fungal communities, fire might affect the availability of symbionts or the potential for symbiont sharing between encroaching trees and woodland vegetation.
To investigate this possibility, we performed a soil bioassay experiment using inoculum from burned and unburned oak woodlands and Douglas‐fir forests. We examined how fire, ecosystem type, and neighboring heterospecific seedlings affect fungal root community assembly of Douglas‐firs and oaks. We asked whether heterospecific seedlings facilitated fungal colonization of seedling roots in non‐native soil, and if so, whether fire influenced this interaction.
External fungal colonization of oak roots was more influenced by fire and ecosystem type than by the presence of a Douglas‐fir, and oaks increased the likelihood that Douglas‐fir roots would be colonized by fungi in oak woodland soil. Yet, fire increased colonization of Douglas‐fir in oak soil, diminishing the otherwise crucial role played by oak facilitation. Fire also strengthened the positive effect of Douglas‐firs on oak root‐associated fungal diversity in Douglas‐fir forest soil.
Prior work shows that fire supports woodland ecosystems by stemming recruitment of encroaching seedlings. Here, we find evidence that it may contrastingly reduce fungal limitation of invasive seedling growth and establishment, otherwise relieved only by facilitation. Future work can investigate how these opposing effects might contribute to the net impact of changes in fire regime on landcover stability.
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Efforts to predict the responses of soil fungal communities to climate change are hindered by limited information on how fungal niches are distributed across environmental hyperspace. We predict the climate sensitivity of North American soil fungal assemblage composition by modelling the ecological niches of several thousand fungal species.
One hundred and thirteen sites in the United States and Canada spanning all biomes except tropical rain forest.
Major Taxa Studied
We combine internal transcribed spacer (ITS) sequences from two continental‐scale sampling networks in North America and cluster them into operational taxonomic units (OTUs) at 97% similarity. Using climate and soil data, we fit ecological niche models (ENMs) based on logistic ridge regression for all OTUs present in at least 10 sites (
n= 8597). To describe the compositional turnover of soil fungal assemblages over climatic gradients, we introduce a novel niche‐based metric of climate sensitivity, the Sørensen climate sensitivity index. Finally, we map climate sensitivity across North America. Results
ENMs have a mean out‐of‐sample predictive accuracy of 73.8%, with temperature variables being strong predictors of fungal distributions. Soil fungal climate niches clump together across environmental space, which suggests common physiological limits and predicts abrupt changes in composition with respect to changes in climate. Soil fungi in North American climates are more likely to be limited by cold and dry conditions than by warm and wet conditions, and ectomycorrhizal fungi generally tolerate colder temperatures than saprotrophic fungi. Sørensen climate sensitivity exhibits a multimodal distribution across environmental space, with a peak in climates corresponding to boreal forests.
The boreal forest occupies an especially precarious region of environmental space for the composition of soil fungal assemblages in North America, as even small degrees of warming could trigger large compositional changes characterized mainly by an influx of warm‐adapted species.
Seedling recruitment can be strongly affected by the composition of nearby plant species. At the neighborhood scale (on the order of tens of meters), adult conspecifics can modify soil chemistry and the presence of host microbes (pathogens and mutualists) across their combined canopy area or rooting zones. At local or small spatial scales (on the order of one to few meters), conspecific seed or seedling density can influence the strength of intraspecific light and resource competition and also modify the density‐dependent spread of natural enemies such as pathogens or invertebrate predators. Intrinsic correlation between proximity to adult conspecifics (i.e., recruitment neighborhood) and local seedling density, arising from dispersal, makes it difficult to separate the independent and interactive factors that contribute to recruitment success. Here, we present a field experiment in which we manipulated both the recruitment neighborhood and seedling density to explore how they interact to influence the growth and survival of
Dryobalanops aromatica, a dominant ectomycorrhizal tree species in a Bornean tropical rainforest. First, we found that both local seedling density and recruitment neighborhood had effects on performance of D. aromaticaseedlings, though the nature of these impacts varied between growth and survival. Second, we did not find strong evidence that the effect of density on seedling survival is dependent on the presence of conspecific adult trees. However, accumulation of mutualistic fungi beneath conspecifics adults does facilitate establishment of D. aromaticaseedlings. In total, our results suggest that recruitment near adult conspecifics was not associated with a performance cost and may have weakly benefitted recruiting seedlings. Positive effects of conspecifics may be a factor facilitating the regional hyperabundance of this species. Synthesis: Our results provide support for the idea that dominant species in diverse forests may escape the localized recruitment suppression that limits abundance in rarer species.
Decomposition has historically been considered a function of climate and substrate but new research highlights the significant role of specific micro‐organisms and their interactions. In particular, wood decay is better predicted by variation in fungal communities than in climate. Multiple links exist: interspecific competition slows decomposition in more diverse fungal communities, whereas trait variation between different communities also affects process rates. Here, we paired field and laboratory experiments using a dispersal gradient at a forest‐shrubland ecotone to examine how fungi affect wood decomposition across scales. We observed that while fungal communities closer to forests were capable of faster decomposition, wood containing diverse fungal communities decomposed more slowly, independent of location. Dispersal‐driven stochasticity in small‐scale community assembly was nested within large‐scale turnover in the regional species pool, decoupling the two patterns. We thus find multiple distinct links between microbes and ecosystem function that manifest across different spatial scales.
Aim Ectomycorrhizal fungi (ECMF) are partners in a globally distributed tree symbiosis implicated in most major ecosystem functions. However, resilience of ECMF to future climates is uncertain. We forecast these changes over the extent of North American Pinaceae forests. Location About 68 sites from North American Pinaceae forests ranging from Florida to Ontario in the east and southern California to Alaska in the west. Taxon Ectomycorrhizal fungi (Asco‐ and Basidiomycetes). Methods We characterized ECMF communities at each site using molecular methods and modelled climatic drivers of diversity and community composition with general additive, generalized dissimilarity models and Threshold Indicator Taxa ANalysis (TITAN). Next, we projected our models across the extent of North American Pinaceae forests and forecast ECMF responses to climate changes in these forests over the next 50 years. Results We predict median declines in ECMF species richness as high as 26% in Pinaceae forests throughout a climate zone comprising more than 3.5 million square kilometres of North America (an area twice that of Alaska state). Mitigation of greenhouse gas emissions can reduce these declines, but not prevent them. The existence of multiple diversity optima along climate gradients suggest regionally divergent trajectories for North American ECMF, which is corroborated by corresponding ECMF community thresholds identified in TITAN models. Warming of forests along the boreal–temperate ecotone results in projected ECMF species loss and declines in the relative abundance of long‐distance foraging ECMF species, whereas warming of eastern temperate forests has the opposite effect. Main Conclusions Our results reveal potentially unavoidable ECMF species‐losses over the next 50 years, which is likely to have profound (if yet unclear) effects on ECMF‐associated biogeochemical cycles.more » « less
Ectomycorrhizal symbiosis is essential for the nutrition of most temperate forest trees and helps regulate the movement of carbon (C) and nitrogen (N) through forested ecosystems. The factors governing the exchange of plant C for fungal N, however, remain obscure.
Because competition and soil resources may influence ectomycorrhizal resource movement, we performed a 10‐month split‐root microcosm study using
Pinus muricataseedlings with Thelephora terrestris, Suillus pungens, or no ectomycorrhizal fungus, under two N concentrations in artificial soil. Fungi competed directly with roots and indirectly with each other. We used stable isotope enrichment to track plant photosynthate and fungal N.
T. terrestris, plants received N commensurate with the C given to their fungal partners. Thelephora terrestriswas a superior mutualist under high‐N conditions. For S. pungens, plant C and fungal N exchange were not coupled. However, in low‐N conditions, plants preferentially allocated C to S. pungensrather than T. terrestris.
Our results suggest that ectomycorrhizal resource transfer depends on competitive and nutritional context. Plants can exchange C for fungal N, but coupling of these resources can depend on the fungal species and soil N. Understanding the diversity of fungal strategies, and how they change with environmental context, reveals mechanisms driving this important symbiosis.
Wildfire affects our planet's biogeochemistry both by burning biomass and by driving changes in ecological communities and landcover. Some plants and ecosystem types are threatened by increasing fire pressure while others respond positively to fire, growing in local and regional abundance when it occurs regularly. However, quantifying total ecosystem response to fire demands consideration of impacts not only on aboveground vegetation, but also on soil microbes like fungi, which influence decomposition and nutrient mineralization. If fire‐resistant soil fungal communities co‐occur with similarly adapted plants, these above‐ and belowground ecosystem components should shift and recover in relative synchrony after burning. If not, fire might decouple ecosystem processes governed by these different communities, affecting total functioning. Here, we use a natural experiment to test whether fire‐dependent ecosystems host unique, fire‐resistant fungal communities. We surveyed burned and unburned areas across two California ecosystem types with differing fire ecologies in the immediate aftermath of a wildfire, finding that the soil fungal communities of fire‐dependent oak woodlands differ from those of neighbouring mixed evergreen forests. We discovered furthermore that the latter are more strongly altered compositionally by fire than the former, suggesting that differences in fungal community structure support divergent community responses to fire across ecosystems. Our results thus indicate that fire‐dependent ecosystems may host fire‐resistant fungal communities.