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.

Erosion of tropical bird diversity over a century is influenced by abundance, diet and subtle climatic tolerances.

Human alteration of landscapes leads to attrition of biodiversity. Recommendations for maximizing retention of species richness typically focus on protection and preservation of large habitat patches. Despite a century of protection from human disturbance, 27% of the 228 bird species initially detected on Barro Colorado Island (BCI), Panama, a large hilltop forest fragment isolated by waters of Gatun Lake, are now absent. Lost species were more likely to be initially uncommon and terrestrial insectivores. Analyses of the regional avifauna, exhaustively inventoried and mapped across 24 subregions, identified strong geographical discontinuities in species distributions associated with a steep transisthmian rainfall gradient. Having lost mostly species preferring humid forests, the BCI species assemblage continues to shift from one originally typical of wetter forests toward one now resembling bird communities in drier forests. Even when habitat remnants are large and protected for 100 years, altered habitat characteristics resulting from isolation produce non-random loss of species linked with their commonness, dietary preferences and subtle climatic sensitivities.

Novel climates reverse carbon uptake of atmospherically dependent epiphytes: Climatic constraints on the iconic boreal forest lichen Evernia mesomorpha.

PREMISE OF THE STUDY: Changing climates are expected to affect the abundance and distribution of global vegetation, especially plants and lichens with an epiphytic lifestyle and direct exposure to atmospheric variation. The study of epiphytes could improve understanding of biological responses to climatic changes, but only if the conditions that elicit physiological performance changes are clearly defined. METHODS: We evaluated individual growth performance of the epiphytic lichen Evernia mesomorpha, an iconic boreal forest indicator species, in the first year of a decade-long experiment featuring whole-ecosystem warming and drying. Field experimental enclosures were located near the southern edge of the species' range. KEY RESULTS: Mean annual biomass growth of Evernia significantly declined 6 percentage points for every +1°C of experimental warming after accounting for interactions with atmospheric drying. Mean annual biomass growth was 14% in ambient treatments, 2% in unheated control treatments, and -9% to -19% (decreases) in energy-added treatments ranging from +2.25 to +9.00°C above ambient temperatures. Warming-induced biomass losses among persistent individuals were suggestive evidence of an extinction debt that could precede further local mortality events. CONCLUSIONS: Changing patterns of warming and drying would decrease or reverse Evernia growth at its southern range margins, with potential consequences for the maintenance of local and regional populations. Negative carbon balances among persisting individuals could physiologically commit these epiphytes to local extinction. Our findings illuminate the processes underlying local extinctions of epiphytes and suggest broader consequences for range shrinkage if dispersal and recruitment rates cannot keep pace.

Limitations of Species Delimitation Based on Phylogenetic Analyses: A Case Study in the Hypogymnia hypotrypa Group (Parmeliaceae, Ascomycota).

Delimiting species boundaries among closely related lineages often requires a range of independent data sets and analytical approaches. Similar to other organismal groups, robust species circumscriptions in fungi are increasingly investigated within an empirical framework. Here we attempt to delimit species boundaries in a closely related clade of lichen-forming fungi endemic to Asia, the Hypogymnia hypotrypa group (Parmeliaceae). In the current classification, the Hypogymnia hypotrypa group includes two species: H. hypotrypa and H. flavida, which are separated based on distinctive reproductive modes, the former producing soredia but absent in the latter. We reexamined the relationship between these two species using phenotypic characters and molecular sequence data (ITS, GPD, and MCM7 sequences) to address species boundaries in this group. In addition to morphological investigations, we used Bayesian clustering to identify potential genetic groups in the H. hypotrypa/H. flavida clade. We also used a variety of empirical, sequence-based species delimitation approaches, including: the "Automatic Barcode Gap Discovery" (ABGD), the Poisson tree process model (PTP), the General Mixed Yule Coalescent (GMYC), and the multispecies coalescent approach BPP. Different species delimitation scenarios were compared using Bayes factors delimitation analysis, in addition to comparisons of pairwise genetic distances, pairwise fixation indices (FST). The majority of the species delimitation analyses implemented in this study failed to support H. hypotrypa and H. flavida as distinct lineages, as did the Bayesian clustering analysis. However, strong support for the evolutionary independence of H. hypotrypa and H. flavida was inferred using BPP and further supported by Bayes factor delimitation. In spite of rigorous morphological comparisons and a wide range of sequence-based approaches to delimit species, species boundaries in the H. hypotrypa group remain uncertain. This study reveals the potential limitations of relying on distinct reproductive strategies as diagnostic taxonomic characters for Hypogymnia and also the challenges of using popular sequence-based species delimitation methods in groups with recent diversification histories.

Non-Native Plant Invasion along Elevation and Canopy Closure Gradients in a Middle Rocky Mountain Ecosystem.

Mountain environments are currently among the ecosystems least invaded by non-native species; however, mountains are increasingly under threat of non-native plant invasion. The slow pace of exotic plant invasions in mountain ecosystems is likely due to a combination of low anthropogenic disturbances, low propagule supply, and extreme/steep environmental gradients. The importance of any one of these factors is debated and likely ecosystem dependent. We evaluated the importance of various correlates of plant invasions in the Wallowa Mountain Range of northeastern Oregon and explored whether non-native species distributions differed from native species along an elevation gradient. Vascular plant communities were sampled in summer 2012 along three mountain roads. Transects (n = 20) were evenly stratified by elevation (~70 m intervals) along each road. Vascular plant species abundances and environmental parameters were measured. We used indicator species analysis to identify habitat affinities for non-native species. Plots were ordinated in species space, joint plots and non-parametric multiplicative regression were used to relate species and community variation to environmental variables. Non-native species richness decreased continuously with increasing elevation. In contrast, native species richness displayed a unimodal distribution with maximum richness occurring at mid-elevations. Species composition was strongly related to elevation and canopy openness. Overlays of trait and environmental factors onto non-metric multidimensional ordinations identified the montane-subalpine community transition and over-story canopy closure exceeding 60% as potential barriers to non-native species establishment. Unlike native species, non-native species showed little evidence for high-elevation or closed-canopy specialization. These data suggest that non-native plants currently found in the Wallowa Mountains are dependent on open canopies and disturbance for establishment in low and mid elevations. Current management objectives including restoration to more open canopies in dry Rocky Mountain forests, may increase immigration pressure of non-native plants from lower elevations into the montane and subalpine zones.

Evolution of complex symbiotic relationships in a morphologically derived family of lichen-forming fungi.

We studied the evolutionary history of the Parmeliaceae (Lecanoromycetes, Ascomycota), one of the largest families of lichen-forming fungi with complex and variable morphologies, also including several lichenicolous fungi. We assembled a six-locus data set including nuclear, mitochondrial and low-copy protein-coding genes from 293 operational taxonomic units (OTUs). The lichenicolous lifestyle originated independently three times in lichenized ancestors within Parmeliaceae, and a new generic name is introduced for one of these fungi. In all cases, the independent origins occurred c. 24 million yr ago. Further, we show that the Paleocene, Eocene and Oligocene were key periods when diversification of major lineages within Parmeliaceae occurred, with subsequent radiations occurring primarily during the Oligocene and Miocene. Our phylogenetic hypothesis supports the independent origin of lichenicolous fungi associated with climatic shifts at the Oligocene-Miocene boundary. Moreover, diversification bursts at different times may be crucial factors driving the diversification of Parmeliaceae. Additionally, our study provides novel insight into evolutionary relationships in this large and diverse family of lichen-forming ascomycetes.

A single phosphorus treatment doubles growth of cyanobacterial lichen transplants.

Lichens are reputedly slow growing and become unhealthy or die in response to supplements of the usual limiting resources, such as water and nitrogen. We found, however, that the tripartite cyanobacterial lichen Lobaria pulmonaria doubled in annual biomass growth after a single 20-minute immersion in a phosphorus solution (K2HPO4), as compared to controls receiving no supplemental phosphorus. This stimulation of cyanolichens by phosphorus has direct relevance to community and population ecology of lichens, including improving models of lichen performance in relation to air quality, improving forest management practices affecting old-growth associated cyanolichens, and understanding the distribution and abundance of cyanolichens on the landscape. Phosphorus may be as important a stimulant to cyanobacterial-rich lichen communities as it is to cyanobacteria in aquatic ecosystems.