The worldwide phenomenon of shrub encroachment in grass-dominated dryland ecosystems is commonly associated with desertification. Studies of the purported desertification effects associated with shrub encroachment are often restricted to relatively few study areas, and document a narrow range of possible impacts upon biota and ecosystem processes. We conducted a study in degraded Mediterranean grasslands dominated by Stipa tenacissima to simultaneously evaluate the effects of shrub encroachment on the structure and composition of multiple biotic community components, and on various indicators of ecosystem function. Shrub encroachment enhanced vascular plant richness, biomass of fungi, actinomycetes and other bacteria, and was linked with greater soil fertility and N mineralization rates. While shrub encroachment may be a widespread phenomenon in drylands, an interpretation that this is an expression of desertification is not universal. Our results suggest that shrub establishment may be an important step in the reversal of desertification processes in the Mediterranean region.

The adaptation of plants to particular soil types has long intrigued biologists. Gypsum soils occupy large areas in many regions of the world and host a striking biological diversity, but their vegetation has been much less studied than that developing over serpentine or saline soils. Herein, we review all aspects of plant life on gypsum ecosystems, discuss the main processes driving their structure and functioning, and highlight the main conservation threats that they face. Plant communities in gypsum habitats typically show distinctive bands at very small spatial scales, which are mainly determined by topography. Plants living on gypsum soils can be classified into three categories: (i) wide gypsophiles are specialists that can penetrate the physical soil crust during early life stages and have physiological adjustments to cope with the chemical limitations imposed by gypsum soils; (ii) narrow gypsophiles are refugee plants which successfully deal with the physical soil crust and can tolerate these chemical limitations but do not show specific adaptations for this type of soils; and (iii) gypsovags are non-specialist gypsum plants that can only thrive in gypsum soils when the physical crust is absent or reduced. Their ability to survive in gypsum soils may also be mediated by below-ground interactions with soil microorganisms. Gypsophiles and gypsovags show efficient germination at low temperatures, seed and fruit heteromorphism within and among populations, and variation in seed dormancy among plants and populations. In gypsum ecosystems, spatio-temporal changes in the composition and structure of above-ground vegetation are closely related to those of the soil seed bank. Biological soil crusts (BSCs) dominated by cyanobacteria, lichens and mosses are conspicuous in gypsum environments worldwide, and are important drivers of ecosystem processes such as carbon and nitrogen cycling, water infiltration and run-off and soil stability. These organisms are also important determinants of the structure of annual plant communities living on gypsum soils. The short-distance seed dispersal of gypsophiles is responsible for the high number of very narrow endemisms typically found in gypsum outcrops, and suggests that these species are evolutionarily old taxa due to the time they need to colonize isolated gypsum outcrops by chance. Climate change and habitat fragmentation negatively affect both plants and BSCs in gypsum habitats, and are among the major threats to these ecosystems. Gypsum habitats and specialists offer the chance to advance our knowledge on restrictive soils, and are ideal models not only to test important evolutionary questions such as tolerance to low Ca/Mg proportions in soils, but also to improve the theoretical framework of community ecology and ecosystem functioning.

The study of species coexistence and community assembly has been a hot topic in ecology for decades. Disentangling the hierarchical role of abiotic and biotic filters is crucial to understand community assembly processes. The most critical environmental factor in semi-arid environments is known to be water availability, and perennials are usually described as nurses that create milder local conditions and expand the niche range of several species. We aimed to broaden this view by jointly evaluating how biological soil crusts (BSCs), water availability, perennial species (presence/absence of Stipa tenacissima) and plant-plant interactions shape a semi-arid annual plant community. The presence and cover of annual species was monitored during three years of contrasting climate. Water stress acted as the primary filter determining the species pool available for plant community assembly. Stipa and BSCs acted as secondary filters by modulating the effects of water availability. At extremely harsh environmental conditions, Stipa exerted a negative effect on the annual plant community, while at more benign conditions it increased annual community richness. Biological soil crusts exerted a contradictory effect depending on climate and on the presence of Stipa, favoring annuals in the most adverse conditions but showing repulsion at higher water availability conditions. Finally, interactions among co-occurring annuals shaped species richness and diversity of the final annual plant assembly. This study sheds light on the processes determining the assembly of annual communities and highlights the importance of Biological Soil Crusts and of interactions among annual plants on the final outcome of the species assembly.

Pioneer plants are adapted to grow in unpredictable environments. These plants have evolved several traits related to seed morphology and germination to cope with this unpredictability. The main aim of this study was to investigate the effect of plant identity and maternal environment on seed mass and germination of Sinapis arvensis. This study was undertaken with S. arvensis plants grown in four experimentally manipulated resource availability scenarios: (i) nitrogen addition, (ii) water addition, (iii) nitrogen and water simultaneously added, and (iv) control. Plants grew under field conditions in an emerging plant community. Germination response was mainly affected by maternal plant identity. This reflects not only genotype quality but also phenotypic plasticity, which allows adaptation to habitat conditions. Seed coat colour together with mean seed mass per plant explained significant fractions of the variability in germination rate. Changes in the germination response because of maternal environmental effects were also detected. Germination rate decreased in seeds from nitrogen-enriched conditions; this may be due to the induction of dormancy caused by high concentration of nitrogen in the seeds. Addition of water to maternal environment caused a decrease in total germination percentage and germination rate of seeds. Thus, our results provide evidence for maternal environmental effects on germination under field conditions. Furthermore, changes in germination timing, as detected in our study, may determine plant development and fitness in unpredictable conditions.

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According to the competitive exclusion principle, species with low competitive abilities should be excluded by more efficient competitors, and yet they generally remain as rare species. Here, we describe the positive and negative spatial association networks of 326 disparate assemblages, showing a general organization pattern that simultaneously supports the primacy of competition and the persistence of rare species. Abundant species monopolize negative associations in about 90% of the assemblages. Contrarily, rare species are mostly involved in positive associations, forming small network modules. Simulations suggest that positive interactions among rare species and microhabitat preferences are the most likely mechanisms underpinning this pattern and rare species persistence. The

This paper reports a study on species richness and composition of Tumbesian dry forest communities. We tested two alternative hypotheses about species assemblage processes in tropical dry forests: (1) species assemblage is determined by the filtering effect of environmental conditions and (2) species assemblage is determined by facilitative processes along the gradient of water availability, and thus, species richness and evenness increase as water becomes limited. In addition, we also explored the effect of climate and soil conditions on species composition in tropical dry forests. Species composition was sampled in 109 plots in terms of cover and tree diameter at breast height. Climatic, edaphic, topographic and anthropogenic degradation variables were obtained for each plot. We used generalized linear models and canonical correspondence analyses to evaluate the effect of environmental variables on species composition, richness and evenness. Water availability negatively affected richness and significantly determined the species assemblage. Species richness increased from ridges to valleys and evenness increased at higher altitudes. Soil characteristics showed no effect on richness and evenness but soil moisture, nitrogen concentration and soil temperature explained significant fractions of species composition. Although timber extraction and livestock in our study area were of low intensity, it negatively affected richness but had only a minor effect on species composition. Our results suggest that species composition in these endangered tropical dry forests may be at least partially explained by the stress-gradient hypothesis, with higher species richness at drier conditions probably induced by facilitation processes.

1. Understanding the mechanisms that underlie species assembly is a central concern in community ecology. Abiotic and biotic filters are probabilistic 'sieves' that allow species with certain functional traits to become a part of the community, or not. We manipulated natural plant assemblies in order to identify variations in the timings of biotic and abiotic filters that determine community trait assemblies.2. We extracted soil portions when the investigated annual plant community was in its seed phase ('community monolith'), thereby maintaining the structure and similar soil characteristics to the field conditions. Community monoliths were subjected to experimental manipulation in terms of the rainfall timing and amount, and perturbations of the biological soil crust (BSC; intact vs. perturbed). We surveyed the experimental community assembly over time based on the functional diversity by considering important functional traits in different life stages.3. We found that autumn droughts acted as abiotic filters by favouring the germination and establishment of species with greater investment in the root biomass.Under severe droughts (66% water reduction), the experimental assemblies were dominated by species with functional traits adapted to water shortage conditions: high leaf dry matter content, low specific leaf area, small individual size, low reproductive ratio and high root:shoot ratio. We identified two roles of BSCs in annual plant species assemblies: (a) as a biotic filter that limited the establishment of species based on seed size, and (b) as a buffer against water stress conditions by reducing soil evapotranspiration. Synthesis.We demonstrated the importance of the timing and amount of rainfall for shaping annual plant communities, and identified germination filters as the main process that determined community assemblies. Our results suggest that the phenotypic integration of functional traits facilitates resistance to drought during the life cycle. The BSC-annual plant relationship shifted from negative, by acting as a germination filter, to positive, by acting as a buffer in later stages.Climatic fluctuations and fine scale biotic determinants of spatial heterogeneity emerged as sources of changes in the community assembly in time and space to