Long-term biocrust responses to wildfires in Washington, USA.

PREMISE: Dryland ecosystems in the western United States are affected by invasive species, wildfires, livestock grazing, and climate change in ways that are difficult to distinguish. Biocrusts perform important ecological roles in these systems and are sensitive to all of these pressures. METHODS: We revisited a Washington, USA, site sampled for biocrusts in 1999 to focus on effects of exotic annual grass invasion and wildfires in the absence of livestock grazing. We examined changes between 1999 and 2020 using a Bayesian directed acyclic graph (DAG) to interpret direct and indirect causal impacts of wildfire on perennial bunchgrasses, exotic annual grasses, and biocrusts. RESULTS: Between 1999 and 2020, exotic annual grass cover increased in all plots and in unburned plots by 16% and 18%, respectively, bunchgrass cover decreased by 21% and 25%, and biocrust cover decreased by 8.9% and 9.8%. Our DAG suggested that decreases in bunchgrass increased exotic annual grass, which reduced biocrust cover. Wildfires did not directly influence changes in bunchgrass, exotic annual grass, or biocrust cover. Areas dominated by exotic annual grass had less abundant and diverse biocrusts than areas with less exotic annual grass. CONCLUSIONS: Biocrust community changes were more strongly related to increasing exotic annual grasses than to wildfires. Changes may relate to other soil disturbances or broad-scale changes in climate or air quality. The minimal influence of wildfire on exotic annual grass and biocrusts suggests that apparent negative impacts of wildfire at other sites may be due to exacerbation by livestock grazing or other surface disturbance.

Grazing disturbance promotes exotic annual grasses by degrading soil biocrust communities.

Exotic invasive plants threaten ecosystem integrity, and their success depends on a combination of abiotic factors, disturbances, and interactions with existing communities. In dryland ecosystems, soil biocrusts (communities of lichens, bryophytes, and microorganisms) can limit favorable microsites needed for invasive species establishment, but the relative importance of biocrusts for landscape-scale invasion patterns remains poorly understood. We examine effects of livestock grazing in habitats at high risk for invasion to test the hypothesis that disturbance indirectly favors exotic annual grasses by reducing biocrust cover. We present some of the first evidence that biocrusts increase site resistance to invasion at a landscape scale and mediate the effects of disturbance. Biocrust species richness, which is reduced by livestock grazing, also appears to promote native perennial grasses. Short mosses, as a functional group, appear to be particularly valuable for preventing invasion by exotic annual grasses. Our study suggests that maintaining biocrust communities with high cover, species richness, and cover of short mosses can increase resistance to invasion. These results highlight the potential of soil surface communities to mediate invasion dynamics and suggest promising avenues for restoration in dryland ecosystems.

Assessing Ecological Risks from Atmospheric Deposition of Nitrogen and Sulfur to US Forests Using Epiphytic Macrolichens.

Critical loads of atmospheric deposition help decision-makers identify levels of air pollution harmful to ecosystem components. But when critical loads are exceeded, how can the accompanying ecological risk be quantified? We use a 90% quantile regression to model relationships between nitrogen and sulfur deposition and epiphytic macrolichens, focusing on responses of concern to managers of US forests: Species richness and abundance and diversity of functional groups with integral ecological roles. Analyses utilized national-scale lichen survey data, sensitivity ratings, and modeled deposition and climate data. We propose 20, 50, and 80% declines in these responses as cut-offs for low, moderate, and high ecological risk from deposition. Critical loads (low risk cut-off) for total species richness, sensitive species richness, forage lichen abundance and cyanolichen abundance, respectively, were 3.5, 3.1, 1.9, and 1.3 kg N and 6.0, 2.5, 2.6, and 2.3 kg S ha-1 yr-1. High environmental risk (80% decline), excluding total species richness, occurred at 14.8, 10.4, and 6.6 kg N and 14.1, 13, and 11 kg S ha-1 yr-1. These risks were further characterized in relation to geography, species of conservation concern, number of species affected, recovery timeframes, climate, and effects on interdependent biota, nutrient cycling, and ecosystem services.

Altered fire regimes cause long-term lichen diversity losses.

Many global ecosystems have undergone shifts in fire regimes in recent decades, such as changes in fire size, frequency, and/or severity. Recent research shows that increases in fire size, frequency, and severity can lead to long-persisting deforestation, but the consequences of shifting fire regimes for biodiversity of other vegetative organisms (such as understory plants, fungi, and lichens) remain poorly understood. Understanding lichen responses to wildfire is particularly important because lichens play crucial roles in nutrient cycling and supporting wildlife in many ecosystems. Lichen responses to fire have been little studied, and most previous research has been limited to small geographic areas (e.g. studies of a single fire), making it difficult to establish generalizable patterns. To investigate long-term effects of fire severity on lichen communities, we sampled epiphytic lichen communities in 104 study plots across California's greater Sierra Nevada region in areas that burned in five wildfires, ranging from 4 to 16 years prior to sampling. The conifer forest ecosystems we studied have undergone a notable increase in fire severity in recent decades, and we sample across the full gradient of fire severity to infer how shifting fire regimes may influence landscape-level biodiversity. We find that low-severity fire has little to no effect on lichen communities. Areas that burned at moderate and high severities, however, have significantly and progressively lower lichen richness and abundance. Importantly, we observe very little postfire lichen recolonization on burned substrates even more than 15 years after fire. Our multivariate model suggests that the hotter, drier microclimates that occur after fire removes forest canopies may prevent lichen reestablishment, meaning that lichens are not likely to recolonize until mature trees regenerate. These findings suggest that altered fire regimes may cause broad and long-persisting landscape-scale biodiversity losses that could ultimately impact multiple trophic levels.

Plant diversity enhances moth diversity in an intensive forest management experiment.

Intensive forest management (IFM) promises to help satisfy increasing global demand for wood but may come at the cost of local reductions to forest biodiversity. IFM often reduces early seral plant diversity as a result of efforts to eliminate plant competition with crop trees. If diversity is a function of bottom-up drivers, theory predicts that specialists at lower trophic levels (e.g., insect herbivores) should be particularly sensitive to reductions in plant diversity. We conducted a stand-level experiment to test bottom-up controls on moth community structure, as mediated by degrees of forest management intensity. Using a dataset of 12,003 moths representing 316 moth species, moth richness decreased only slightly, if at all, as herbicide intensity increased (P = 0.062); the moderate treatment, which is most commonly applied in the northwestern USA, was estimated to have 4.72 (±2.14 SE, P = 0.039) fewer species than the control. Structural equation modeling revealed strong support for an effect of herbicide on plant abundance, which influenced plant species richness and subsequently moth species richness. Moth species richness was associated with plant species richness and followed a power law function (z = 0.42, P = 0.006), which is surprisingly consistent with a recent large-scale experiment in agricultural systems, and provides support for bottom-up drivers of moth community structure. Moth abundance was not influenced by the direct effects of silvicultural herbicide treatments. Site-level effects and variation in pre-harvest vegetation communities resulted in residual broadleaf and herbaceous vegetation in even the most intensive treatment. Even at low densities, these residual deciduous and herbaceous plants supported higher than expected moth abundance and richness. We conclude that forest management practices that retain early seral vegetation diversity are the most likely to conserve moth communities.

Native and exotic plant cover vary inversely along a climate gradient 11 years following stand-replacing wildfire in a dry coniferous forest, Oregon, USA.

Community re-assembly following future disturbances will often occur under warmer and more moisture-limited conditions than when current communities assembled. Because the establishment stage is regularly the most sensitive to climate and competition, the trajectory of recovery from disturbance in a changing environment is uncertain, but has important consequences for future ecosystem functioning. To better understand how ongoing warming and rising moisture limitation may affect recovery, we studied native and exotic plant composition 11 years following complete stand-replacing wildfire in a dry coniferous forest spanning a large gradient in climatic moisture deficit (CMD) from warm and dry low elevation sites to relatively cool and moist higher elevations sites. We then projected future precipitation, temperature and CMD at our study locations for four scenarios selected to encompass a broad range of possible future conditions for the region. Native perennials dominated relatively cool and moist sites 11 years after wildfire, but were very sparse at the warmest and driest (high CMD) sites, particularly when combined with high topographic sun exposure. In contrast, exotic species (primarily annual grasses) were dominant or co-dominant at the warmest and driest sites, especially with high topographic sun exposure. All future scenarios projected increasing temperature and CMD in coming decades (e.g., from 4.5% to 29.5% higher CMD by the 2080's compared to the 1971-2000 average), even in scenarios where growing season (May-September) precipitation increased. These results suggest increasing temperatures and moisture limitation could facilitate longer term (over a decade) transitions toward exotic-dominated communities after severe wildfire when a suitable exotic seed source is present.