Characteristics of Wetting-Induced Bacteriophage Blooms in Biological Soil Crust.

Biological soil crusts (biocrusts) are photosynthetic "hot spots" in deserts and cover ∼12% of the Earth's terrestrial surface, and yet they face an uncertain future given expected shifts in rainfall events. Laboratory wetting of biocrust communities is known to cause a bloom of Firmicutes which rapidly become dominant community members within 2 days after emerging from a sporulated state. We hypothesized that their bacteriophages (phages) would respond to such a dramatic increase in their host's abundance. In our experiment, wetting caused Firmicutes to bloom and triggered a significant depletion of cyanobacterial diversity. We used genome-resolved metagenomics to link phage to their hosts and found that the bloom of the genus Bacillus correlated with a dramatic increase in the number of Caudovirales phages targeting these diverse spore-formers (r = 0.762). After 2 days, we observed dramatic reductions in the relative abundances of Bacillus, while the number of Bacillus phages continued to increase, suggestive of a predator-prey relationship. We found predicted auxiliary metabolic genes (AMGs) associated with sporulation in several Caudovirales genomes, suggesting that phages may influence and even benefit from sporulation dynamics in biocrusts. Prophage elements and CRISPR-Cas repeats in Firmicutes metagenome-assembled genomes (MAGs) provide evidence of recent infection events by phages, which were corroborated by mapping viral contigs to their host MAGs. Combined, these findings suggest that the blooming Firmicutes become primary targets for biocrust Caudovirales phages, consistent with the classical "kill-the-winner" hypothesis.IMPORTANCE This work forms part of an overarching research theme studying the effects of a changing climate on biological soil crust (biocrust) in the Southwestern United States. To our knowledge, this study was the first to characterize bacteriophages in biocrust and offers a view into the ecology of phages in response to a laboratory wetting experiment. The phages identified here represent lineages of Caudovirales, and we found that the dynamics of their interactions with their Firmicutes hosts explain the collapse of a bacterial bloom that was induced by wetting. Moreover, we show that phages carried host-altering metabolic genes and found evidence of proviral infection and CRISPR-Cas repeats within host genomes. Our results suggest that phages exert controls on population density by lysing dominant bacterial hosts and that they further impact biocrust by acquiring host genes for sporulation. Future research should explore how dominant these phages are in other biocrust communities and quantify how much the control and lysis of blooming populations contributes to nutrient cycling in biocrusts

Dynamics of organic carbon losses by water erosion after biocrust removal

In arid and semiarid ecosystems, plant interspaces are frequently covered by communities of cyanobacteria, algae, lichens and mosses, known as biocrusts. These crusts often act as runoff sources and are involved in soil stabilization and fertility, as they prevent erosion by water and wind, fix atmospheric C and N and contribute large amounts of C to soil. Their contribution to the C balance as photosynthetically active surfaces in arid and semiarid regions is receiving growing attention. However, very few studies have explicitly evaluated their contribution to organic carbon (OC) lost from runoff and erosion, which is necessary to ascertain the role of biocrusts in the ecosystem C balance. Furthermore, biocrusts are not resilient to physical disturbances, which generally cause the loss of the biocrust and thus, an increase in runoff and erosion, dust emissions, and sediment and nutrient losses. The aim of this study was to find out the influence of biocrusts and their removal on dissolved and sediment organic carbon losses. One-hour extreme rainfall simulations (50 mm h-1) were performed on small plots set up on physical soil crusts and three types of biocrusts, representing a development gradient, and also on plots where these crusts were removed from. Runoff and erosion rates, dissolved organic carbon (DOC) and organic carbon bonded to sediments (SdOC) were measured during the simulated rain. Our results showed different SdOC and DOC for the different biocrusts and also that the presence of biocrusts substantially decreased total organic carbon (TOC) (average 1.80±1.86 g m-2) compared to physical soil crusts (7.83±3.27 g m-2). Within biocrusts, TOC losses decreased as biocrusts developed, and erosion rates were lower. Thus, erosion drove TOC losses while no significant direct relationships were found between TOC losses and runoff. In both physical crusts and biocrusts, DOC and SdOC concentrations were higher during the first minutes after runoff began and decreased over time as nutrient-enriched fine particles were washed away by runoff water. Crust removal caused a strong increase in water erosion and TOC losses. The strongest impacts on TOC losses after crust removal occurred on the lichen plots, due to the increased erosion when they were removed. DOC concentration was higher in biocrust-removed soils than in intact biocrusts, probably because OC is more strongly retained by BSC structures, but easily blown away in soils devoid of them. However, SdOC concentration was higher in intact than removed biocrusts associated with greater OC content in the top crust than in the soil once the crust is scraped off. Consequently, the loss of biocrusts leads to OC impoverishment of nutrient-limited interplant spaces in arid and semiarid areas and the reduction of soil OC heterogeneity, essential for vegetation productivity and functioning of this type of ecosystems.This work was partially supported by several research projects: COSTRAS (RNM 3614), funded by the Consejería de Innovación, Ciencia y Empresa (Junta de Andalucía) including ERD (European Union of Regional Development) Funds, BACARCOS (CGL2011–29429) funded by Spanish national Plan for Research, Development and Innovation and including European Union of Regional Development Funds ERDF funds

Reproduction and Dispersal of Biological Soil Crust Organisms

Biological soil crusts (BSCs) consist of a diverse and highly integrated community of organisms that effectively colonize and collectively stabilize soil surfaces. BSCs vary in terms of soil chemistry and texture as well as the environmental parameters that combine to support unique combinations of organisms—including cyanobacteria dominated, lichen-dominated, and bryophyte-dominated crusts. The list of organismal groups that make up BSC communities in various and unique combinations include—free living, lichenized, and mycorrhizal fungi, chemoheterotrophic bacteria, cyanobacteria, diazotrophic bacteria and archaea, eukaryotic algae, and bryophytes. The various BSC organismal groups demonstrate several common characteristics including—desiccation and extreme temperature tolerance, production of various soil binding chemistries, a near exclusive dependency on asexual reproduction, a pattern of aerial dispersal over impressive distances, and a universal vulnerability to a wide range of human-related perturbations. With this publication, we provide literature-based insights as to how each organismal group contributes to the formation and maintenance of the structural and functional attributes of BSCs, how they reproduce, and how they are dispersed. We also emphasize the importance of effective application of molecular and microenvironment sampling and assessment tools in order to provide cogent and essential answers that will allow scientists and land managers to better understand and manage the biodiversity and functional relationships of soil crust communities

Rapidly Restoring Biological Soil Crusts and Ecosystem Functions in a Severely Disturbed Desert Ecosystem

Restoring biological soil crusts (biocrusts) in degraded drylands can contribute to recovery of ecosystem functions that have global implications, including erosion resistance and nutrient cycling. To examine techniques for restoring biocrusts, we conducted a replicated, factorial experiment on recently abandoned road surfaces by applying biocrust inoculation (salvaged and stored dry for two years), salvaged topsoil, an abiotic soil amendment (wood shavings), and planting of a dominant perennial shrub (Ambrosia dumosa). Eighteen months after treatments, we measured biocrust abundance and species composition, soil chlorophyll a content and fertility, and soil resistance to erosion. Biocrust addition significantly accelerated biocrust recovery on disturbed soils, including increasing lichen and moss cover and cyanobacteria colonization. Compared to undisturbed controls, inoculated plots had similar lichen and moss composition, recovered 43% of total cyanobacteria density, had similar soil chlorophyll content, and exhibited recovery of soil fertility and soil stability. Inoculation was the only treatment that generated lichen and moss cover. Topsoil application resulted in partial recovery of the cyanobacteria community and soil properties. Compared to untreated disturbed plots, topsoil application without inoculum increased cyanobacteria density by 186% and moderately improved soil chlorophyll and ammonium content and soil stability. Topsoil application produced 22% and 51% of the cyanobacteria density g−1 soil compared to undisturbed and inoculated plots, respectively. Plots not treated with either topsoil or inoculum had significantly lower cyanobacteria density, soil chlorophyll and ammonium concentrations, and significantly higher soil nitrate concentration. Wood shavings and Ambrosia had no influence on biocrust lichen and moss species recovery but did affect cyanobacteria composition and soil fertility. Inoculation of severely disturbed soil with native biocrusts rapidly restored biocrust communities and soil stability such that restored areas were similar to undisturbed desert within three years. Using salvaged biocrust as inoculum can be an effective tool in ecological restoration because of its efficacy and simple implementation. Although salvaging biocrust material can be technically difficult and potentially costly, utilizing opportunities to salvage material in planned future disturbance can provide additional land management tools

Effects of Rate of Drying, Life History Phase, and Ecotype on the Ability of the Moss Bryum Argenteum to Survive Desiccation Events and the Influence on Conservation and Selection of Material for Restoration

Desiccation stress is frequently experienced by the moss Bryum argenteum and can influence survival, propagation and niche selection. We attempted to disentangle the interacting factors of life history phase (five categories) and rate of desiccation (time allotted for induction of desiccation tolerance) for 13 ecotypes of B. argenteum. Using chlorophyll fluorescence as a stress index, we determined how these parameters influenced desiccation tolerance. Rate of drying and life phase significantly affected desiccation tolerance. The reaction norms of desiccation tolerance displayed by the 13 ecotypes showed a substantial degree of variation in phenotypic plasticity. We observed differences in survival and fluorescence between rapid and slow drying events in juveniles. These same drying applications did not produce as large of a response for adult shoots (which consistently displayed high values). Some juvenile and protonemal ecotypes, such as those from the southwest United States, possessed higher innate tolerance to rapid drying, and greater resilience compared to ecotypes sourced from mesic localities in the United States. These results show a complex nuanced response to desiccation with ecotypes displaying a range of responses to desiccation reflecting both inherently different capacities for tolerating desiccation as well as variation in capacity for phenotypic plasticity. Our results suggest that we should expect few short-term effects of climate change due to high desiccation tolerance of adult shoots, but significant adverse long-term effects on colony establishment due to low tolerance of protonema and juvenile shoots. Further, we would recommend that future studies using mosses for habitat restoration of aridlands consider the desiccation tolerance capacity of individual ecotypes used for cultivation and later re-introduction. Understanding how mosses respond to desiccation is essential to interpret ecological roles, habitat preferences, selective pressures, and responses to climate change, and to estimate the potential effects of climate changes on bryophyte species and populations

The potential of permanent gullies in Europe as geomorphosites

Geotourism is a useful way to educate societies in the field of geomorphology and natural hazards. Geosites, including geomorphosites, represent the basis for the development of this type of tourism. This study describes twelve representative gully regions within nine European countries. The characteristics of 42 permanent gullies, gully systems and badland landscapes are presented and a geotouristic assessment of these gullies was made, based on scientific, educational, functional and touristic indicators. This assessment demonstrates a large difference between the selected gully sites, particularly with regard to functional and tourist values. The geotouristic potential of gullies is the highest in Italy and Spain and the lowest in Romania and Latvia. In some countries, permanent gullies are not regarded as geotouristic attractions at all, while in others they constitute a significant element of their tourism development strategy. Based on the analysis of scientific values of the gully sites, educational lessons to be learned were identified that are mainly related to i) gullies as a geological window, ii) present-day geomorphological processes, and iii) stages of historical gully erosion reflecting past human-environment interactions. These must be part of a broader strategy for the development of geotourism in gully regions. Gullies are potential geosites within existing or planned geoparks. Values of gullies for other forms of tourism (active, cultural, nature), should be also emphasized

Six Collective Challenges for Sustainability of Almería Greenhouse Horticulture

Globally, current food consumption and trade are placing unprecedented demand on agricultural systems and increasing pressure on natural resources, requiring tradeoffs between food security and environmental impacts especially given the tension between market-driven agriculture and agro-ecological goals. In order to illustrate the wicked social, economic and environmental challenges and processes to find transformative solutions, we focus on the largest concentration of greenhouses in the world located in the semi-arid coastal plain of South-east Spain. Almería family farming, predominantly cooperative, greenhouse intensive production, commenced after the 1960s and has resulted in very significant social and economic benefits for the region, while also having important negative environmental and biodiversity impacts, as well as creating new social challenges. The system currently finds itself in a crisis of diminishing economic benefits and increasing environmental and social dilemmas. Here, we present the outcomes of multi-actor, transdisciplinary research to review and provide collective insights for solutions-oriented research on the sustainability of Almeria’s agricultural sector. The multi-actor, transdisciplinary process implemented collectively, and supported by scientific literature, identified six fundamental challenges to transitioning to an agricultural model that aims to ameliorate risks and avoid a systemic collapse, whilst balancing a concern for profitability with sustainability: (1) Governance based on a culture of shared responsibility for sustainability, (2) Sustainable and efficient use of water, (3) Biodiversity conservation, (4) Implementing a circular economy plan, (5) Technology and knowledge transfer, and (6) Image and identity. We conclude that the multi-actor transdisciplinary approach successfully facilitated the creation of a culture of shared responsibility among public, private, academic, and civil society actors. Notwithstanding plural values, challenges and solutions identified by consensus point to a nascent acknowledgement of the strategic necessity to locate agricultural economic activity within social and environmental spheres.This paper demonstrates the need to establish transdisciplinary multi-actor work-schemes to continue collaboration and research for the transition to an agro-ecological model as a means to remain competitive and to create value

Biological soil crust effects and responses in arid ecosystems: recent advances at the species level

La costra biológica del suelo (CBS) es un componente complejo del ecosistema que engloba diferentes organismos (líquenes, musgos, hepáticas, cianobacterias, hongos, algas) presentes en las primeras capas de suelo. La CBS se encuentra en una amplia variedad de ecosistemas, aunque generalmente es más abundante en ecosistemas donde la cobertura de plantas vasculares es escasa, como los ecosistemas áridos. En estos ecosistemas, la CBS contribuye considerablemente a su biodiversidad y funcionamiento. Debido a la gran dificultad para la identificación de especies de estas comunidades, la mayoría de la investigación sobre la CBS se ha desarrollado a escala de comunidad y grupo morfológico. A este nivel, se ha podido observar el gran potencial de estas comunidades de contribuir a la estructura y dinámica del ecosistema: interaccionan con las primeras capas del suelo y con otros organismos, participan en la fijación de carbono y nitrógeno, así como en procesos hidrológicos y en el ciclo de nutrientes. Sin embargo, avances recientes en el conocimiento de la CBS arrojan interesantes y marcadas diferencias en la ecología y el papel funcional de las distintas especies que la componen, con las consecuentes implicaciones en la gestión y conservación de estas comunidades y de los ecosistemas que habitan. En particular, se han observado respuestas específicas en términos de presencia, abundancia y frecuencia ante diversos factores ambientales (variables climáticas, tipo de sustrato, presencia de plantas vasculares y perturbación por pastoreo – recuperación natural), así como un efecto a nivel de especie sobre las propiedades del suelo.Biological soil crusts (BSCs) constitute a complex component of the ecosystem formed by different organisms (lichens, mosses, liverworts, cyanobacteria, fungi, algae) associated with soil surface. These communities are present in a wide variety of ecosystems; however, their abundance is generally higher in arid environments with sparse vegetation cover. In these ecosystems, BSCs greatly contribute to biodiversity and ecosystem functioning. Due to technical difficulties in species identification, most studies on BSCs have been carried out at community and morphotype levels. These studies have emphasized the potential role of BSCs in defining ecosystem structure and functioning by: interacting with topsoil layers and other soil organisms, participating in carbon and nitrogen fixation, and also in hydrological and nutrient cycling. Notwithstanding, recent advances in our knowledge about BSCs show substantial and interesting differences in the ecology and functional roles of BSC species, with marked implications in the management and conservation of these communities and their ecosystems. Particularly, it has been observed that BSC presence, abundance and frequency respond differently to diverse environmental factors (climatic variables, soil type, presence of vascular plants, and grazing disturbance - natural recovery) at the species level, and also do BSC effects on topsoil properties.Este trabajo ha sido parcialmente financiado por la Comunidad Autónoma de Madrid (Proyecto REMEDINAL2, S2009/AMB-1783) y el Ministerio de Ciencia e Innovación (Proyecto EPICON, CGL2010-22049)