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Abstract: The emergence of alternative stable states in forest systems has significant implications for the functioning and structure of the terrestrial biosphere, yet empirical evidence remains scarce. Here, we combine global forest biodiversity observations and simulations to test for alternative stable states in the presence of evergreen and deciduous forest types. We reveal a bimodal distribution of forest leaf types across temperate regions of the Northern Hemisphere that cannot be explained by the environment alone, suggesting signatures of alternative forest states. Moreover, we empirically demonstrate the existence of positive feedbacks in tree growth, recruitment and mortality, with trees having 4–43% higher growth rates, 14–17% higher survival rates and 4–7 times higher recruitment rates when they are surrounded by trees of their own leaf type. Simulations show that the observed positive feedbacks are necessary and sufficient to generate alternative forest states, which also lead to dependency on history (hysteresis) during ecosystem transition from evergreen to deciduous forests and vice versa. We identify hotspots of bistable forest types in evergreen-deciduous ecotones, which are likely driven by soil-related positive feedbacks. These findings are integral to predicting the distribution of forest biomes, and aid to our understanding of biodiversity, carbon turnover, and terrestrial climate feedbacks.

Earth harbours an extraordinary plant phenotypic diversity1 that is at risk from ongoing global changes2,3. However, it remains unknown how increasing aridity and livestock grazing pressure—two major drivers of global change4–6—shape the trait covariation that underlies plant phenotypic diversity1,7. Here we assessed how covariation among 20 chemical and morphological traits responds to aridity and grazing pressure within global drylands. Our analysis involved 133,769 trait measurements spanning 1,347 observations of 301 perennial plant species surveyed across 326 plots from 6 continents. Crossing an aridity threshold of approximately 0.7 (close to the transition between semi-arid and arid zones) led to an unexpected 88% increase in trait diversity. This threshold appeared in the presence of grazers, and moved toward lower aridity levels with increasing grazing pressure. Moreover, 57% of observed trait diversity occurred only in the most arid and grazed drylands, highlighting the phenotypic uniqueness of these extreme environments. Our work indicates that drylands act as a global reservoir of plant phenotypic diversity and challenge the pervasive view that harsh environmental conditions reduce plant trait diversity8–10. They also highlight that many alternative strategies may enable plants to cope with increases in environmental stress induced by climate change and land-use intensification.Analysis of 20 chemical and morphological plant traits at diverse sites across 6 continents shows that the transition from semi-arid to arid zones is associated with an unexpected 88% increase in trait diversity. [ABSTRACT FROM AUTHOR]

Climate change will affect the way biodiversity influences the stability of plant communities. Although biodiversity, associated species asynchrony, and species stability could enhance community stability, the understanding of potential nonlinear shifts in the biodiversity–stability relationship across a wide range of aridity (measured as the aridity index, the precipitation/potential evapotranspiration ratio) gradients and the underlying mechanisms remain limited. Using an 8‐year dataset from 687 sites in Mongolia, which included 5496 records of vegetation and productivity, we found that the temporal stability of plant communities decreased more rapidly in more arid areas than in less arid areas. The result suggests that future aridification across terrestrial ecosystems may adversely affect community stability. Additionally, we identified nonlinear shifts in the effects of species richness and species synchrony on temporal community stability along the aridity gradient. Species synchrony was a primary driver of community stability, which was consistently negatively affected by species richness while being positively affected by the synchrony between C3 and C4 species across the aridity gradient. These results highlight the crucial role of C4 species in stabilizing communities through differential responses to interannual climate variations between C3 and C4 species. Notably, species richness and the synchrony between C3 and C4 species independently regulated species synchrony, ultimately affecting community stability. We propose that maintaining plant communities with a high diversity of C3 and C4 species will be key to enhancing community stability across Mongolian grasslands. Moreover, species synchrony, species stability, species richness and the synchrony between C3 and C4 species across the aridity gradient consistently mediated the impacts of aridity on community stability. Hence, strategies aimed at promoting the maintenance of biological diversity and composition will help ecosystems adapt to climate change or mitigate its adverse effects on ecosystem stability. [ABSTRACT FROM AUTHOR]

Plant–soil biodiversity interactions are fundamental for the functioning of terrestrial ecosystems. Yet, the existence of a set of globally distributed topsoil microbial and small invertebrate organisms consistently associated with land plants (i.e., their consistent soil‐borne microbiome), together with the environmental preferences and functional capabilities of these organisms, remains unknown. We conducted a standardized field survey under 150 species of land plants, including 58 species of bryophytes and 92 of vascular plants, across 124 locations from all continents. We found that, despite the immense biodiversity of soil organisms, the land plants evaluated only shared a small fraction (less than 1%) of all microbial and invertebrate taxa that were present across contrasting climatic and soil conditions and vegetation types. These consistent taxa were dominated by generalist decomposers and phagotrophs and their presence was positively correlated with the abundance of functional genes linked to mineralization. Finally, we showed that crossing environmental thresholds in aridity (aridity index of 0.65, i.e., the transition from mesic to dry ecosystems), soil pH (5.5; i.e., the transition from acidic to strongly acidic soils), and carbon (less than 2%, the lower limit of fertile soils) can result in drastic disruptions in the associations between land plants and soil organisms, with potential implications for the delivery of soil ecosystem processes under ongoing global environmental change. [ABSTRACT FROM AUTHOR]

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Self-organized spatial patterns are a common feature of complex systems, ranging from microbial communities to mussel beds and drylands. While the theoretical implications of these patterns for ecosystem-level processes, such as functioning and resilience, have been extensively studied, empirical evidence remains scarce. To address this gap, we analyzed global drylands along an aridity gradient using remote sensing, field data, and modeling. We found that the spatial structure of the vegetation strengthens as aridity increases, which is associated with the maintenance of a high level of soil multifunctionality, even as aridity levels rise up to a certain threshold. The combination of these results with those of two individual-based models indicate that self-organized vegetation patterns not only form in response to stressful environmental conditions but also provide drylands with the ability to adapt to changing conditions while maintaining their functioning, an adaptive capacity which is lost in degraded ecosystems. Self-organization thereby plays a vital role in enhancing the resilience of drylands. Overall, our findings contribute to a deeper understanding of the relationship between spatial vegetation patterns and dryland resilience. They also represent a significant step forward in the development of indicators for ecosystem resilience, which are critical tools for managing and preserving these valuable ecosystems in a warmer and more arid world [ABSTRACT FROM AUTHOR]

Crossing certain aridity thresholds in global drylands can lead to abrupt decays of ecosystem attributes such as plant productivity, potentially causing land degradation and desertification. It is largely unknown, however, whether these thresholds can be altered by other key global change drivers known to affect the water-use efficiency and productivity of vegetation, such as elevated CO2 and nitrogen (N). Using >5000 empirical measurements of plant biomass, we showed that crossing an aridity (1–precipitation/potential evapotranspiration) threshold of ∼0.50, which marks the transition from dry sub-humid to semi-arid climates, led to abrupt declines in aboveground biomass (AGB) and progressive increases in root:shoot ratios, thus importantly affecting carbon stocks and their distribution. N addition significantly increased AGB and delayed the emergence of its aridity threshold from 0.49 to 0.55 (P  < 0.05). By coupling remote sensing estimates of leaf area index with simulations from multiple models, we found that CO2 enrichment did not alter the observed aridity threshold. By 2100, and under the RCP 8.5 scenario, we forecast a 0.3% net increase in the global land area exceeding the aridity threshold detected under a scenario that includes N deposition, in comparison to a 2.9% net increase if the N effect is not considered. Our study thus indicates that N addition could mitigate to a great extent the negative impact of increasing aridity on plant biomass in drylands. These findings are critical for improving forecasts of abrupt vegetation changes in response to ongoing global environmental change. [ABSTRACT FROM AUTHOR]

Fairy circles (FCs) are regular vegetation patterns found in drylands of Namibia and Western Australia. It is virtually unknown whether they are also present in other regions of the world and which environmental factors determine their distribution. We conducted a global systematic survey and found FC-like vegetation patterns in 263 sites from 15 countries and three continents, including the Sahel, Madagascar, and Middle-West Asia. FC-like vegetation patterns are found in environments characterized by a unique combination of soil (including low nutrient levels and high sand content) and climatic (arid regions with high temperatures and high precipitation seasonality) conditions. In addition to these factors, the presence of specific biological elements (termite nests) in certain regions also plays a role in the presence of these patterns. Furthermore, areas with FC-like vegetation patterns also showed more stable temporal productivity patterns than those of surrounding areas. Our study presents a global atlas of FCs and provides unique insights into the ecology and biogeography of these fascinating vegetation patterns. [ABSTRACT FROM AUTHOR]

Soil biota influences nutrient cycling and climate regulation and represents an important fraction of global biodiversity, yet we know very little about how this soil biota responds to habitat fragmentation and degradation of habitat quality.We studied the response of different soil trophic groups (microbes and soil fauna), and their trophic structure, to changes in their habitat derived from forest long‐term management and extensive tree die‐off in a Mediterranean ecosystem. Specifically, we evaluated changes in (i) habitat size, (ii) habitat resource availability and heterogeneity and (iii) habitat connectivity. To do this, we sampled the soil biota of 43 holm oak trees (and five open interspaces) differing in size, quality, heterogeneity, connectivity and the effect of die‐off (healthy or affected). We sorted soil biota by trophic group and related their richness to habitat characteristics.Seven of the 12 trophic groups evaluated increased their species richness with soil organic carbon content, which was the most frequently selected driver of soil biota (both microbial and faunal richness). Habitat connectivity positively affected the richness of larger organisms (fauna) and plant attributes (richness, productivity and specific leaf area) also showed significant but contrasting effects depending on the group evaluated.Due to the idiosyncratic responses of different groups, the entire trophic structure (microbes and fauna) was affected by a more complex set of factors than most trophic groups in isolation, including interactions between habitat size and resource availability or connectivity. A major factor influencing habitat resource availability was the die‐off of the dominant tree species (drastically altering tree productivity). We found weaker and more negative relationships between trophic groups under trees suffering from die‐off than beneath healthy trees, particularly between microbial rather than faunal groups.Synthesis. We provide a comprehensive assessment of the response of key members of the soil food web to habitat fragmentation and tree die‐off (landscape‐level plant–soil interactions), illustrating the major role of soil carbon, habitat connectivity and tree die‐off in driving soil biodiversity and trophic structure. [ABSTRACT FROM AUTHOR]

Soil micronutrients are capital for the delivery of ecosystem functioning and food provision worldwide. Yet, despite their importance, the global biogeography and ecological drivers of soil micronutrients remain virtually unknown, limiting our capacity to anticipate abrupt unexpected changes in soil micronutrients in the face of climate change. Here, we analyzed >1300 topsoil samples to examine the global distribution of six metallic micronutrients (Cu, Fe, Mn, Zn, Co and Ni) across all continents, climates and vegetation types. We found that warmer arid and tropical ecosystems, present in the least developed countries, sustain the lowest contents of multiple soil micronutrients. We further provide evidence that temperature increases may potentially result in abrupt and simultaneous reductions in the content of multiple soil micronutrients when a temperature threshold of 12–14°C is crossed, which may be occurring on 3% of the planet over the next century. Altogether, our findings provide fundamental understanding of the global distribution of soil micronutrients, with direct implications for the maintenance of ecosystem functioning, rangeland management and food production in the warmest and poorest regions of the planet. [ABSTRACT FROM AUTHOR]

Aim: Fungi are major drivers of ecosystem functioning. Increases in aridity are known to negatively impact fungal community composition in dryland ecosystems globally; yet, much less is known on the potential influence of other environmental drivers, and whether these relationships are linear or nonlinear. Time period: 2017–2021. Location: Global. Major taxa studied: Fungi. Methods: We re‐analysed multiple datasets from different dryland biogeographical regions, for a total of 912 samples and 1,483 taxa. We examined geographical patterns in community diversity and composition, and spatial, edaphic and climatic factors driving them. Results: UV index, climate seasonality, and sand content were the most important environmental predictors of community shifts, showing the strongest association with the richness of putative plant pathogens and saprobes. Important nonlinear relationships existed with each of these fungal guilds, with increases in UV and temperature seasonality above 7.5 and 900 SD (standard deviation x 100 of the mean monthly temperature), respectively, being associated with an increased probability of plant pathogen and unspecified saprotroph occurrence. Conversely, these environmental parameters had a negative relationship with litter and soil saprotroph richness. Consequently, these ecological groups might be particularly sensitive to shifts in UV radiation and climate seasonality, which is likely to disturb current plant–soil dynamics in drylands. Main conclusions: Our synthesis integrates fungal community data from drylands across the globe, allowing the investigation of fungal distribution and providing the first evidence of shifts in fungal diversity and composition of key fungal ecological groups along diverse spatial, climatic and edaphic gradients in these widely distributed ecosystems. Our findings imply that shifts in soil structure and seasonal climatic patterns induced by global change will have disproportionate consequences for the distribution of fungal groups linked to vegetation and biogeochemical cycling in drylands, with implications for plant–soil interactions in drylands. [ABSTRACT FROM AUTHOR]

The constant provision of plant productivity is integral to supporting the liability of ecosystems and human wellbeing in global drylands. Drylands are paradigmatic examples of systems prone to experiencing abrupt changes in their functioning. Indeed, space-fortime substitution approaches suggest that abrupt changes in plant productivity are widespread, but this evidence is less clear using observational time series or experimental data at a large scale. Studying the prevalence and, most importantly, the unknown drivers of abrupt (rather than gradual) dynamical patterns in drylands may help to unveil hotspots of current and future dynamical instabilities in drylands. Using a 20-y global satellite-derived temporal assessment of dryland Normalized Difference Vegetation Index (NDVI), we show that 50% of all dryland ecosystems exhibiting gains or losses of NDVI are characterized by abrupt positive/negative temporal dynamics. We further show that abrupt changes are more common among negative than positive NDVI trends and can be found in global regions suffering recent droughts, particularly around critical aridity thresholds. Positive abrupt dynamics are found most in ecosystems with low seasonal variability or high aridity. Our work unveils the high importance of climate variability on triggering abrupt shifts in vegetation and it provides missing evidence of increasing abruptness in systems intensively managed by humans, with low soil organic carbon contents, or around specific aridity thresholds. These results highlight that abrupt changes in dryland dynamics are very common, especially for productivity losses, pinpoint global hotspots of dryland vulnerability, and identify drivers that could be targeted for effective dryland management. [ABSTRACT FROM AUTHOR]

1. Rapid technological advancements and increasing data availability have improved the capacity to monitor and evaluate Earth's ecology via remote sensing. However, remote sensing is notoriously ‘blind’ to fine-scale ecological processes such as interactions among plants, which encompass a central topic in ecology. 2. Here, we discuss how remote sensing technologies can help infer plant–plant interactions and their roles in shaping plant-based systems at individual, community and landscape levels. At each of these levels, we outline the key attributes of ecosystems that emerge as a product of plant–plant interactions and could possibly be detected by remote sensing data. We review the theoretical bases, approaches and prospects of how inference of plant–plant interactions can be assessed remotely. 3. At the individual level, we illustrate how close-range remote sensing tools can help to infer plant–plant interactions, especially in experimental settings. At the community level, we use forests to illustrate how remotely sensed community structure can be used to infer dominant interactions as a fundamental force in shaping plant communities. At the landscape level, we highlight how remotely sensed attributes of vegetation states and spatial vegetation patterns can be used to assess the role of local plant–plant interactions in shaping landscape ecological systems. 4. Synthesis. Remote sensing extends the domain of plant ecology to broader and finer spatial scales, assisting to scale ecological patterns and search for generic rules. Robust remote sensing approaches are likely to extend our understanding of how plant–plant interactions shape ecological processes across scales—from individuals to landscapes. Combining these approaches with theories, models, experiments, data-driven approaches and data analysis algorithms will firmly embed remote sensing techniques into ecological context and open new pathways to better understand biotic interactions. [ABSTRACT FROM AUTHOR]

Unraveling the biogeographic pattern of soil fungal decomposers along temperature gradients—in smooth linearity or an abrupt jump—can help us connect the global carbon cycle to global warming. Through a standardized global field survey, we identify the existence of temperature thresholds that control the global distribution of soil fungal decomposers, leading to abrupt reductions in their proportion (i.e., the relative abundance in the fungal community) immediately after crossing particular air and soil temperature thresholds. For example, small increases over the mean annual temperature threshold of ~9°C result in abrupt reductions in their proportion, paralleling a similar temperature threshold for soil carbon content. We further find that the proportion of soil fungal decomposers is more sensitive to temperature increases under arid conditions. Given the positive correlation between the global distributions of fungal decomposers and soil heterotrophic respiration, the reported temperature‐driven abrupt reductions in fungal decomposers could further suppress their driven soil decomposition processes and reduce carbon fluxes from soils to the atmosphere with implications for climate change feedback. This work not only advances the current knowledge on the global distribution of soil fungal decomposers, but also highlights that small changes in temperature around certain thresholds can lead to potential unexpected consequences in global carbon cycling under projected climate change. [ABSTRACT FROM AUTHOR]

With ongoing climate change, the probability of crossing environmental thresholds promoting abrupt changes in ecosystem structure and functioning is higher than ever. In drylands (areas where it rains <65% of what could be potentially evaporated), recent research has shown how the crossing of three aridity thresholds [at aridity (1‐Aridity Index) values of 0.54, 0.70 and 0.80] leads to abrupt changes on ecosystem structural and functional attributes. Despite the importance of these findings and their implications to develop effective monitoring and adaptation actions to combat climate change and desertification, we lack a proper understanding of the mechanisms unleashing these abrupt shifts.Here we review multiple mechanisms that may explain the existence of aridity thresholds observed across global drylands, and discuss the potential amplification mechanisms that may underpin hypothetical abrupt temporal shifts with climate change.We propose that each aridity threshold is caused by different and specific mechanisms. The first threshold is mainly caused by physiological mechanisms of plant adaptation to water shortages. The second threshold is unleashed by different mechanisms involving soil processes and plant–soil interactions such as soil erosion, plant community shifts and nutrient cycling and circulation. The collapse of vegetation observed once the third aridity threshold (0.8) is crossed is caused by the mechanisms related to the survival limits of plants that may cause sudden cover and diversity losses and plant–atmospheric feedbacks that link vegetation collapse with further climate aridification.By identifying, revising and linking relevant mechanisms to each aridity threshold observed, we provide a set of specific hypotheses and identify knowledge gaps concerning the study of threshold emergence in drylands. We were also able to establish plausible factors that are context dependent and may influence the occurrence of abrupt ecosystem changes in time. Our review may help to focus future research efforts on aridity thresholds and to develop strategies to monitor, adapt to or even revert abrupt ecosystem changes across global drylands. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Summary: Despite their extent and socio‐ecological importance, a comprehensive biogeographical synthesis of drylands is lacking. Here we synthesize the biogeography of key organisms (vascular and nonvascular vegetation and soil microorganisms), attributes (functional traits, spatial patterns, plant–plant and plant–soil interactions) and processes (productivity and land cover) across global drylands. These areas have a long evolutionary history, are centers of diversification for many plant lineages and include important plant diversity hotspots. This diversity captures a strikingly high portion of the variation in leaf functional diversity observed globally. Part of this functional diversity is associated with the large variation in response and effect traits in the shrubs encroaching dryland grasslands. Aridity and its interplay with the traits of interacting plant species largely shape biogeographical patterns in plant–plant and plant–soil interactions, and in plant spatial patterns. Aridity also drives the composition of biocrust communities and vegetation productivity, which shows large geographical variation. We finish our review by discussing major research gaps, which include: studying regular vegetation spatial patterns; establishing large‐scale plant and biocrust field surveys assessing individual‐level trait measurements; knowing whether the impacts of plant–plant and plant–soil interactions on biodiversity are predictable; and assessing how elevated CO2 modulates future aridity conditions and plant productivity. [ABSTRACT FROM AUTHOR]

Rift Valley fever phlebovirus (RVFV) causes an emerging zoonotic disease and is mainly transmitted by Culex and Aedes mosquitoes. While Aedes aegypti-dengue virus (DENV) is the most studied model, less is known about the genes involved in infection-responses in other mosquito-arboviruses pairing. The main objective was to investigate the molecular responses of Cx. pipiens to RVFV exposure focusing mainly on genes implicated in innate immune responses. Mosquitoes were fed with blood spiked with RVFV. The fully-engorged females were pooled at 3 different time points: 2 hours post-exposure (hpe), 3- and 14-days post-exposure (dpe). Pools of mosquitoes fed with non-infected blood were also collected for comparisons. Total RNA from each mosquito pool was subjected to RNA-seq analysis and a de novo transcriptome was constructed. A total of 451 differentially expressed genes (DEG) were identified. Most of the transcriptomic alterations were found at an early infection stage after RVFV exposure. Forty-eight DEG related to immune infection-response were characterized. Most of them were related with the RNAi system, Toll and IMD pathways, ubiquitination pathway and apoptosis. Our findings provide for the first time a comprehensive view on Cx. pipiens-RVFV interactions at the molecular level. The early depletion of RNAi pathway genes at the onset of the RVFV infection would allow viral replication in mosquitoes. While genes from the Toll and IMD immune pathways were altered in response to RVFV none of the DEG were related to the JAK/STAT pathway. The fact that most of the DEG involved in the Ubiquitin-proteasome pathway (UPP) or apoptosis were found at an early stage of infection would suggest that apoptosis plays a regulatory role in infected Cx. pipiens midguts. This study provides a number of target genes that could be used to identify new molecular targets for vector control. Author summary: Rift valley fever (RVF) is an emerging zoonotic disease and it is caused by RVFV. This virus is commonly transmitted in endemic areas between wild ruminants and mosquitoes, mainly by mosquitoes of Culex and Aedes genus. Starting from the year 2000, several outbreaks have been reported outside Sub Saharan Africa, in countries facing the Mediterranean Sea (Egypt), or Yemen and Saudi Arabia. Available vaccines for ruminants present limited efficacy or residual pathogenic effects. Consequently, new strategies are urgently required to limit the expansion of this zoonotic virus. The main objective of this work is to investigate transcriptional alterations of Cx. pipiens to RVFV focusing mainly on genes implicated in conventional innate immunity pathways, RNAi mechanisms and the apoptotic process in order to evaluate the involvement of these genes in viral infection. The immune altered genes here described could be potential targets to control RVFV infection in mosquitoes. Some of the genes related to the immune defense response were previously described in others mosquito-arbovirus models, as well as in Drosophila and human. To our knowledge, this study highlights for the first time the Cx. pipiens-RVFV interactions in terms of defense infection-response and provides information for developing in the future new approaches to prevent and control the expansion of the virus. [ABSTRACT FROM AUTHOR]

The spatial arrangement of perennial vegetation is critical for ecosystem function in drylands. While much is known about how vegetation patches respond to grazing and abiotic conditions, the size dynamics of individual plants is mostly limited to theoretical studies. We measured the size distribution (mean, variance, skewness) and density of individual grasses, and grass species composition at 451 sites spanning a range of grazing intensities across three broad vegetation communities in semi‐arid eastern Australia. We assessed the relative role of grazing by livestock (cattle and sheep), native (kangaroos) and introduced (rabbits) free ranging herbivores, and several environmental measures (productivity, diversity, composition and groundstorey plant cover) on the size distribution and density of individual grasses. We found mean grass size and density were more sensitive to shifts in grazing intensity and environmental conditions than size variance or the frequency of the smallest individuals (skewness), and shifts were mostly driven by site productivity and cattle and kangaroo grazing. Sheep grazing only reduced mean grass size, and rabbit grazing had no consistent effects. Importantly, we found that site productivity and species composition altered the impacts of grazing on grass density and size distribution. For example, increasing cattle grazing led to larger grasses in low productivity sites. It also led to larger, denser, more variable‐sized grasses among grass species from sites with finer soil texture. Increasing kangaroo grazing led to smaller, denser individuals among grass species from sites with coarse soil texture. At high diversity sites kangaroo grazing led to denser, more homogenised grass sizes with a lower frequency of small individuals. Understanding the in situ response of individual plant sizes gives us insights into the processes driving shifts in perennial vegetation patchiness, improving our ability to predict how the spatial arrangement of ecosystems might change under global change scenarios. [ABSTRACT FROM AUTHOR]