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Earth harbours an extraordinary plant phenotypic diversity 1 that is at risk from ongoing global changes 2,3 . However, it remains unknown how increasing aridity and livestock grazing pressure-two major drivers of global change 4-6 -shape the trait covariation that underlies plant phenotypic diversity 1,7 . 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 diversity 8-10 . 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., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Whereas temporal variability of plant phenology in response to climate change has already been well studied, the spatial variability of phenology is not well understood. Given that phenological shifts may affect biotic interactions, there is a need to investigate how the variability in environmental factors relates to the spatial variability in herbaceous species' phenology by at the same time considering their functional traits to predict their general and species-specific responses to future climate change. In this project, we analysed phenology records of 148 herbaceous species, which were observed for a single year by the PhenObs network in 15 botanical gardens. For each species, we characterised the spatial variability in six different phenological stages across gardens. We used boosted regression trees to link these variabilities in phenology to the variability in environmental parameters (temperature, latitude and local habitat conditions) as well as species traits (seed mass, vegetative height, specific leaf area and temporal niche) hypothesised to be related to phenology variability. We found that spatial variability in the phenology of herbaceous species was mainly driven by the variability in temperature but also photoperiod was an important driving factor for some phenological stages. In addition, we found that early-flowering and less competitive species characterised by small specific leaf area and vegetative height were more variable in their phenology. Our findings contribute to the field of phenology by showing that besides temperature, photoperiod and functional traits are important to be included when spatial variability of herbaceous species is investigated., (© 2024. The Author(s).)

Invasive plants typically escape specialist herbivores but are often attacked by generalist herbivores in their introduced ranges. The shifting defense hypothesis suggests that this will cause invasive plants to evolve lower resistance against specialists, higher resistance against generalists, and greater tolerance to herbivore damage. However, the duration and direction of selective pressures can shape the evolutionary responses of resistance and tolerance for invasive plants. Two critical factors are (1) residence time (length of time that an invasive species has been in its introduced range) and (2) specialist herbivore reassociation (attack by purposely or accidentally introduced specialists). Yet, these two factors have not been considered simultaneously in previous quantitative syntheses. Here, we performed a meta-analysis with 367 effect sizes from 70 studies of 35 invasive plant species from native and invasive populations. We tested how the residence time of invasive plant species and specialist reassociation in their introduced ranges affected evolutionary responses of defenses against specialists and generalists, including herbivore resistance traits (physical barriers, digestibility reducers and toxins), resistance effects (performance of and damage caused by specialists or generalists) and tolerance to damage (from specialists or generalists). We found that residence time and specialist reassociation each significantly altered digestibility reducers, specialist performance, generalist damage, and tolerance to specialist damage. Furthermore, residence time and specialist reassociation strongly altered toxins and generalist performance, respectively. When we restricted consideration to invasive plant species with both longer residence times and no reassociation with specialists, invasive populations had lower resistance to specialists, similar resistance to generalists, and higher tolerance to damage from both herbivore types, compared with native populations. We conclude that the duration and direction of selective pressure shape the evolutionary responses of invasive plants. Under long-term (long residence time) and stable (no specialist reassociation) selective pressure, invasive plants generally decrease resistance to specialists and increase tolerance to generalist damage that provides mixed support for the shifting defense hypothesis., (© 2024 The Ecological Society of America.)

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought., Competing Interests: Competing interests statement:The authors declare no competing interest.

Belowground niche partitioning presents a key mechanism for maintaining species coexistence and diversity. Its importance is currently reinforced by climate change that alters soil hydrological conditions. However, experimental tests examining the magnitude of its change under climate change are scarce. We combined measurements of oxygen stable isotopes to infer plant water-uptake depths and extreme drought manipulation in grasslands. Belowground niche partitioning was evidenced by different water-uptake depths of co-occurring species under ambient and extreme drought conditions despite an increased overlap among species due to a shift to shallower soil layers under drought. A co-occurrence of contrasting strategies related to the change of species water-uptake depth distribution was likely to be key for species to maintain some extent of belowground niche partitioning and could contribute to stabilizing coexistence under drought. Our results suggest that belowground niche partitioning could mitigate negative effects on diversity imposed by extreme drought under future climate., (© 2023 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Climate change is continuously intensifying droughts. Extreme droughts are expected to reduce soil water content and thus, ecosystem functioning such as above-ground primary productivity. Nonetheless, results of experimental drought studies vary from no impact to a significant decrease in soil water content and/or productivity. We experimentally imposed extreme drought as 30 % and 50 % precipitation reductions using rainout shelters for four years in temperate grasslands and in the forest understory. We studied the concurrent impact of two intensities of extreme drought on the soil water content and above-ground primary productivity in the last experimental year (resistance). Furthermore, we observed resilience as the extent to which both variables differ from ambient conditions after the removal of the 50 % reduction. We show a systematic difference in response to extreme experimental drought between grasslands and the forest understory irrespective of the intensity of the extreme drought. Namely, extreme drought resulted in a significant decrease of the soil water content and productivity in grasslands but not in the forest understory. Interestingly, the negative impacts in the grasslands did not persist as evidenced by the fact that soil water content and productivity were similar to ambient conditions after the removal of the drought. Our results indicate that extreme drought on small spatial scales does not necessarily result in a concurrent soil water decrease in the forest understory, while this is the case for grasslands, with respective consequences for the resistance of productivity. Grasslands, however, can be resilient. Our study highlights that considering the response of the soil water content is key to understanding divergent productivity responses to extreme drought among different ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Plant recruitment interactions (i.e., what recruits under what) shape the composition, diversity, and structure of plant communities. Despite the huge body of knowledge on the mechanisms underlying recruitment interactions among species, we still know little about the structure of the recruitment networks emerging in ecological communities. Modeling and analyzing the community-level structure of plant recruitment interactions as a complex network can provide relevant information on ecological and evolutionary processes acting both at the species and ecosystem levels. We report a data set containing 143 plant recruitment networks in 23 countries across five continents, including temperate and tropical ecosystems. Each network identifies the species under which another species recruits. All networks report the number of recruits (i.e., individuals) per species. The data set includes >850,000 recruiting individuals involved in 118,411 paired interactions among 3318 vascular plant species across the globe. The cover of canopy species and open ground is also provided. Three sampling protocols were used: (1) The Recruitment Network (RN) protocol (106 networks) focuses on interactions among established plants ("canopy species") and plants in their early stages of recruitment ("recruit species"). A series of plots was delimited within a locality, and all the individuals recruiting and their canopy species were identified; (2) The paired Canopy-Open (pCO) protocol (26 networks) consists in locating a potential canopy plant and identifying recruiting individuals under the canopy and in a nearby open space of the same area; (3) The Georeferenced plot (GP) protocol (11 networks) consists in using information from georeferenced individual plants in large plots to infer canopy-recruit interactions. Some networks incorporate data for both herbs and woody species, whereas others focus exclusively on woody species. The location of each study site, geographical coordinates, country, locality, responsible author, sampling dates, sampling method, and life habits of both canopy and recruit species are provided. This database will allow researchers to test ecological, biogeographical, and evolutionary hypotheses related to plant recruitment interactions. There are no copyright restrictions on the data set; please cite this data paper when using these data in publications., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Under climate change, extreme droughts will limit water availability for plants. However, the species-specific responses make it difficult to draw general conclusions. We hypothesized that changes in species' abundance in response to extreme drought can be best explained by a set of water economic traits under ambient conditions in combination with the ability to adjust these traits towards higher drought resistance. We conducted a 4-year field experiment in temperate grasslands using rainout shelters with 30% and 50% rainfall reduction. We quantified the response as the change in species abundance between ambient conditions and the rainfall reduction. Abundance response to extreme drought was best explained by a combination of traits in ambient conditions and their functional adjustment, most likely reflecting plasticity. Smaller leaved species decreased less in abundance under drought. With increasing drought intensity, we observed a shift from drought tolerance, i.e., an increase in leaf dry matter content, to avoidance, i.e., a less negative turgor loss point (TLP) in ambient conditions and a constancy in TLP under drought. We stress the importance of using a multidimensional approach of variation in multiple traits and the importance of considering a range of drought intensities to improve predictions of species' response to climate change., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

In recent years, the effects of plastic contamination on soil and plants have received growing attention. Plastic can affect soil water content and thus may interact with the effects of drought on soil and plants. However, the effects of plastic on soil are highly context-dependent, and interactions with drought have been hardly tested. We conducted two greenhouse experiments to test the combined effects of plastic fragments (of varying size and concentration), water availability and soil texture, on soil water content and performance of the plant Arabidopsis thaliana. Plastic fragments had stronger negative effects on soil water content in low water availability, and the shape of this response (linear vs. unimodal) was mediated by soil texture. Conversely, increasing concentration of plastic had positive effects on plant growth. We suggest that plastic fragments introduce fracture points within soil aggregates. This increases number and size of soil pores favoring water loss but also facilitating root growth. Our results suggest complex interactive effects of plastic and drought, that may lead to a decoupling of plant and soil response. These processes should be taken into account in ecological studies and agricultural practices., (© 2022. The Author(s).)

Phenology has emerged as key indicator of the biological impacts of climate change, yet the role of functional traits constraining variation in herbaceous species' phenology has received little attention. Botanical gardens are ideal places in which to investigate large numbers of species growing under common climate conditions. We ask whether interspecific variation in plant phenology is influenced by differences in functional traits. We recorded onset, end, duration and intensity of initial growth, leafing out, leaf senescence, flowering and fruiting for 212 species across five botanical gardens in Germany. We measured functional traits, including plant height, absolute and specific leaf area, leaf dry matter content, leaf carbon and nitrogen content and seed mass and accounted for species' relatedness. Closely related species showed greater similarities in timing of phenological events than expected by chance, but species' traits had a high degree of explanatory power, pointing to paramount importance of species' life-history strategies. Taller plants showed later timing of initial growth, and flowered, fruited and underwent leaf senescence later. Large-leaved species had shorter flowering and fruiting durations. Taller, large-leaved species differ in their phenology and are more competitive than smaller, small-leaved species. We assume climate warming will change plant communities' competitive hierarchies with consequences for biodiversity., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Researchers use both experiments and observations to study the impacts of climate change on ecosystems, but results from these contrasting approaches have not been systematically compared for droughts. Using a meta-analysis and accounting for potential confounding factors, we demonstrate that aboveground biomass responded only about half as much to experimentally imposed drought events as to natural droughts. Our findings indicate that experimental results may underestimate climate change impacts and highlight the need to integrate results across approaches., (© 2022. The Author(s).)

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1-km 2 resolution for 0-5 and 5-15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1-km 2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (-0.7 ± 2.3°C). The observed substantial and biome-specific offsets emphasize that the projected impacts of climate and climate change on near-surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Plants have been shown to change their foraging behaviour in response to resource heterogeneity. However, an unexplored hypothesis is that foraging could be induced by environmental stressors, such as herbivory, which might increase the demand for particular resources, such as those required for herbivore defence. This study examined the way simulated herbivory affects both root foraging for and uptake of cadmium (Cd), in the metal-hyperaccumulating plant Arabidopsis halleri , which uses this heavy metal as herbivore defence. Simulated herbivory elicited enhanced relative allocation of roots to Cd-rich patches as well as enhanced Cd uptake, and these responses were exhibited particularly by plants from non-metalliferous origin, which have lower metal tolerance. By contrast, plants from a metalliferous origin, which are more tolerant to Cd, did not show any preference in root allocation, yet enhanced Cd sharing between ramets when exposed to herbivory. These results suggest that foraging for heavy metals, as well as their uptake and clonal-sharing, could be stimulated in A. halleri by herbivory impact. Our study provides first support for the idea that herbivory can induce not only defence responses in plants but also affect their foraging, resource uptake and clonal sharing responses.

Plants can respond to competition with a myriad of physiological or morphological changes. Competition has also been shown to affect the foraging decisions of plants belowground. However, a completely unexplored idea is that competition might also affect plants' foraging for specific elements required to inhibit the growth of their competitors. In this study, we examined the effect of simulated competition on root foraging and accumulation of heavy metals in the metal hyperaccumulating perennial plant Arabidopsis halleri, whose metal accumulation has been shown to provide allelopathic ability. A. halleri plants originating from both metalliferous and non-metalliferous soils were grown in a "split-root" setup with one root in a high-metal pot and the other in a low-metal one. The plants were then assigned to either simulated light competition or no-competition (control) treatments, using vertical green or clear plastic filters, respectively. While simulated light competition did not induce greater root allocation into the high-metal pots, it did result in enhanced metal accumulation by A. halleri, particularly in the less metal-tolerant plants, originating from non-metalliferous soils. Interestingly, this accumulation response was particularly enhanced for zinc rather than cadmium. These results provide support to the idea that the accumulation of metals by hyperaccumulating plants can be facultative and change according to their demand following competition., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The plant economics spectrum hypothesizes a correlation among resource-use related traits along one single axis, which determines species' growth rates and their ecological filtering along resource gradients. This concept has been mostly investigated and shown in perennial species, but has rarely been tested in annual species. Annuals evade unfavorable seasons as seeds and thus may underlie different constraints, with consequences for interspecific trait-trait, trait-growth, and trait-environment relations. To test the hypotheses of the plant economics spectrum in annual species, we measured twelve resource-use related leaf and root traits in 30 winter annuals from Israel under controlled conditions. Traits and their coordinations were related to species' growth rates (for 19 species) and their distribution along a steep rainfall gradient. Contrary to the hypotheses of the plant economics spectrum, in the investigated annuals traits were correlated along two independent axes, one of structural traits and one of carbon gain traits. Consequently, species' growth rates were related to carbon gain traits, but independent from structural traits. Species' distribution along the rainfall gradient was unexpectedly neither associated with species' scores along the axes of carbon gain or structural traits nor with growth rate. Nevertheless, root traits were related with species' distribution, indicating that they are relevant for species' filtering along rainfall gradients in winter annuals. Overall, our results showed that the functional constraints hypothesized by the plant economics spectrum do not apply to winter annuals, leading to unexpected trait-growth and trait-rainfall relations. Our study thus cautions to generalize trait-based concepts and findings between life-history strategies. To predict responses to global change, trait-based concepts should be explicitly tested for different species groups., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Kurze, Engelbrecht, Bilton, Tielbörger and Álvarez-Cansino.)

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long-term average thermal conditions at coarse spatial resolutions only. Hence, many climate-forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold-air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free-air temperatures, microclimatic ground and near-surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near-surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes., (© 2020 John Wiley & Sons Ltd.)

Facilitation studies typically compare plants under differential stress levels with and without neighbors, while the density of neighbors has rarely been addressed. However, recent empirical studies indicate that facilitation may be density-dependent too and peak at intermediate neighbor densities. Here, we propose a conceptual model to incorporate density-dependence into theory about changes of plant-plant interactions under stress. To test our predictions, we combine an individual-based model incorporating both facilitative response and effect, with an experiment using salt stress and Arabidopsis thaliana. Theoretical and experimental results are strikingly consistent: (1) the intensity of facilitation peaks at intermediate density, and this peak shifts to higher densities with increasing stress; (2) this shift further modifies the balance between facilitation and competition such that the stress-gradient hypothesis applies only at high densities. Our model suggests that density-dependence must be considered for predicting plant-plant interactions under environmental change.

Plant-plant interactions are reciprocal and include effects on and response to neighbours. Distinct traits confer competitive effect and response ability, but how specific traits determine effect and response in facilitative interactions has not been studied experimentally. We utilized the model species Arabidopsis thaliana to test for trait dependence of facilitative interactions. Salt-sensitive (sos) mutants or salt-tolerant wild-types were exposed to an experimental salinity gradient with and without intraspecific neighbours and the intensity of plant-plant interactions was measured for three performance variables. We tested whether salt tolerance can predict facilitative effect and response and whether a tradeoff exists between competitive ability and tolerance to stress. Interactions shifted very clearly from negative to positive with increasing stress. Salt-sensitive genotypes were less negatively affected by competition but more dependent on facilitation than were wild-types, indicating a tradeoff between competitive ability and stress tolerance. Surprisingly, sensitive genotypes imposed stronger facilitative effects, despite being much smaller under stress, probably because they retrieved more salt from the soil. Stress tolerance defined facilitative effect and response via distinct mechanisms. We advocate more controlled experiments with model species to advance our understanding of the trait dependence of biotic interactions and their consequences for community organization., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Admixture is the hybridization between populations within one species. It can increase plant fitness and population viability by alleviating inbreeding depression and increasing genetic diversity. However, populations are often adapted to their local environments and admixture with distant populations could break down local adaptation by diluting the locally adapted genomes. Thus, admixed genotypes might be selected against and be outcompeted by locally adapted genotypes in the local environments. To investigate the costs and benefits of admixture, we compared the performance of admixed and within-population F1 and F2 generations of the European plant Lythrum salicaria in a reciprocal transplant experiment at three European field sites over a 2-year period. Despite strong differences between site and plant populations for most of the measured traits, including herbivory, we found limited evidence for local adaptation. The effects of admixture depended on experimental site and plant population, and were positive for some traits. Plant growth and fruit production of some populations increased in admixed offspring and this was strongest with larger parental distances. These effects were only detected in two of our three sites. Our results show that, in the absence of local adaptation, admixture may boost plant performance, and that this is particularly apparent in stressful environments. We suggest that admixture between foreign and local genotypes can potentially be considered in nature conservation to restore populations and/or increase population viability, especially in small inbred or maladapted populations.

Global warming and reduced precipitation may trigger large-scale species losses and vegetation shifts in ecosystems around the world. However, currently lacking are practical ways to quantify the sensitivity of species and community composition to these often-confounded climatic forces. Here we conducted long-term (16 yr) nocturnal-warming (+0.6°C) and reduced precipitation (-20% soil moisture) experiments in a Mediterranean shrubland. Climatic niche groups (CNGs) - species ranked or classified by similar temperature or precipitation distributions - informatively described community responses under experimental manipulations. Under warming, CNGs revealed that only those species distributed in cooler regions decreased. Correspondingly, under reduced precipitation, a U-shaped treatment effect observed in the total community was the result of an abrupt decrease in wet-distributed species, followed by a delayed increase in dry-distributed species. Notably, while partially correlated, CNG explanations of community response were stronger for their respective climate parameter, suggesting some species possess specific adaptations to either warming or drought that may lead to independent selection to the two climatic variables. Our findings indicate that when climatic distributions are combined with experiments, the resulting incorporation of local plant evolutionary strategies and their changing dynamics over time leads to predictable and informative shifts in community structure under independent climate change scenarios., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Intraspecific hybridization between diverged populations can enhance fitness via various genetic mechanisms. The benefits of such admixture have been proposed to be particularly relevant in biological invasions, when invasive populations originating from different source populations are found sympatrically. However, it remains poorly understood if admixture is an important contributor to plant invasive success and how admixture effects compare between invasive and native ranges. Here, we used experimental crosses in Lythrum salicaria , a species with well-established history of multiple introductions to Eastern North America, to quantify and compare admixture effects in native European and invasive North American populations. We observed heterosis in between-population crosses both in native and invasive ranges. However, invasive-range heterosis was restricted to crosses between two different Eastern and Western invasion fronts, whereas heterosis was absent in geographically distant crosses within a single large invasion front. Our results suggest that multiple introductions have led to already-admixed invasion fronts, such that experimental crosses do not further increase performance, but that contact between different invasion fronts further enhances fitness after admixture. Thus, intra-continental movement of invasive plants in their introduced range has the potential to boost invasiveness even in well-established and successfully spreading invasive species., Competing Interests: Compliance with ethical standardsThe authors declare that they have no conflict of interest.

Plants can plastically respond to light competition in three strategies, comprising vertical growth, which promotes competitive dominance; shade tolerance, which maximises performance under shade; or lateral growth, which offers avoidance of competition. Here, we test the hypothesis that plants can 'choose' between these responses, according to their abilities to competitively overcome their neighbours. We study this hypothesis in the clonal plant Potentilla reptans using an experimental setup that simulates both the height and density of neighbours, thus presenting plants with different light-competition scenarios. Potentilla reptans ramets exhibit the highest vertical growth under simulated short-dense neighbours, highest specific leaf area (leaf area/dry mass) under tall-dense neighbours, and tend to increase total stolon length under tall-sparse neighbours. These responses suggest shifts between 'confrontational' vertical growth, shade tolerance and lateral-avoidance, respectively, and provide evidence that plants adopt one of several alternative plastic responses in a way that optimally corresponds to prevailing light-competition scenarios.

To understand the biological effects of climate change, it is essential to take into account species' evolutionary responses to their changing environments. Ongoing climate change is resulting in species shifting their geographical distribution ranges poleward. We tested whether a successful range expanding plant has rapidly adapted to the regional conditions in its novel range, and whether adaptation could be driven by herbivores. Furthermore, we investigated if enemy release occurred in the newly colonized areas and whether plant origins differed in herbivore resistance. Plants were cloned and reciprocally transplanted between three experimental sites across the range. Effects of herbivores on plant performance were tested by individually caging plants with either open or closed cages. There was no indication of (regional) adaptation to abiotic conditions. Plants originating from the novel range were always larger than plants from the core distribution at all experimental sites, with or without herbivory. Herbivore damage was highest and not lowest at the experimental sites in the novel range, suggesting no release from enemy impact. Genotypes from the core were more damaged compared to genotypes from newly colonized areas at the most northern site in the novel range, which was dominated by generalist slug herbivory. We also detected subtle shifts in chemical defenses between the plant origins. Genotypes from the novel range had more inducible defenses. Our results suggest that plants that are expanding their range with climate change may evolve increased vigor and altered herbivore resistance in their new range, analogous to invasive plants.

Bet-hedging via between-year seed dormancy is a costly strategy for plants in unpredictable environments. Theoretically, fitness costs can be reduced through a parental environmental effect when the environment is partly predictable. We tested whether populations from environments that differ in predictability diverged in parental effects on seed dormancy. Common garden-produced seeds of the two annual plant species Biscutella didyma and Bromus fasciculatus collected along an aridity gradient were grown under 12 irrigation treatments. Offspring germination was evaluated and related to environmental correlations between generations and their fitness consequences at the four study sites. One species exhibited strong seed dormancy that increased with unpredictability in seasonal precipitation. The parental effect on seed dormancy also increased proportionally with the environmental correlation between precipitation in the parental season and seedling density in the following season; this correlation increased from mesic to arid environments. Because fitness was negatively related to density, this parental effect may be adaptive. However, the lack of dormancy in the second species indicates that bet-hedging is not the only strategy for annual plants in arid environments. Our results provide the first evidence for clinal variation in the relative strength of parental effects along environmental gradients., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Well-defined productivity-precipitation relationships of ecosystems are needed as benchmarks for the validation of land models used for future projections. The productivity-precipitation relationship may be studied in two ways: the spatial approach relates differences in productivity to those in precipitation among sites along a precipitation gradient (the spatial fit, with a steeper slope); the temporal approach relates interannual productivity changes to variation in precipitation within sites (the temporal fits, with flatter slopes). Precipitation-reduction experiments in natural ecosystems represent a complement to the fits, because they can reduce precipitation below the natural range and are thus well suited to study potential effects of climate drying. Here, we analyse the effects of dry treatments in eleven multiyear precipitation-manipulation experiments, focusing on changes in the temporal fit. We expected that structural changes in the dry treatments would occur in some experiments, thereby reducing the intercept of the temporal fit and displacing the productivity-precipitation relationship downward the spatial fit. The majority of experiments (72%) showed that dry treatments did not alter the temporal fit. This implies that current temporal fits are to be preferred over the spatial fit to benchmark land-model projections of productivity under future climate within the precipitation ranges covered by the experiments. Moreover, in two experiments, the intercept of the temporal fit unexpectedly increased due to mechanisms that reduced either water loss or nutrient loss. The expected decrease of the intercept was observed in only one experiment, and only when distinguishing between the late and the early phases of the experiment. This implies that we currently do not know at which precipitation-reduction level or at which experimental duration structural changes will start to alter ecosystem productivity. Our study highlights the need for experiments with multiple, including more extreme, dry treatments, to identify the precipitation boundaries within which the current temporal fits remain valid., (© 2016 John Wiley & Sons Ltd.)

The number of individuals within a population or community and their body size can be associated with changes in resource supply. While these relationships may provide a key to better understand the role of abiotic vs. biotic constraints in animal communities, little is known about the way size and abundance of organisms change along resource gradients. Here, we studied this interplay in ants, addressing two hypotheses with opposite predictions regarding variation in population densities along resource gradients- the 'productivity hypothesis' and the 'productivity-based thinning hypothesis'. The hypotheses were tested in two functional groups of ground-dwelling ants that are directly primary consumers feeding on seeds: specialized seed-eaters and generalist species. We examined variations in colony density and foraging activity (a size measurement of the forager caste) in six ant assemblages along a steep productivity gradient in a semi-arid region, where precipitation and plant biomass vary 6-fold over a distance of 250km. An increase in the density or foraging activity of ant colonies along productivity gradients is also likely to affect competitive interactions among colonies, and consequently clinal changes in competition intensity were also examined. Ant foraging activity increased with productivity for both functional groups. However, colony density revealed opposing patterns: it increased with productivity for the specialized seed-eaters, but decreased for the generalist species. Competition intensity, evaluated by spatial partitioning of species at food baits and distribution of colonies, was uncorrelated with productivity in the specialized seed-eaters, but decreased with increasing productivity in the generalists. Our results provide support for two contrasting hypotheses regarding the effect of resource availability on the abundance of colonial organisms- the 'productivity hypothesis' for specialized seed-eaters and the 'productivity-based thinning hypothesis' for generalist species. These results also stress the importance of considering the role of functional groups in studies of community structure.

Altered rainfall regimes will greatly affect the response of plant species to climate change. However, little is known about how direct effects of changing precipitation on plant performance may depend on other abiotic factors and biotic interactions. We used reciprocal transplants between climatically very different sites with simultaneous manipulation of soil, plant population origin, and neighbor conditions to evaluate local adaptation and possible adaptive response of four Eastern Mediterranean annual plant species to climate change. The effect of site on plant performance was negligible, but soil origin had a strong effect on fecundity, most likely due to differential water retaining ability. Competition by neighbors strongly reduced fitness. We separated the effects of the abiotic and biotic soil properties on plant performance by repeating the field experiment in a greenhouse under homogenous environmental conditions and including a soil biota manipulation treatment. As in the field, plant performance differed among soil origins and neighbor treatments. Moreover, we found plant species-specific responses to soil biota that may be best explained by the differential sensitivity to negative and positive soil biota effects. Overall, under the conditions of our experiment with two contrasting sites, biotic interactions had a strong effect on plant fitness that interacted with and eventually overrode climate. Because climate and biotic interactions covary, reciprocal transplants and climate gradient studies should consider soil biotic interactions and abiotic conditions when evaluating climate change effects on plant performance.

Many species will need to adapt to the observed climate change in order to persist. However, research about adaptation or phenotypic plasticity in response to climate change is rare. In particular, field studies are lacking that impose artificial selection for a sufficiently long time to elicit changes in phenotypic and genotypic structure of populations. Here, we present findings for an 8-year field experiment with 16 annual plant species that tested potentially adaptive phenotypic responses to precipitation change. In both a Mediterranean and a semi-arid site, annual precipitation was manipulated (±30%) and phenotypic response was recorded. We measured flowering time as a key trait related to climatic conditions and biomass and survival as fitness correlates. Differences in traits among treatments were compared to trait shifts between sites, according to space-for-time approaches. In the drier site, phenology was accelerated, but within that site, experimental drought delayed phenology, probably as a plastic response to delayed ontogenetic development. Biomass was smaller in the dry treatments of that site, but it was also reduced in irrigated plots in both sites, indicating more intense competition. The shifts from limitation by drought to limitation by competition corresponded to patterns along the gradient. This also implies a larger negative impact of climate change in the drier site. Our results suggest that experimental selection in the field caused directional responses in most species, but these were not necessarily adaptive. Furthermore, competitive release imposed by climate change may revert direct negative effects of rainfall change in determining plant performance.

Background and Aims: A common hypothesis to explain plants' invasive success is that release from natural enemies in the introduced range selects for reduced allocation to resistance traits and a subsequent increase in resources available for growth and competitive ability (evolution of increased competitive ability, EICA). However, studies that have investigated this hypothesis have been incomplete as they either did not test for all aspects of competitive ability or did not select appropriate competitors., Methods: Here, the prediction of increased competitive ability was examined with the invasive plant Lythrum salicaria (purple loosestrife) in a set of common-garden experiments that addressed these aspects by carefully distinguishing between competitive effect and response of invasive and native plants, and by using both intraspecific and interspecific competition settings with a highly vigorous neighbour, Urtica dioica (stinging nettle), which occurs in both ranges., Key Results: While the intraspecific competition results showed no differences in competitive effect or response between native and invasive plants, the interspecific competition experiment revealed greater competitive response and effect of invasive plants in both biomass and seed production., Conclusions: The use of both intra- and interspecific competition experiments in this study revealed opposing results. While the first experiment refutes the EICA hypothesis, the second shows strong support for it, suggesting evolutionarily increased competitive ability in invasive populations of L. salicaria. It is suggested that the use of naturally co-occurring heterospecifics, rather than conspecifics, may provide a better evaluation of the possible evolutionary shift towards greater competitive ability., (© The Author 2014. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

For evaluating climate change impacts on biodiversity, extensive experiments are urgently needed to complement popular non-mechanistic models which map future ecosystem properties onto their current climatic niche. Here, we experimentally test the main prediction of these models by means of a novel multi-site approach. We implement rainfall manipulations--irrigation and drought--to dryland plant communities situated along a steep climatic gradient in a global biodiversity hotspot containing many wild progenitors of crops. Despite the large extent of our study, spanning nine plant generations and many species, very few differences between treatments were observed in the vegetation response variables: biomass, species composition, species richness and density. The lack of a clear drought effect challenges studies classifying dryland ecosystems as most vulnerable to global change. We attribute this resistance to the tremendous temporal and spatial heterogeneity under which the plants have evolved, concluding that this should be accounted for when predicting future biodiversity change.

Due to global warming, species are expanding their range to higher latitudes. Some range expanding plants have become invasive in their new range. The Evolution of Increased Competitive Ability (EICA) hypothesis and the Shifting Defense Hypothesis (SDH) predict altered selection on plant defenses in the introduced range of invasive plants due to changes in herbivore pressures and communities. Here, we investigated chemical defenses (glucosinolates) of five native and seven invasive populations of the Eurasian invasive range expanding plant, Rorippa austriaca. Further, we studied feeding preferences of a generalist and a specialist herbivore among the populations. We detected eight glucosinolates in the leaves of R. austriaca. 8-Methylsulfinyloctyl glucosinolate was the most abundant glucosinolate in all plants. There were no overall differences between native and invasive plants in concentrations of glucosinolates. However, concentrations among populations within each range differed significantly. Feeding preference between the populations by a generalist herbivore was negatively correlated with glucosinolate concentrations. Feeding by a specialist did not differ between the populations and was not correlated with glucosinolates. Possibly, local differences in herbivore communities within each range may explain the differences in concentrations of glucosinolates among populations. Little support for the predictions of the EICA hypothesis or the SDH was found for the glucosinolate defenses of the studied native and invasive R. austriaca populations.

Disturbances' role in shaping communities is well documented but highly disputed. We suggest replacing the overused two-trait trade-off approach with a functional group scheme, constructed from combinations of four key traits that represent four classes of species' responses to disturbances. Using model results and field observations from sites affected by two highly different disturbances, we demonstrated that popular dichotomous trade-offs are not sufficient to explain community dynamics, even if some emerge under certain conditions. Without disturbances, competition was only sufficient to predict species survival but not relative success, which required some escape mechanism (e.g., long-term dormancy). With highly predictable and large-scale disturbances, successful species showed a combination of high individual tolerance to disturbance and, more surprisingly, high competitive ability. When disturbances were less predictable, high individual tolerance and long-term seed dormancy were favored, due to higher environmental uncertainty. Our study demonstrates that theories relying on a small number of predefined trade-offs among traits (e.g., competition-colonization trade-off) may lead to unrealistic results. We suggest that the understanding of disturbance-community relationships can be significantly improved by employing sets of relevant trait assemblies instead of the currently common approach in which trade-offs are assumed in advance.

Desert species respond strongly to infrequent, intense pulses of precipitation. Consequently, indigenous flora has developed a rich repertoire of life-history strategies to deal with fluctuations in resource availability. Examinations of how future climate change will affect the biota often forecast negative impacts, but these-usually correlative-approaches overlook precipitation variation because they are based on averages. Here, we provide an overview of how variable precipitation affects perennial and annual desert plants, and then implement an innovative, mechanistic approach to examine the effects of precipitation on populations of two desert plant species. This approach couples robust climatic projections, including variable precipitation, with stochastic, stage-structured models constructed from long-term demographic datasets of the short-lived Cryptantha flava in the Colorado Plateau Desert (USA) and the annual Carrichtera annua in the Negev Desert (Israel). Our results highlight these populations' potential to buffer future stochastic precipitation. Population growth rates in both species increased under future conditions: wetter, longer growing seasons for Cryptantha and drier years for Carrichtera. We determined that such changes are primarily due to survival and size changes for Cryptantha and the role of seed bank for Carrichtera. Our work suggests that desert plants, and thus the resources they provide, might be more resilient to climate change than previously thought.

Background and Aims: The enemy release hypothesis assumes that invasive plants lose their co-evolved natural enemies during introduction into the new range. This study tested, as proposed by the evolution of increased competitive ability (EICA) hypothesis, whether escape from enemies results in a decrease in defence ability in plants from the invaded range. Two straightforward aspects of the EICA are examined: (1) if invasives have lost their enemies and their defence, they should be more negatively affected by their full natural pre-invasion herbivore spectrum than their native conspecifics; and (2) the genetic basis of evolutionary change in response to enemy release in the invasive range has not been taken sufficiently into account., Methods: Lythrum salicaria (purple loosestrife) from several populations in its native (Europe) and invasive range (North America) was exposed to all above-ground herbivores in replicated natural populations in the native range. The experiment was performed both with plants raised from field-collected seeds as well as with offspring of these where maternal effects were removed., Key Results: Absolute and relative leaf damage was higher for introduced than for native plants. Despite having smaller height growth rate, invasive plants attained a much larger final size than natives irrespective of damage, indicating large tolerance rather than effective defence. Origin effects on response to herbivory and growth were stronger in second-generation plants, suggesting that invasive potential through enemy release has a genetic basis., Conclusions: The findings support two predictions of the EICA hypothesis - a genetically determined difference between native and invasive plants in plant vigour and response to enemies - and point to the importance of experiments that control for maternal effects and include the entire spectrum of native range enemies.

Morphological plasticity is a striking characteristic of plants in natural communities. In the context of foraging behavior particularly, root plasticity has been documented for numerous species. Root plasticity is known to mitigate competitive interactions by reducing the overlap of the individuals' rhizospheres. But despite its obvious effect on resource acquisition, plasticity has been generally neglected in previous empirical and theoretical studies estimating interaction intensity among plants. In this study, we developed a semi-mechanistic model that addresses this shortcoming by introducing the idea of compensatory growth into the classical-zone-of influence (ZOI) and field-of-neighborhood (FON) approaches. The model parameters describing the belowground plastic sphere of influence (PSI) were parameterized using data from an accompanying field experiment. Measurements of the uptake of a stable nutrient analogue at distinct distances to the neighboring plants showed that the study species responded plastically to belowground competition by avoiding overlap of individuals' rhizospheres. An unexpected finding was that the sphere of influence of the study species Bromus hordeaceus could be best described by a unimodal function of distance to the plant's center and not with a continuously decreasing function as commonly assumed. We employed the parameterized model to investigate the interplay between plasticity and two other important factors determining the intensity of competitive interactions: overall plant density and the distribution of individuals in space. The simulation results confirm that the reduction of competition intensity due to morphological plasticity strongly depends on the spatial structure of the competitive environment. We advocate the use of semi-mechanistic simulations that explicitly consider morphological plasticity to improve our mechanistic understanding of plant interactions.

The eastern Mediterranean faces a severe water crisis: water supply decreases due to climate change, while demand increases due to rapid population growth. The GLOWA Jordan River project generates science-based management strategies for maximizing water productivity under global climate change. We use a novel definition of water productivity as the full range of services provided by landscapes per unit blue (surface) and green (in plants and soil) water. Our combined results from climatological, ecological, economic and hydrological studies suggest that, in Israel, certain landscapes provide high returns as ecosystem services for little input of additional blue water. Specifically, cultural services such as recreation may by far exceed that of food production. Interestingly, some highly valued landscapes (e.g. rangeland) appear resistant to climate change, making them an ideal candidate for adaptive land management. Vice versa, expanding irrigated agriculture is unlikely to be sustainable under global climate change. We advocate the inclusion of a large range of ecosystem services into integrated land and water resources management. The focus on cultural services and integration of irrigation demand will lead to entirely different but productive water and land allocation schemes that may be suitable for withstanding the problems caused by climate change.

Bet hedging is a means to increase fitness in environments that vary unpredictably in space and time. In such environments, models predict a trade-off between the bet-hedging strategies dispersal and dormancy, while the increasing importance of risk reduction with decreasing predictability should lead to an increase in dispersal and dormancy along gradients of environmental predictability. However, so far there has been no experimental study to test these predictions in the field. Here, we used a set of novel field experiments that enabled us to quantify and separate seedling recruitment from three sources: local reproduction, dormancy, and dispersal. The study included the entire plant community from five environments differing considerably in predictability. Evidence for both the existence of a trade-off between dispersal and dormancy within environments and their increased use in unpredictable environments was very weak. The importance of dispersal for population and community dynamics in our system was extremely low relative to dormancy and local reproduction. This indicates that the role of dispersal for buffering environmental variation may be negligible compared with other risk-reducing strategies. Our findings highlight the urgent need for multispecies and multisite experiments in empirical tests of theoretical predictions.

The role of local adaptation and factors other than climate in determining extinction probabilities of species under climate change has not been yet explicitly studied. Here we performed a field experiment with annual plants growing along a steep climatic gradient in Israel to isolate climatic effects for local trait expression. The focus trait was seed dormancy, for which many theoretical predictions exist regarding climate-driven optimal germination behaviour. We evaluated how germination is consistent with theory, indicating local adaptation to current and changing climatic conditions, and how it varies among species and between natural and standardised soil conditions. We reciprocally sowed seeds from three or four origins for each of three annual species, Biscutella didyma, Bromus fasciculatus and Hymenocarpos circinnatus, in their home and neighbouring sowing locations along an aridity gradient. Our predictions were: lower germination fraction for seeds from more arid origins, and higher germination at wetter sowing locations for all seed origins. By sowing seeds in both local and standard soil, we separated climatic effects from local conditions. At the arid sowing location, two species supported the prediction of low germination of drier seed origins, but differences between seed origins at the other sites were not substantial. There were no clear rainfall effects on germination. Germination fractions were consistently lower on local soil than on standard soil, indicating the important role of soil type and neighbour conditions for trait expression. Local environmental conditions may override effects of climate and so should be carefully addressed in future studies testing for the potential of species to adapt or plastically respond to climate change.

Dormant seeds in permanent soil seed banks change their germination probability with age, but little is known about the causes of this behavior. Since current adaptive models do not consider the age structure of the seed bank, the sole explanation is that changes in germination probability with age are due to passive accidental factors or to decay of seeds. Here we present a theory, which shows that such changes might be adaptive. In particular, in density-regulated populations, the evolutionarily stable fraction of germinating seeds in each age class increases with age whenever age-dependent mortality of dormant seeds remains constant or increases with age.