Warming reverses directionality in the richness-abundance relationship in ephemeral Mediterranean plant communities.

Recent findings in forests worldwide have demonstrated how directionality in the richness-abundance causality shifts along global climate gradients: The so-called more-species hypothesis (richness determines abundance) prevails in Earth's most productive climates, whereas the opposite, the so-called more-individuals hypothesis (abundance determines richness), is more likely to prevail in climatically harsh conditions. Since temporal variability is the norm, a critical question is whether this directionality shift is also a function of temporal climatic fluctuations locally. Here, we analyze whether directionality in the richness-abundance relationship is contingent on temporal variability over 10 annual consecutive realizations in ephemeral plant assemblages. Our results support the idea that the more-species hypothesis prevailed in the most benign years, whereas the more-individuals hypothesis did so during less productive years, which were significantly linked to the warmest years. These results support the idea that rising temperatures can reverse directionality in the richness-abundance relationship in these annual plant communities, and therefore, climate warming can have a significant effect on the relationship between diversity and ecosystem functions, such as productivity, by altering the prevalence of primary mechanisms involved in species assembly.

Recent and ancient evolutionary events shaped plant elemental composition of edaphic endemics: a phylogeny-wide analysis of Iberian gypsum plants.

The analysis of plant elemental composition and the underlying factors affecting its variation are a current hot topic in ecology. Ecological adaptation to atypical soils may shift plant elemental composition. However, no previous studies have evaluated its relevance against other factors such as phylogeny, climate or individual soil conditions. We evaluated the effect of the phylogeny, environment (climate, soil), and affinity to gypsum soils on the elemental composition of 83 taxa typical of Iberian gypsum ecosystems. We used a new statistical procedure (multiple phylogenetic variance decomposition, MPVD) to decompose total explained variance by different factors across all nodes in the phylogenetic tree of target species (covering 120 million years of Angiosperm evolution). Our results highlight the relevance of phylogeny on the elemental composition of plants both at early (with the development of key preadaptive traits) and recent divergence times (diversification of the Iberian gypsum flora concurrent with Iberian gypsum deposit accumulation). Despite the predominant phylogenetic effect, plant adaptation to gypsum soils had a strong impact on the elemental composition of plants, particularly on sulphur concentrations, while climate and soil effects were smaller. Accordingly, we detected a convergent evolution of gypsum specialists from different lineages on increased sulphur and magnesium foliar concentrations.

Diverse phylogenetic neighborhoods enhance community resistance to drought in experimental assemblages.

Although the role played by phylogeny in the assembly of plant communities remains as a priority to complete the theory of species coexistence, experimental evidence is lacking. It is still unclear to what extent phylogenetic diversity is a driver or a consequence of species assembly processes. We experimentally explored how phylogenetic diversity can drive the community level responses to drought conditions in annual plant communities. We manipulated the initial phylogenetic diversity of the assemblages and the water availability in a common garden experiment with two irrigation treatments: average natural rainfall and drought, formed with annual plant species of gypsum ecosystems of Central Spain. We recorded plant survival and the numbers of flowering and fruiting plants per species in each assemblage. GLMMs were performed for the proportion of surviving, flowering, fruiting plants per species and for total proportion of surviving species and plants per pot. In water limited conditions, high phylogenetic diversity favored species coexistence over time with higher plant survival and more flowering and fruiting plants per species and more species and plants surviving per pot. Our results agree with the existence of niche complementarity and the convergence of water economy strategies as major mechanisms for promoting species coexistence in plant assemblages in semiarid Mediterranean habitats. Our findings point to high phylogenetic diversity among neighboring plants as a plausible feature underpinning the coexistence of species, because the success of each species in terms of surviving and producing offspring in drought conditions was greater when the initial phylogenetic diversity was higher. Our study is a step forward to understand how phylogenetic relatedness is connected to the mechanisms determining the maintenance of biodiversity.

Effect of aridity on species assembly in gypsum drylands: a response mediated by the soil affinity of species.

Previous studies found that plant communities on infertile soils are relatively resistant to climatic variation due to stress tolerance adaptations. However, the species assemblies in gypsum soil habitats require further investigation. Thus, we considered the following questions. (1) Do harsher arid conditions determine the characteristics of the species that form plant assemblages? (2) Is the selection of the species that assemble in arid conditions mediated by their ability to grow on gypsum soils? (3) Is the selection of species that assemble in harsher conditions related to phylogenetically conserved functional traits? Perennial plant communities were analysed in 89 gypsum-soil sites along a 400 km climate gradient from the central to southeastern Iberian Peninsula. Each local assemblage was analysed in 30 × 30 m plots and described based on taxonomic, functional (soil plant affinity) and phylogenetic parameters. The mean maximum temperatures in the hottest month, mean annual precipitation and their interaction terms were used as surrogates for the aridity conditions in generalized linear models. In the hottest locations, the gypsophily range narrowed and the mean gypsophily increased at the community level, thereby suggesting the filtering of species and the dominance of soil specialists in the actual plant assemblies. Drier sites had higher taxonomic diversity. The species that formed the perennial communities were close in evolutionary terms at the two ends of the aridity gradient. The mean maximum temperatures in the hottest month had the main abiotic filtering effect on perennial plant communities, which was mediated by the ability of species to grow on gypsum soils, and thus gypsum specialists dominated the species assemblies in the hottest locations. In contrast, the perennial communities on gypsum soils were relatively resistant to changes in precipitation. Our findings suggest that the warmer environmental conditions predicted by global change models will favour gypsum specialists over generalists.

Impacts of climate, soil and biotic interactions on the interplay of the different facets of alpine plant diversity.

Disentangling the processes that drive plant community assembly is critical for understanding the patterns of plant diversity. We studied how different abiotic and biotic factors shape the interplay between the facets of alpine plant diversity, functional (FD), phylogenetic (PD) and taxonomic diversity (TD), in three different mountain ranges with contrasting evolutionary histories and climate conditions (Pyrenees and Mediterranean-type mountains in central Spain and Chilean Andes). We hypothesized that the causal links vary in strength and sign across regions. We used species inventories, functional trait data, and a phylogeny from 84 plant communities spread throughout three high-mountain alpine grasslands. Structural equation models were used to test our causal hypotheses on the relationships observed between the three diversity facets, and the abiotic (elevation, potential solar radiation and soil total nitrogen) and biotic factors (C-score). Despite our causal model presented a high variability in each mountain range, TD always decreased with increasing elevation (sum of direct and indirect effects). We also found some patterns suggesting that assembly processes could be climatically/biogeographically structured such as the negative relationship between FD and elevation found in Mediterranean mountains and the negative relationship between FD and TD found in both Spanish mountain ranges (independently of their different climates). A remarkable finding of this study is that ecological factors such as soil total nitrogen and elevation indirectly alter the relationships between the diversity facets. Our results suggest that diversity facets are simultaneously affected by different ecological and biogeographical/evolutionary processes, resulting in some general trends but also in parallel idiosyncratic patterns. Our findings highlight that although FD stand out by its explanatory power of community processes, TD and PD provide a complementary and necessary view that should not be disregarded in the attempt to globally explain community assembly processes.

Positive associations among rare species and their persistence in ecological assemblages.

According to the competitive exclusion principle, species with low competitive abilities should be excluded by more efficient competitors; 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. On the other hand, 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 probable mechanisms underpinning this pattern and rare species persistence. The consistent results across taxa and geography suggest a general explanation for the maintenance of biodiversity in competitive environments.

Alpine vegetation dataset from three contrasting mountain ranges differing in climate and evolutionary history.

Vegetation above treeline constitutes one of the most vulnerable ecosystems to climate warming and other drivers of Global Change. Given the panorama of such an uncertain future facing these plant communities, it is critical to know how they respond to environmental changes and improve the knowledge on the potential impacts of climate change on their distribution. Recently, with the impressive development of trait-based approaches, relevant progress has been made to better understand the relationships between environmental conditions and plant communities. In this data paper, we describe data on abundances of 327 alpine plant species across 430 subplots of 2.4 m × 2.4 m in three mountain ranges (Sierra de Guadarrama and Pyrenees in Spain, and the Central Andes in Chile). We provide data on different environmental variables that represent variation in abiotic conditions and operate at different spatial scales (e.g., climatic, topographic and soil conditions). Data on six plant functional traits are also shown, which were measured on ten individuals of each species, following standard protocols. We provided the dataset as tables in the supplementary material. This information could be used to analyse the relationship between the alpine vegetation and changes in environmental conditions, and ultimately, to understand ecosystem functioning and guide conservation strategies of theses threatened and valuable ecosystems.

Winter is coming: plant freezing resistance as a key functional trait for the assembly of annual Mediterranean communities.

Background and Aims: In Mediterranean annual plants, germination mainly occurs during the autumn and only those seedlings that survive winter freezing can flower and produce seedlings in spring. Surprisingly, the effect of freezing events as an abiotic determinant of these communities remains unexplored. The present study aimed to investigate how freezing events affect annual Mediterranean communities and whether their functional structure as related to freezing resistance is linked to the main biotic and abiotic determinants of these communities. Methods: In 120 plots located on a semi-arid Mediterranean steppe (Spain), the community functional structure related to the lethal temperature causing 50 % frost damage (LT50 trait) in seedlings was estimated and summarized as the community-weighted mean (CWM-LT50) and its functional diversity (FD-LT50). Plots were stratified according to distance to rabbit shelters and latrines as a proxy for rabbit density, proximity to Stipa tenacissima and spring water availability, where annual species abundance was recorded in all plots over three consecutive years. Key Results: Annual species were able to resist a threshold temperature of -4 °C and most had LT50 values around the absolute minimum temperature (-9.5 °C) in the three years. Higher rabbit densities led to lower CWM-LT50 and higher FD-LT50 values. Plots close to Stipa tussocks had higher CWM-LT50 values whereas water availability had no effects. Conclusions: High freezing resistance was extended among winter annual species, suggesting the presence of an association between historical environmental filtering and low winter temperatures. However, the community functional structure related to freezing resistance remained variable among scenarios with differences in herbivory pressure and distance to perennial vegetation. The trends observed indicate that traits that allow plants to deal with herbivory may also promote freezing resistance, and that tussocks can act as nurses via microclimatic amelioration of harsher winter conditions.

Factors driving species assemblage in Mediterranean soil seed banks: from the large to the fine scale.

BACKGROUND AND AIMS: Many studies have analysed the mechanisms that determine plant coexistence in standing vegetation, but the determinants of soil seed bank species assemblies have rarely been studied. In gypsum soil communities, aerial vegetation and seed banks are tightly connected in space and time, but the mechanisms involved in their organization may differ. The aim of this study is to understand the relative importance of biotic and abiotic factors controlling soil seed bank composition and structure. METHODS: Persistent and complete (i.e. persistent plus transient) soil seed banks were investigated at two spatial scales in a very species-rich semi-arid community dominated by annuals. A water addition treatment equivalent to 50 % annual increase in average precipitation (abiotic factor) was applied for two consecutive years, and the relationships of the soil seed bank to the biological soil crust (BSC), above-ground vegetation and the presence of Stipa tenacissima tussocks (biotic factors) were simultaneously evaluated. KEY RESULTS: As expected, the standing vegetation was tightly related to seed abundance, species richness and composition in both seed banks. Remarkably, BSC cover was linked to a decrease in seed abundance and species richness in the persistent seed bank, and it even determined complete seed bank composition at the fine spatial scale. However, this effect disappeared at coarser scales, probably because of the high spatial heterogeneity induced by BSCs. In contrast to findings on standing vegetation, Stipa and the irrigation treatment for two consecutive years had no effect on soil seed banks. CONCLUSIONS: Soil seed bank assemblies in our semi-arid plant community were the result of above-ground vegetation dynamics and of the direct filtering processes on seed fate operated by the spatially heterogeneous BSCs. Cover of BSCs was negatively correlated with seed abundance and species richness, and affected seed species composition in the soil. Changes in species composition and enrichment when the BSC cover is low suggest that BSCs promote a fine scale niche differentiation in the soil seed bank and thereby potentially enhance species coexistence and high species diversity in these communities.

Plant life on gypsum: a review of its multiple facets.

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.

Assemblage of a semi-arid annual plant community: abiotic and biotic filters act hierarchically.

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.

Shrub encroachment can reverse desertification in semi-arid Mediterranean grasslands.

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.

Environmental scales on the reproduction of a gypsophyte: a hierarchical approach.

BACKGROUND AND AIMS: Environmental variability at several scales can determine plant reproductive success. The main goal of this work was to model the reproductive flexibility of a semi-arid specialist considering different scales of environmental variability. METHODS: A 2-year field study was performed on the determinants of the female reproductive success of Helianthemum squamatum, an Iberian gypsophyte, considering two scales of environmental variability: differences between two contrasting slope aspects; and, on individual scale, the neighbouring microenvironment. Generalized linear mixed models were used to evaluate simultaneously the potential effects of environmental variability at both scales, together with flowering phenology and plant size on the reproductive output of H. squamatum. The following reproductive response variables were considered: number of flowers, fruit-set, number of viable and aborted seeds per fruit, and number of seeds per plant. KEY RESULTS: Contrary to expectations, environmental variability exerted a weak or even absent effect on the reproductive variables considered, while flowering phenology and plant size, which did not vary between slopes, played a major role. Surprisingly, the absolute reproductive variables were even higher in the extremely dry year of 2003, although only on the south-facing slope. The relatively milder conditions of the north-facing slope did not involve any advantage to this species in terms of reproductive output. CONCLUSIONS: The species seemed to be considerably well adapted to the environmental unpredictability characteristic of Mediterranean systems, considering its ability to maintain reproduction across contrasting environments and contrasting climatic conditions. These findings make us face the question of what must be considered stressful conditions in the case of a stress-tolerant specialist.