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Lichens are combinations of two symbiotic organisms, a green alga or cyanobacterium and a fungus. They grow in nearly all terrestrial ecosystems and survive in habitats, which are very dry or cold, or too poor in nutrients to maintain vegetation growth. Because lichens grow on visible surfaces and exhibit spectral properties, which are clearly different from, for example, vegetation, it is possible to distinguish them in remote sensing data. In this first systematic review article on remote sensing of lichens, we analyze and summarize which lichen species or genera, and in which habitats and geographical regions, have been remotely sensed, and which remote sensing or spectroscopic technologies have been used. We found that laboratory or in situ measured spectra of over 70 lichen species have been reported to date. We show that studies on remote sensing of lichens fall under seven broad themes: (1) collection of lichen spectra for quantification of lichen species or characteristics, (2) pollution monitoring with lichens as ecological indicators, (3) geological and lithological mapping, (4) desert and dryland monitoring, (5) animal habitat monitoring, (6) land cover or vegetation mapping, and (7) surface energy budget modeling. [ABSTRACT FROM AUTHOR]

Abstract Lichens are combinations of two symbiotic organisms, a green alga or cyanobacterium and a fungus. They grow in nearly all terrestrial ecosystems and survive in habitats, which are very dry or cold, or too poor in nutrients to maintain vegetation growth. Because lichens grow on visible surfaces and exhibit spectral properties, which are clearly different from, for example, vegetation, it is possible to distinguish them in remote sensing data. In this first systematic review article on remote sensing of lichens, we analyze and summarize which lichen species or genera, and in which habitats and geographical regions, have been remotely sensed, and which remote sensing or spectroscopic technologies have been used. We found that laboratory or in situ measured spectra of over 70 lichen species have been reported to date. We show that studies on remote sensing of lichens fall under seven broad themes: (1) collection of lichen spectra for quantification of lichen species or characteristics, (2) pollution monitoring with lichens as ecological indicators, (3) geological and lithological mapping, (4) desert and dryland monitoring, (5) animal habitat monitoring, (6) land cover or vegetation mapping, and (7) surface energy budget modeling.

Dryland ecosystems can be constrained by low soil fertility. Within drylands, the soil nutrient and organic carbon (C) cycling that does occur is often mediated by soil surface communities known as biological soil crusts (biocrusts), which cycle C and nutrients in the top ca. 0–2 cm of soil. However, the degree to which biocrusts are influencing soil fertility and biogeochemical cycling in deeper, subsurface mineral soils is unclear. The movement of dissolved resources from biocrusts to deeper soil layers in leachate may be one of the main mechanisms through which biocrust fertility is transferred downward towards deeper microbial communities and plant roots occurring within mineral soil. Here we examined the role of biocrust leachate in contributing to subsurface nutrient and soluble C pools and subsurface microbial cycling. We collected biocrusts from three biocrust successional stages and explored resource pools in situ at multiple soil depths, while collecting leachate and measuring nutrient and organic C concentrations and metabolite composition from each successional stage in the laboratory. After four leachate collections, we conducted an incubation of mineral soil collected from below each biocrust successional stage to measure heterotrophic microbial CO2 flux and biomass. Overall, our findings observed that the degree of nutrient and C connectivity between biocrusts and the sub-crust mineral soil depended on the biocrust successional stage and the element being considered, and the influence of biocrust successional stage on mineral soil CO2 flux is likely related to long-term resource build up. Together, our results suggest that the influence of biocrust leachate on subsurface mineral soil is complex and context dependent, but, over longer time periods and at later successional stages, can have measurable effects on dryland soil biogeochemical cycling with feedbacks to resource availability and CO2 flux.

The ranges of many species globally have already shifted to maintain climatic equilibrium in the face of climate change.Biocrusts—soil surface dwelling communities of lichens, bryophytes and microbes—play important functional roles in many ecosystems, particularly in drylands. Compared to better studied animal and plant taxa, dryland biocrusts have different establishment requirements and have never been assessed for historical range shifts.Here, we revisited the sites (N = 204) of a 25‐year‐old biocrust survey across a large area (400,000 km2) of drylands in south‐eastern Australia. We used quadratic models to quantify changes in the climate niches of 15 lichen, eight moss and five liverwort taxa, as well as biocrust cover and richness.Our models showed that the observed climatic niches of most taxa have become hotter and drier in the past quarter century, yet the responses of the vast majority of taxa are consistent with remaining in the same geographic space. A similar pattern was observed at the community level, where the peak of biocrust cover and richness now occurs in a hotter, drier environment. Notable exceptions were the liverwort Riccia lamellosa and lichens in the genera Cladonia and Xanthoparmelia, which showed signs of contraction at their arid range edges.Unlike more mobile taxa, most biocrust species have yet to shift geographically and may already be lagging behind the pace of climate change. One explanation for the mortality lag is that long‐term climate variability in the system is extensive, which may have selected for the ability to withstand multi‐year warm periods as long as there is an eventual return to milder conditions. However, no forecasts of future climate include a return to milder conditions, suggesting there will be an eventual loss of ecosystem multifunctionality at the contracting front. Expansion lags are most likely due to delays in the mortality of competing vascular plants.Synthesis: Our study provides a valuable contribution to the knowledge of range shifts in understudied taxa and highlights a future need to promote the expansion of biocrusts to maintain the provision of ecosystem functions and services across their range. [ABSTRACT FROM AUTHOR]

Aim: Lichens and mosses play important functional roles in all terrestrial ecosystems, particularly in tundra and drylands. As with all taxa, to maintain their current niche in a changing climate, lichens and mosses will have to migrate. However, there are no published estimates of future habitat suitability or necessary rates of migration for members of these groups at the global scale. Taxon: Lichens and mosses. Location: Global. Methods: Using global occurrence data, we conducted ensemble distribution models in the 'biomod2' R package, parameterised with a range of climatic, land use and soil variables, to estimate current and future (2100) habitat suitability in 16 abundant species of lichen and moss. Results: Without considering dispersal limitation, suitable area was forecast to expand for eight species and decline for four species. For species with predominantly boreo‐arctic distributions, suitable area typically declined at the temperate range edge and expanded across the High Arctic. Future suitable area available to dryland‐adapted species generally declined overall, likely relating to the desiccation‐tolerant physiology of lichens and mosses. The average migration rates required for species to disperse into new suitable habitat ranged from 1.7 (Placidium squamulosum) to 9.0 km year−1 (Syntrichia ruralis), although most species will need to migrate >16 km year−1 to completely fill their potential future suitable habitat. Main Conclusions: For mosses and lichens, as with all species, migration will be an important part of the adjustment to a warmer climate, but realisation of these potential migrations will require both rare dispersal events and habitat that is suitable in non‐climatic dimensions. Current evidence on dispersal in these groups suggests that these geographical shifts may be unlikely to be realised without intervention, especially in landscapes that are highly modified by humans. [ABSTRACT FROM AUTHOR]

Lichens are one of the most iconic and ubiquitous symbioses known, widely valued as indicators of environmental quality and, more recently, climate change. Our understanding of lichen responses to climate has greatly expanded in recent decades, but some biases and constraints have shaped our present knowledge. In this review we focus on lichen ecophysiology as a key to predicting responses to present and future climates, highlighting recent advances and remaining challenges. Lichen ecophysiology is best understood through complementary whole‐thallus and within‐thallus scales. Water content and form (vapor or liquid) are central to whole‐thallus perspectives, making vapor pressure differential (VPD) a particularly informative environmental driver. Responses to water content are further modulated by photobiont physiology and whole‐thallus phenotype, providing clear links to a functional trait framework. However, this thallus‐level perspective is incomplete without also considering within‐thallus dynamics, such as changing proportions or even identities of symbionts in response to climate, nutrients, and other stressors. These changes provide pathways for acclimation, but their understanding is currently limited by large gaps in our understanding of carbon allocation and symbiont turnover in lichens. Lastly, the study of lichen physiology has mainly prioritized larger lichens at high latitudes, producing valuable insights but underrepresenting the range of lichenized lineages and ecologies. Key areas for future work include improving geographic and phylogenetic coverage, greater emphasis on VPD as a climatic factor, advances in the study of carbon allocation and symbiont turnover, and the incorporation of physiological theory and functional traits in our predictive models. [ABSTRACT FROM AUTHOR]

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Multiple ecological drivers, along with forest age, determine the species composition of boreal forest ecosystems. However, the role of age in successional changes in forests cannot be understood without taking site conditions, the disturbance regime and forest structure into account. In this study, we ask two research questions: 1. What is the relationship between forest age and overall species composition in older near-natural spruce forests, i.e. forests of age beyond harvest maturity? 2. Do species associated with different forest habitats respond similarly to variation in forest age? Data were collected in 257 Norway spruce dominated 0.25 ha plots from three study areas in Southeastern and Central Norway. Species inventories were conducted for lichens and bryophytes on trees and rocks, vascular plants on the forest floor, and for deadwood-associated bryophytes and polypore fungi. Although NMDS ordination analyses of the total species composition identified a main axis related to the age of the oldest trees in two of the study areas, variation partitioning analyses showed that age explained a small fraction of variation of the species composition compared to site conditions, logging history, forest structure, and differences between the sites in all habitats. The unique variation explained by forest age species was, however, significant for all habitats. The fraction of variation in species composition explained by forest age was the largest for lichens and bryophytes on trees, and for deadwood-associated bryophytes and polypore fungi. Our results suggest that practical mapping of near-natural forests for management purposes inventories should include site conditions, forest structure and between site differences in addition to forest age. [Display omitted] • Site conditions and forest structure explained most of the species composition. • There is unique variation in the species assemblages explained by age. • Age was most influential for epiphytes, deadwood-associated bryophytes and polypores. • Representative mapping of near-natural spruce forest should include age variables. [ABSTRACT FROM AUTHOR]

Lichens and bryophytes are ubiquitous in terrestrial habitats, sensitive to environmental changes, and they contribute to ecosystem functions and biogeochemical cycles. Intact forest ecosystems host distinct biotic communities that are associated with microhabitat and structural diversity at stand and landscape scales. In the boreal region of eastern Canada, insect outbreaks and clearcutting (the dominant timber harvesting method) are common disturbances. However, while insect outbreaks from native species can maintain ecological continuity, clearcutting disrupts it and homogenizes forest structure. We compared lichen and bryophyte communities between old-growth forest stands and 50 years-old stands regenerated from clearcuts and insect outbreaks. From these communities, liverwort and deadwood specialist species richness was higher in insect outbreak and old-growth than in clearcut forest stands. Stand type explained 10.3 % and 7.0 % of the variability observed in liverwort and deadwood specialist community composition, respectively. Clearcut stands were responsible for most of this variability since communities had less unique species and lacked some of the most common species found in old-growth and insect outbreak stands. Indeed, many species associated with deadwood and ecological continuity appeared unable to recolonize forest stands between planned intervals of clearcut rotations (i.e. ~50 to 60 years in our study area). However, deadwood volume did not differ between stand types and could not explain any part of the observed variability in the composition of communities. We suggest that forests regenerating from insect outbreaks are a suitable alternative for biological conservation in regions where old-growth forests are rare, especially for liverworts and deadwood specialists. • Liverworts and deadwood specialists are the most sensitive to clearcuts. • Insect outbreak and old-growth stands host highly similar communities. • Current deadwood volume did not influence the composition of communities. • Successive clearcut rotations may lead to biodiversity erosion. • Insect outbreaks present an alternative to old-growth for biological conservation. [ABSTRACT FROM AUTHOR]

Human activity and climate change are increasing the spread of species across the planet, threatening biodiversity and ecosystem functions. Invasion engineers, such as birds, facilitate plant growth through manuring of soil, while native vegetation influences plant germination by creating suitable microhabitats which are especially valuable in cold and dry polar regions. Here we tested how penguin‐derived nitrogen, several common Antarctic moss species and warming affect seed germination and growth of the non‐native grass Agrostis capillaris under laboratory conditions. Experimental settings included a simulation of contemporary season‐specific Antarctic light and temperature (2°C) conditions and a +5°C warming scenario. Mosses (Andreaea depressinervis, A. regularis, Sanionia uncinata and Chorisodontium aciphyllum) incorporated a range of nitrogen content and isotopic nitrogen signatures (δ15N) due to variation in sampling proximity to penguin colonies. Moss species greatly affected time to germination with consequences for further growth under the simulated Antarctic conditions. Grass seeds germinated 10 days earlier among A. regularis compared to S. uncinata and C. aciphyllum and 26 days earlier compared to A. depressinervis. Moss‐specific effects are likely related to microclimatic differences within the moss canopy. Warming reduced this moss influence. Grass emerged on average 20 days earlier under warming, leading to increased leaf count (88%), plant height (112%) and biomass (145%). Positive correlations were identified between moss and grass nitrogen content (r = 0.377), grass biomass (r = 0.332) and height (r = 0.742) with stronger effects under the warming scenario. Transfer of nitrogen from moss to grass was confirmed by δ15N (r = 0.803). Overall, the results suggest a shift from temperature‐limited to N‐limited growth of invasive plants under increased warming in the maritime Antarctic. [ABSTRACT FROM AUTHOR]

• Different biocrust successional stages leach differing nutrients into sub-crust soil. • Moss-dominated biocrust leached more metabolites than other biocrust types. • Lightly-pigmented biocrusts leach the most nitrogen into sub-crust soils. • Nutrient pools below biocrusts are similar between light and darkpigmented crusts. • Leachate content does not affect short term CO 2 release from sub-crust soils. Dryland ecosystems can be constrained by low soil fertility. Within drylands, the soil nutrient and organic carbon (C) cycling that does occur is often mediated by soil surface communities known as biological soil crusts (biocrusts), which cycle C and nutrients in the top ca. 0–2 cm of soil. However, the degree to which biocrusts are influencing soil fertility and biogeochemical cycling in deeper, subsurface mineral soils is unclear. The movement of dissolved resources from biocrusts to deeper soil layers in leachate may be one of the main mechanisms through which biocrust fertility is transferred downward towards deeper microbial communities and plant roots occurring within mineral soil. Here we examined the role of biocrust leachate in contributing to subsurface nutrient and soluble C pools and subsurface microbial cycling. We collected biocrusts from three biocrust successional stages and explored resource pools in situ at multiple soil depths, while collecting leachate and measuring nutrient and organic C concentrations and metabolite composition from each successional stage in the laboratory. After four leachate collections, we conducted an incubation of mineral soil collected from below each biocrust successional stage to measure heterotrophic microbial CO 2 flux and biomass. Overall, our findings observed that the degree of nutrient and C connectivity between biocrusts and the sub-crust mineral soil depended on the biocrust successional stage and the element being considered, and the influence of biocrust successional stage on mineral soil CO 2 flux is likely related to long-term resource build up. Together, our results suggest that the influence of biocrust leachate on subsurface mineral soil is complex and context dependent, but, over longer time periods and at later successional stages, can have measurable effects on dryland soil biogeochemical cycling with feedbacks to resource availability and CO 2 flux. [ABSTRACT FROM AUTHOR]