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Biocrusts are a worldwide phenomenon, contributing substantially to ecosystem functioning. Their growth and survival depend on multiple environmental factors, including climatic ones, and the relations of these factors to physiological processes. Responses of biocrusts to individual environmental factors have been examined in a large number of field and laboratory experiments. These observational data, however, have rarely been assembled into a comprehensive, consistent framework that allows quantitative exploration of the roles of multiple environmental factors and physiological properties for the performance of biocrusts, in particular across climatic regions. Here we used a data-driven mechanistic modelling framework to simulate the carbon balance of biocrusts, a key measure of their growth and survival. We thereby assessed the relative importance of physiological and environmental factors for the carbon balance at six study sites that differ in climatic conditions. Moreover, we examined the role of seasonal acclimation of physiological properties using our framework, since the effects of this process on the carbon balance of biocrusts are poorly constrained so far. We found substantial effects of air temperature, CO2 concentration, and physiological parameters that are related to respiration on biocrust carbon balance, which differ, however, in their patterns across regions. The ambient CO2 concentration is the most important factor for biocrusts from drylands, while air temperature has the strongest impact at alpine and temperate sites. Metabolic respiration cost plays a more important role than optimum temperature for gross photosynthesis at the alpine site; this is not the case, however, in drylands and temperate regions. Moreover, we estimated a small annual carbon gain of 1.5 gm-2yr-1 by lichen-dominated biocrust and 1.9 gm-2yr-1 by moss-dominated biocrust at a dryland site, while the biocrusts lost a large amount of carbon at some of the temperate sites (e.g. −92.1 for lichen-dominated and −74.7 gm-2yr-1 for moss-dominated biocrust). These strongly negative values contradict the observed survival of the organisms at the sites and may be caused by the uncertainty in environmental conditions and physiological parameters, which we assessed in a sensitivity analysis. Another potential explanation for this result may be the lack of acclimation in the modelling approach, since the carbon balance can increase substantially when testing for seasonally varying parameters in the sensitivity analysis. We conclude that the uncertainties in air temperature, CO2 concentration, respiration-related physiological parameters, and the absence of seasonal acclimation in the model for humid temperate and alpine regions may be a relevant source of error and should be taken into account in future approaches that aim at estimating the long-term biocrust carbon balance based on ecophysiological data.

1. Manipulative experiments typically show a decrease in dryland biocrust cover and altered species composition under climate change. Biocrust‐forming lichens, such as the globally distributed Diploschistes diacapsis, are particularly affected and show a decrease in cover with simulated climate change. However, the underlying mechanisms are not fully understood, and long‐term interacting effects of different drivers are largely unknown due to the short‐term nature of the experimental studies conducted so far. 2. We addressed this gap and successfully parameterised a process‐based model for D. diacapsis to quantify how changing atmospheric CO2, temperature, rainfall amount and relative humidity affect its photosynthetic activity and cover. We also mimicked a long‐term manipulative climate change experiment to understand the mechanisms underlying observed patterns in the field. 3. The model reproduced observed experimental findings: warming reduced lichen cover, whereas less rainfall had no effect on lichen performance. This warming effect was caused by the associated decrease in relative humidity and non‐rainfall water inputs, which are major water sources for biocrust‐forming lichens. Warming alone, however, increased cover because higher temperatures promoted photosynthesis during early morning hours with high lichen activity. When combined, climate variables showed non‐additive effects on lichen cover, and effects of increased CO2 levelled off with decreasing levels of relative humidity. 4. Synthesis. Our results show that a decrease in relative humidity, rather than an increase in temperature, may be the key factor for the survival of the lichen D. diacapsis under climate change and that effects of increased CO2 levels might be offset by a reduction in non‐rainfall water inputs in the future. Because of a global trend towards warmer and drier air and the widespread global distribution of D. diacapsis, this will affect lichen‐dominated dryland biocrust communities and their role in regulating ecosystem functions worldwide. This research was supported by the Collaborative Research Centre 973 (www.sfb973.de) of the German Research Foundation (DFG) and by the European Research Council (grant agreement no. 647038 (BIODESERT)). P. Porada appreciates funding by Deutsche Forschungsgemeinschaft (DFG, German Research Foundation)—408092731. F.T. Maestre acknowledges support from Generalitat Valenciana (CIDEGENT/2018/041) and the Alexander Von Humboldt Foundation. J. Raggio acknowledges the ERA-Net BiodivERsA program as Soil Crust InterNational (SCIN) and The Spanish Ministerio de Economía y Competitividad (MINECO) project numbers PRI-PIMBDV-2011-0874 and CRYPTOCOVER, CTM2015-64728-C21-R.

Despite the important role that biocrust communities play in maintaining ecosystem structure and functioning in drylands world-wide, few studies have evaluated how climate change will affect them. Using data from an 8-yr-old manipulative field experiment located in central Spain, we evaluated how warming, rainfall exclusion and their combination affected the dynamics of biocrust communities in areas that initially had low (< 20%, LIBC plots) and high (> 50%, HIBC plots) biocrust cover. Warming reduced the richness (35 6%), diversity (25 8%) and cover (82 5%) of biocrusts in HIBC plots. The presence and abundance of mosses increased with warming through time in these plots, although their growth rate was much lower than the rate of lichen death, resulting in a net loss of biocrust cover. On average, warming caused a decrease in the abundance (64 7%) and presence (38 24%) of species in the HIBC plots. Over time, lichens and mosses colonized the LIBC plots, but this process was hampered by warming in the case of lichens. The observed reductions in the cover and diversity of lichen-dominated biocrusts with warming will lessen the capacity of drylands such as that studied here to sequester atmospheric CO2 and to provide other key ecosystem services associated to these communities.

Here we report details of the European research initiative “Soil Crust International” (SCIN) focusing on the biodiversity of biological soil crusts (BSC, composed of bacteria, algae, lichens, and bryophytes) and on functional aspects in their specific environment. Known as the so-called “colored soil lichen community” (Bunte Erdflechtengesellschaft), these BSCs occur all over Europe, extending into subtropical and arid regions. Our goal is to study the uniqueness of these BSCs on the regional scale and investigate how this community can cope with large macroclimatic differences. One of the major aims of this project is to develop biodiversity conservation and sustainable management strategies for European BSCs. To achieve this, we established a latitudinal transect from the Great Alvar of Öland, Sweden in the north over Gössenheim, Central Germany and Hochtor in the Hohe Tauern National Park, Austria down to the badlands of Tabernas, Spain in the south. The transect stretches over 20 latitude and 2,300 m in altitude, including natural (Hochtor, Tabernas) and semi-natural sites that require maintenance such as by grazing activities (Öland, Gössenheim). At all four sites BSC coverage exceeded 30 % of the referring landscape, with the alpine site (Hochtor) reaching the highest cyanobacterial cover and the two semi-natural sites (Öland, Gössenheim) the highest bryophyte cover. Although BSCs of the four European sites share a common set of bacteria, algae (including cyanobacteria) lichens and bryophytes, first results indicate not only climate specific additions of species, but also genetic/phenotypic uniqueness of species between the four sites. While macroclimatic conditions are rather different, microclimatic conditions and partly soil properties seem fairly homogeneous between the four sites, with the exception of water availability. Continuous activity monitoring of photosystem II revealed the BSCs of the Spanish site as the least active in terms of photosynthetic active periods. (VLID)4424548

Here we report details of the European research initiative “Soil Crust International” (SCIN) focusing on the biodiversity of biological soil crusts (BSC, composed of bacteria, algae, lichens, and bryophytes) and on functional aspects in their specific environment. Known as the so-called “colored soil lichen community” (Bunte Erdflechtengesellschaft), these BSCs occur all over Europe, extending into subtropical and arid regions. Our goal is to study the uniqueness of these BSCs on the regional scale and investigate how this community can cope with large macroclimatic differences. One of the major aims of this project is to develop biodiversity conservation and sustainable management strategies for European BSCs. To achieve this, we established a latitudinal transect from the Great Alvar of Öland, Sweden in the north over Gössenheim, Central Germany and Hochtor in the Hohe Tauern National Park, Austria down to the badlands of Tabernas, Spain in the south. The transect stretches over 20° latitude and 2,300 m in altitude, including natural (Hochtor, Tabernas) and semi-natural sites that require maintenance such as by grazing activities (Öland, Gössenheim). At all four sites BSC coverage exceeded 30 % of the referring landscape, with the alpine site (Hochtor) reaching the highest cyanobacterial cover and the two semi-natural sites (Öland, Gössenheim) the highest bryophyte cover. Although BSCs of the four European sites share a common set of bacteria, algae (including cyanobacteria) lichens and bryophytes, first results indicate not only climate specific additions of species, but also genetic/phenotypic uniqueness of species between the four sites. While macroclimatic conditions are rather different, microclimatic conditions and partly soil properties seem fairly homogeneous between the four sites, with the exception of water availability. Continuous activity monitoring of photosystem II revealed the BSCs of the Spanish site as the least active in terms of photosynthetic active periods. Version of record

1. Manipulative experiments typically show a decrease in dryland biocrust cover and altered species composition under climate change. Biocrust-forming lichens, such as the globally distributed Diploschistes diacapsis, are particularly affected and show a decrease in cover with simulated climate change. However, the underlying mechanisms are not fully understood, and long-term interacting effects of different drivers are largely unknown due to the short-term nature of the experimental studies conducted so far. 2. We addressed this gap and successfully parameterised a process-based model for D. diacapsis to quantify how changing atmospheric CO2, temperature, rainfall amount and relative humidity affect its photosynthetic activity and cover. We also mimicked a long-term manipulative climate change experiment to understand the mechanisms underlying observed patterns in the field. 3. The model reproduced observed experimental findings: warming reduced lichen cover, whereas less rainfall had no effect on lichen performance. This warming effect was caused by the associated decrease in relative humidity and nonrainfall water inputs, which are major water sources for biocrust-forming lichens. Warming alone, however, increased cover because higher temperatures promoted photosynthesis during early morning hours with high lichen activity. When combined, climate variables showed non-additive effects on lichen cover, and effects of increased CO2 levelled off with decreasing levels of relative humidity. 4. Synthesis. Our results show that a decrease in relative humidity, rather than an increase in temperature, may be the key factor for the survival of the lichen D. diacapsis under climate change and t hat effects of increased CO2 levels might be offset by a reduction in non-rainfall water inputs in the future. Because of a global trend towards warmer and drier air and the widespread global distribution of D. diacapsis, this will affect lichen-dominated dryland biocrust communities and their role in regulating ecosystem functions worldwide.