Showing papers in "Journal of Ecology in 2013"

Plant–soil feedbacks: the past, the present and future challenges

Abstract: Summary Plant–soil feedbacks is becoming an important concept for explaining vegetation dynamics, the invasiveness of introduced exotic species in new habitats and how terrestrial ecosystems respond to global land use and climate change. Using a new conceptual model, we show how critical alterations in plant–soil feedback interactions can change the assemblage of plant communities. We highlight recent advances, define terms and identify future challenges in this area of research and discuss how variations in strengths and directions of plant–soil feedbacks can explain succession, invasion, response to climate warming and diversity-productivity relationships. While there has been a rapid increase in understanding the biological, chemical and physical mechanisms and their interdependencies underlying plant–soil feedback interactions, further progress is to be expected from applying new experimental techniques and technologies, linking empirical studies to modelling and field-based studies that can include plant–soil feedback interactions on longer time scales that also include long-term processes such as litter decomposition and mineralization. Significant progress has also been made in analysing consequences of plant–soil feedbacks for biodiversity-functioning relationships, plant fitness and selection. To further integrate plant–soil feedbacks into ecological theory, it will be important to determine where and how observed patterns may be generalized, and how they may influence evolution. Synthesis. Gaining a greater understanding of plant–soil feedbacks and underlying mechanisms is improving our ability to predict consequences of these interactions for plant community composition and productivity under a variety of conditions. Future research will enable better prediction and mitigation of the consequences of human-induced global changes, improve efforts of restoration and conservation and promote sustainable provision of ecosystem services in a rapidly changing world.

Identification of 100 fundamental ecological questions

Abstract: Summary 1. Fundamental ecological research is both intrinsically interesting and provides the basic knowledge required to answer applied questions of importance to the management of the natural world. The 100th anniversary of the British Ecological Society in 2013 is an opportune moment to reflect on the current status of ecology as a science and look forward to high-light priorities for future work.

Linking litter decomposition of above‐ and below‐ground organs to plant–soil feedbacks worldwide

Abstract: Conceptual frameworks relating plant traits to ecosystem processes such as organic matter dynamics are progressively moving from a leaf-centred to a whole-plant perspective. Through the use of meta-analysis and global literature data, we quantified the relative roles of litters from above- and below-ground plant organs in ecosystem labile organic matter dynamics. We found that decomposition rates of leaves, fine roots and fine stems were coordinated across species worldwide although less strongly within ecosystems. We also show that fine roots and stems had lower decomposition rates relative to leaves, with large differences between woody and herbaceous species. Further, we estimated that on average below-ground litter represents approximately 33 and 48% of annual litter inputs in grasslands and forests, respectively. These results suggest a major role for below-ground litter as a driver of ecosystem organic matter dynamics. We also suggest that, given that fine stem and fine root litters decompose approximately 1.5 and 2.8 times slower, respectively, than leaf litter derived from the same species, cycling of labile organic matter is likely to be much slower than predicted by data from leaf litter decomposition only. Synthesis. Our results provide evidence that within ecosystems, the relative inputs of above- versus below-ground litter strongly control the overall quality of the litter entering the decomposition system. This in turn determines soil labile organic matter dynamics and associated nutrient release in the ecosystem, which potentially feeds back to the mineral nutrition of plants and therefore plant trait values and plant community composition. © 2013 The Authors. Journal of Ecology © 2013 British Ecological Society.

Latitudinal gradients as natural laboratories to infer species' responses to temperature

Abstract: Macroclimatic variation along latitudinal gradients provides an excellent natural laboratory to investigate the role of temperature and the potential impacts of climate warming on terrestrial organisms. Here, we review the use of latitudinal gradients for ecological climate change research, in comparison with altitudinal gradients and experimental warming, and illustrate their use and caveats with a meta-analysis of latitudinal intraspecific variation in important life-history traits of vascular plants. We first provide an overview of latitudinal patterns in temperature and other abiotic and biotic environmental variables in terrestrial ecosystems. We then assess the latitudinal intraspecific variation present in five key life-history traits [plant height, specific leaf area (SLA), foliar nitrogen:phosphorus (N:P) stoichiometry, seed mass and root:shoot (R:S) ratio] in natural populations or common garden experiments across a total of 98 plant species. Intraspecific leaf N:P ratio and seed mass significantly decreased with latitude in natural populations. Conversely, the plant height decreased and SLA increased significantly with latitude of population origin in common garden experiments. However, less than a third of the investigated latitudinal transect studies also formally disentangled the effects of temperature from other environmental drivers which potentially hampers the translation from latitudinal effects into a temperature signal. Synthesis. Latitudinal gradients provide a methodological set-up to overcome the drawbacks of other observational and experimental warming methods. Our synthesis indicates that many life-history traits of plants vary with latitude but the translation of latitudinal clines into responses to temperature is a crucial step. Therefore, especially adaptive differentiation of populations and confounding environmental factors other than temperature need to be considered. More generally, integrated approaches of observational studies along temperature gradients, experimental methods and common garden experiments increasingly emerge as the way forward to further our understanding of species and community responses to climate warming.

Plant functional diversity and carbon storage - an empirical test in semi-arid forest ecosystems

Abstract: Summary 1. Carbon storage in vegetation and soil underpins climate regulation through carbon sequestration. Because plant species differ in their ability to capture, store and release carbon, the collective functional characteristics of plant communities (functional diversity) should be a major driver of carbon accumulation in terrestrial ecosystems. 2. Three major components of plant functional diversity could be put forward as drivers of carbon storage in ecosystems: the most abundant functional trait values, the variety of functional trait values and the abundance of particular species that could have additional effects not incorporated in the first two components. 3. We tested for associations between these components and carbon storage across 16 sites in the Chaco forest of Argentina under the same climate and on highly similar parental material. The sites differed in their plant functional diversity caused by different long-term land-use regimes. 4. We measured six plant functional traits in 27 species and weighted them by the species abundance at each site to calculate the community-weighted mean (CWM) and the functional divergence (FDvar) of each single trait and of multiple traits (FDiv). We also measured plant and soil carbon storage. Using a stepwise multiple regression analysis, we assessed which of the functional diversity components best explained carbon storage. 5. Both CWM and FDvar of plant height and wood-specific gravity, but no leaf traits, were retained as predictors of carbon storage in multiple models. Relationships of FDvar of stem traits and FDiv with carbon storage were all negative. The abundance of five species improved the predictive power of some of the carbon storage models. 6. Synthesis. All three major components of plant functional diversity contributed to explain carbon storage. What matters the most to carbon storage in these ecosystems is the relative abundance of plants with tall, and to a lesser extent dense, stems with a narrow range of variation around these values. No consistent link was found between carbon storage and the leaf traits usually associated with plant resource use strategy. The negative association of trait divergence with carbon storage provided no evidence in support to niche complementarity promoting carbon storage in these forest ecosystems.

Contrasting changes in taxonomic, phylogenetic and functional diversity during a long‐term succession: insights into assembly processes

Abstract: Summary 1. Theory predicts that the processes generating biodiversity after disturbance will change during succession. Comparisons of phylogenetic and functional (alpha and beta) diversity with taxonomic diversity can provide insights into the extent to which community assembly is driven by deterministic or stochastic processes, but comparative approaches have yet to be applied to successional systems. 2. We characterized taxonomic, phylogenetic and functional plant (alpha and beta) diversity within and between four successional stages in a > 270-year-long arable-to-grassland chronosequence. Null models were used to test whether functional and phylogenetic turnover differed from random expectations, given the levels of species diversity. 3. The three facets of diversity showed different patterns of change during succession. Between early and early-mid succession, species richness increased but there was no increase in functional or phylogenetic diversity. Higher than predicted levels of functional similarity between species within the early and early-mid successional stages, indicate that abiotic filters have selected for sets of functionally similar species within sites. Between late-mid and late succession, there was no further increase in species richness, but a significant increase in functional alpha diversity, suggesting that functionally redundant species were replaced by functionally more dissimilar species. Functional turnover between stages was higher than predicted, and higher than within-stage turnover, indicating that different assembly processes act at different successional stages. 4. Synthesis. Analysis of spatial and temporal turnover in different facets of diversity suggests that deterministic processes generate biodiversity during post-disturbance ecosystem development and that the relative importance of assembly processes has changed over time. Trait-mediated abiotic filtering appears to play an important role in community assembly during the early and early-mid stages of arable-to-grassland succession, whereas the relative importance of competitive exclusion appears to have increased towards the later successional stages. Phylogenetic diversity provided a poor reflection of functional diversity and did not contribute to inferences about underlying assembly processes. Functionally deterministic assembly suggests that it may be possible to predict future post-disturbance changes in biodiversity, and associated ecosystem attributes, on the basis of species’ functional traits but not phylogeny.

Relative contributions of plant traits and soil microbial properties to mountain grassland ecosystem services

Abstract: Summary 1. Plant functional diversity and soil microbial community composition are tightly coupled. Changes in these interactions may influence ecosystem functioning. Links between plant functional diversity, soil microbial communities and ecosystem functioning have been demonstrated in experiments using plant monocultures and mixtures, using broad plant and microbial functional groups, but have not been examined in diverse natural plant communities. 2. We quantified the relative effects of plant and microbial functional properties on key ecosystem functions. We measured plant functional diversity, soil microbial community composition and parameters associated with nitrogen (N) cycling and key nutrient cycling processes at three grassland sites in different parts of Europe. Because plant structure and function strongly influence soil microbial communities, we determined relationships between ecosystem properties, plant traits and soil community characteristics following a sequential approach in which plant traits were fitted first, followed by the additional effects of soil micro-organisms. 3. We identified a continuum from standing green biomass and standing litter, linked mostly with plant traits, to potential N mineralization and potential leaching of soil inorganic N, linked mostly with microbial properties. Plant and microbial functional parameters were equally important in explaining % organic matter content in soil. A parallel continuum ran from plant height, linked with above-ground biomass, to plant quality effects captured by the leaf economics spectrum, which were linked with the recycling of carbon (C) and N. 4. More exploitative species (higher specific leaf area, leaf N concentrations and lower leaf dry matter content) and taller swards, along with soil microbial communities dominated by bacteria, with rapid microbial activities, were linked with greater fodder production, but poor C and N retention. Conversely, dominance by conservative species (with opposite traits) and soil microbial communities dominated by fungi, and bacteria with slow activities, were usually linked with low production, but greater soil C storage and N retention. 5. Synthesis – Grassland production, C sequestration and soil N retention are jointly related to plant and microbial functional traits. Managing grasslands for selected, or multiple, ecosystem services will thus require a consideration of the joint effects of plant and soil communities. Further understanding of the mechanisms that link plant and microbial functional traits is essential to achieve this.

Scale-dependent relationships between tree species richness and ecosystem function in forests

Abstract: 1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity.

Plant functional effects on ecosystem services

Abstract: Summary The prominent new place of ecosystem services in environmental policy, land management and land planning requires that the best ecological knowledge be applied to ecosystem service quantification. Given strong evidence that functional diversity underpins the delivery of key ecosystem services, assessments of these services may progress rapidly using a trait-based approach. The trait-based approach shows promising results, especially for plant trait effects on primary production and some processes associated with carbon and nitrogen cycling in grasslands. However, there is a need to extend the proof of concept for a wider range of ecosystems and ecosystem services and to incorporate not only the functional characteristics of plants but those of other organisms with which plants interact for the provision of ecosystem services. The five papers in this Special Feature illustrate how some of the key conceptual and methodological challenges can be resolved, and provide a range of case studies across three continents. Relevant plant functional traits depict different axes of variation including stature, the leaf economics spectrum, and associated or independent variations in root or stem traits. The application of the trait approach to ecosystem processes underpinned by interactions between plants and other biota is illustrated for soil micro-organisms and granivorous invertebrates. There is strong evidence for the biomass ratio hypothesis (i.e. prevalent effects of the traits of dominant species through the community-weighted mean), along with less prevalent and more complex effects of heterogeneous trait values between species (i.e. functional divergence). Synthesis. Together, the five papers in this Special Feature illustrate how trait-based approaches may help elucidate the complexity of ecological mechanisms operating in the field to determine ecosystem service delivery. To address scientific and management questions about the provision of multiple services, progress is needed in understanding how functional trade-offs and synergies within organisms scale up to interactions between ecosystem services. Service-oriented ecosystem management within the context of global change, or ecological restoration, remains a major challenge, but trait-based understanding opens new avenues towards more generic, integrated approaches.

Plant apparency, an overlooked driver of associational resistance to insect herbivory

Abstract: Summary Herbivore regulation is one of the services provided by plant diversity in terrestrial ecosystems. It has been suggested that tree diversity decreases insect herbivory in forests, but recent studies have reported opposite patterns, indicating that tree diversity can trigger associational resistance or susceptibility. The mechanisms underlying the tree diversity–resistance relationship thus remain a matter of debate. We assessed insect herbivory on pedunculate oak saplings (Quercus robur) in a large-scale experiment in which we manipulated tree diversity and identity by mixing oaks, birch and pine species. Tree diversity at the plot scale had no effect on damage due to leaf chewers, but abundance of leaf miners decreased with increasing tree diversity. The magnitude of this associational resistance increased with host dilution, consistent with the ‘resource concentration hypothesis’. At a smaller scale, we estimated tree apparency as the difference in total height between focal oak saplings and their nearest neighbouring trees. Levels of oak infestation with leaf miners decreased significantly with decreasing tree apparency. As the probability of having taller neighbours increased with tree diversity, notably due to the increase in the proportion of faster growing nonhost trees, such as birches and pines, tree apparency may be seen as a ‘hidden’, sampling effect of tree diversity. Synthesis. These findings suggest that greater host dilution and lower tree apparency contribute to associational resistance in young trees. They also highlight the importance of taking plant size into account as a covariate, to avoid misleading interpretations about the biodiversity–resistance relationship.

Biological Flora of the British Isles: Robinia pseudoacacia

Abstract: Summary This account presents information on all aspects of the biology of Robinia pseudoacacia L. that are relevant to understanding its ecological characteristics and behaviour. The main topics are presented within the standard framework of the Biological Flora of the British Isles: distribution, habitat, communities, responses to biotic factors, responses to environment, structure and physiology, phenology, floral and seed characters, herbivores and disease, and history and conservation. Robinia pseudoacacia, false acacia or black locust, is a deciduous, broad-leaved tree native to North America. The medium-sized, fast-growing tree is armed with spines, and extensively suckering. It has become naturalized in grassland, semi-natural woodlands and urban habitats. The tree is common in the south of the British Isles and in many other regions of Europe. Robinia pseudoacacia is a light-demanding pioneer species, which occurs primarily in disturbed sites on fertile to poor soils. The tree does not tolerate wet or compacted soils. In contrast to its native range, where it rapidly colonizes forest gaps and is replaced after 15–30 years by more competitive tree species, populations in the secondary range can persist for a longer time, probably due to release from natural enemies. Robinia pseudoacacia reproduces sexually, and asexually by underground runners. Disturbance favours clonal growth and leads to an increase in the number of ramets. Mechanical stem damage and fires also lead to increased clonal recruitment. The tree benefits from di-nitrogen fixation associated with symbiotic rhizobia in root nodules. Estimated symbiotic nitrogen fixation rates range widely from 23 to 300 kg ha−1 year−1. The nitrogen becomes available to other plants mainly by the rapid decay of nitrogen-rich leaves. Robinia pseudoacacia is host to a wide range of fungi both in the native and introduced ranges. Megaherbivores are of minor significance in Europe but browsing by ungulates occurs in the native range. Among insects, the North American black locust gall midge (Obolodiplosis robiniae) is specific to Robinia and is spreading rapidly throughout Europe. In parts of Europe, Robinia pseudoacacia is considered an invasive non-indigenous plant and the tree is controlled. Negative impacts include shading and changes of soil conditions as a result of nitrogen fixation.

Specialist species of wood-inhabiting fungi struggle while generalists thrive in fragmented boreal forests

Abstract: Summary The loss of suitable habitats is one of the main causes behind the loss of species and communities. Habitat fragmentation, that is, the division of the remaining habitat into small and isolated fragments, often co-occurs with the process of habitat loss. The spatial division of habitats decreases connectivity among local populations and generally has a negative effect on population viability, but it can also have a positive effect for some species, for example, due to released competition pressure. In both animals and plants, certain characteristics such as low dispersal ability and narrow ecological niche are known to be associated with fragmentation vulnerability, but in fungi, systematic analyses have so far been lacking. With their small and highly dispersive spores, fungi could be mainly resource-limited, not dispersal-limited. In this study, we analysed spatial occurrence data on 119 species of wood-inhabiting fungi to identify the species characteristics that are associated with high extinction risk and fragmentation vulnerability in particular. We modelled resource use and connectivity dependence separately for each species using the presence–absence data on 98 318 dead trees in 496 sites located on a gradient in the duration and intensity of land use in eastern Fennoscandia. We then related species' responses to connectivity to their resource-use patterns, life-history characteristics and red-list status. Our results show that red-listed species are highly specialized in their resource use and suffer from loss of connectivity at three spatial scales: along the large-scale gradient, at the landscape scale and at the scale of a forest stand. In contrast, many of the non-red-listed generalist species are actually more likely to occur (per resource unit) in fragmented managed forests than well-connected natural forests. Synthesis. We show that the expected number of red-listed species per a fixed amount of similar resources (dead trees) can be even more than 10 times higher in well-connected than in fragmented surroundings, and thus, protecting high-quality areas that are well connected is conservationally more effective than protecting small fragments distributed across the landscape.

Evidence that acidification-induced declines in plant diversity and productivity are mediated by changes in below-ground communities and soil properties in a semi-arid steppe

Abstract: Summary 1. Anthropogenic acid deposition–induced soil acidification is one of the major threats to biodiversity, ecosystem functioning and services. Few studies, however, have explored in detail how above-ground changes in plant species richness and productivity resulting from soil acidification are mediated by effects on below-ground biota and soil properties. 2. To increase our understanding of this linkage, we collected data on below- and above-ground communities and soil properties in a 3-year field experiment with seven levels of acid addition rate to build-up broad intensities of soil acidification in the semi-arid Inner Mongolian grassland. 3. Acid addition directly elevated concentrations of soil Al 3+ ions, decreased the base cations Ca 2+ , Mg 2+ and Na + , and increased soil moisture and available phosphorus. Acid addition also appears to have altered the soil microbial community via changes in H + and Al 3+ ions and altered the nematode community via changes in H + ions and soil moisture. 4. The observed changes in soil N availability (i.e. net N mineralization, NO � -N and NH þ -N) could be explained by mediating changes in the H + and Al 3+ ions, microbial community (i.e. community structure, bacteria and fungi/bacteria as indicated by phospholipid fatty acids analysis) and the nematode community (i.e. total abundance, taxa richness and maturity index). 5. Declines in plant species richness and productivity were greater at high intensities of soil acidification in the second sampling year than in the first sampling year. The changes in plant community observed were mostly explained by soil nutrient pathways (e.g. N availability or base mineral cations), which were in turn regulated by the soil microbial or nematode communities as well as by the direct effects of the increase in H + or Al 3+ ions. 6. Synthesis. Our results suggest that the below-ground microbial and nematode communities are more sensitive to soil acidification than the plant communities are, and further that soil acidification–induced changes in plants are mediated by changes in below-ground communities and soil nutrients. These findings improve our understanding of the links between below- and above-ground communities in the Inner Mongolia grassland, especially in the context of anthropogenic acid enrichment.

Plant β-diversity in fragmented rain forests: testing floristic homogenization and differentiation hypotheses

Abstract: Summary Land-use change is the main driver of global biodiversity loss, but its relative impact on species turnover (β-diversity) across multiple spatial scales remains unclear. Plant communities in fragmented rain forests can undergo declines (floristic homogenization) or increases (floristic differentiation) in β-diversity. We tested these alternative hypotheses analysing a large vegetation data base from a hierarchically nested sampling design (450 plots in 45 forest patches in 3 landscapes with different deforestation levels) at Los Tuxtlas rain forest, Mexico. Differences in β-diversity across spatial scales (i.e. among plots, among patches, and among landscapes) were analysed using multiplicative diversity decompositions of Hill numbers. Plant β-diversity among plots within forest patches decreased in landscapes with higher deforestation levels, leading to floristic homogenization within patches. This homogenization process can be explained by the loss of rare and shade-tolerant plant species, and the recruitment and dominance of disturbance-adapted species, and can limit the accumulation of species (γ-diversity) in landscapes with higher deforestation. Nevertheless, the landscape with the highest deforestation level showed the highest floristic differentiation among patches. This landscape showed the greatest isolation distances between patches; a landscape spatial pattern that can limit the interchange of seeds (and species) between patches. Because the study patches are undergoing secondary succession following disturbances (e.g. logging, edge effects), different disturbance regimes and increased distance among patches could lead to higher β-diversity. Synthesis. These findings indicate that patterns of floristic homogenization and differentiation depend on the landscape configuration and on the spatial scale of analysis. At the landscape scale, our results suggest that, in accordance with non-equilibrium dynamics and the landscape-divergence hypothesis, patches located in landscapes with different forest cover and different connectivity can experience contrasting successional pathways due to increasing levels of compositional differentiation between patches. These novel findings add further uncertainties to the maintenance of biodiversity in severely deforested tropical landscapes and have key ecological implications for biodiversity conservation planning.

Microclimate in forests with varying leaf area index and soil moisture: potential implications for seedling establishment in a changing climate

Abstract: Summary 1. Forest microclimate is crucial for the growth and survival of tree seedlings and understorey vegetation. This high ecological relevance contrasts with the poor functional and quantitative understanding of how the properties of forest ecosystems influence forest microclimate. 2. In a long-term (1998–2011) trial, we investigated how temporal patterns of microclimate below sparse and dense forest canopy related to those of nearby open areas and how this relationship was influenced by soil moisture and seasonality. Air temperature (T), vapour pressure deficit (VPD), soil matrix potential and leaf area index (LAI) were measured in a unique set-up of below-canopy and open-area meteorological stations at eleven distinct forest ecosystems, characteristic of subalpine and temperate climate zones. Data from these plots were analysed for the moderating capacity of the canopy, that is, the differences between below-canopy and open-area microclimate, with respect to (i) long-term means, (ii) dynamics within homogeneous moist- vs. dry-soil periods and (iii) diurnal patterns. 3. The long-term mean moderating capacity of the canopy was up to 3.3 °C for daily Tmax and 0.52 kPa for daily VPDmax, of which soil moisture status alone accounted for up to 1.2 ° C( Tmax) and 0.21 kPa (VPDmax). Below dense canopy (LAI > 4), the moderating capacity was generally higher when soils were dry and increased during dry-soil periods, particularly in spring and somewhat less in summer. The opposite pattern was found below sparse canopy (LAI < 4). At the diurnal level, moderating capacity below dense canopy was strongest in mid-afternoon and during dry-soil conditions, whereas peak moderation below sparse canopy occurred in mid-morning and during moist-soil conditions. 4. Synthesis. Our results suggest a threshold canopy density, which is probably linked to sitespecific water availability, below which the moderating capacity of forest ecosystems switches from supportive to unsupportive for seedling establishment. Under supportive moderating capacity, we understand a stronger mitigation during physiologically most demanding conditions for plant growth. Such a threshold canopy density sheds new light on forest resilience to climate change. Climate change may alter forest canopy density in a way that precludes successful establishment of tree species and ultimately changes forest ecosystem structure and functioning.

Tree species diversity increases fine root productivity through increased soil volume filling

Abstract: Summary 1. Although fine roots (< 2 mm in diameter) account for a major share of the production of terrestrial ecosystems, diversity effects on fine root productivity and their mechanisms remain unclear. 2. We hypothesized that: (i) fine root productivity increases with tree species diversity, (ii) higher fine root productivity is a result of greater soil volume filling due to species-specific patterns of root placement and proliferation, and (iii) differences in fine root productivity and soil volume filling associated with tree species diversity are more pronounced in summer when plants are physiologically active and demand for water and nutrients is at its greatest. 3. We investigated the effects of tree species diversity on fine root productivity and soil volume filling of boreal forest stands that have grown naturally for 85 years on similar sites. 4. Annual fine root production was 19–83% higher in evenly mixed- than single-species-dominated stands, and increased with tree species evenness, but not tree species richness. Fine root biomass was higher in evenly mixed- than single-species-dominated stands in summer months, but not in spring or fall. Higher fine root productivity in evenly mixed- than single-species-dominated stands was realized by filling more soil volume horizontally and vertically in the forest floor in the mixtures of deep- and shallow-rooted species vs. the deeper mineral soil in the mixtures of deeprooted species. 5. Synthesis. Our results provide some of the first direct evidence for below-ground species complementarity in heterogeneous natural forests, by demonstrating that tree species evenness increases fine root productivity by filling/exploiting the soil environment more completely in space and time, driven by differences in the inherent rooting traits of the component species and variations of root growth within species.

Variability in functional traits mediates plant interactions along stress gradients

Abstract: Summary Environmental gradients may influence a plant's physiological status and morphology, which in turn may affect plant–plant interactions However, little is known about the relationship between environmental variation, physiological and morphological variability of plants and variation in the balance between competition and facilitation Mountain ranges in dry environments have opposing altitudinal environmental gradients of temperature and aridity, which limit plant growth at high and low elevations This makes them particularly suitable for exploring the relationships between environmental conditions, plant phenotype and plant–plant interactions We hypothesized that different environmental stressors will differently affect the physiological status of a nurse plant This, then, manifests itself as variation in nurse plant morphological traits, which in turn mediates plant–plant interactions by altering microhabitat conditions for the nurse and associated species In an observational study, we measured a series of functional traits of Arenaria tetraquetra cushions as indicators of its physiological status (eg specific leaf area, relative water content) and morphology (eg cushion compactness, size) Measurements were taken along the entire elevation range where A tetraquetra occurs Furthermore, we analysed how these functional traits related to soil properties beneath cushions and the number of associated species and individuals compared with open areas Cushions at high elevation showed good physiological status; they were compact and large, had higher soil water and organic matter content compared with open areas and showed the strongest facilitation effect of the whole elevation gradient – that is, the highest increase in species richness and abundance of beneficiaries compared with open areas Physiological data at low elevation indicated stressful abiotic conditions for A tetraquetra, which formed loose and small cushions These cushions showed less improved soil conditions and had reduced facilitative effects compared with those at high elevation Synthesis Functional traits of the nurse species varied distinctively along the two opposing stress gradients, in parallel to the magnitude of differences in microenvironmental conditions between cushions and the surrounding open area, and also to the facilitation effect of cushions Our data, therefore, provides a strong demonstration of the generally overlooked importance of a nurse plant's vigour and morphology for its facilitative effects

Hierarchical responses of plant-soil interactions to climate change:consequences for the global carbon cycle

Abstract: Interactions between plant and soil communities play a major role in determining the impact of climate change on ecosystem functioning and the carbon cycle, and the mechanisms involved operate over a wide range of spatial and temporal scales. We present a framework for understanding the consequences of climate-induced changes in plant–soil feedback for the carbon cycle. The framework describes a hierarchy of mechanisms by which changes in climate impact on ecosystem carbon dynamics at three levels of response, namely individual and community reordering and species immigration and loss. For each level, we identify the mechanisms by which climate change impacts on plant–soil interactions with consequences for the carbon cycle. We also demonstrate that the potential for decoupling of plant–soil interactions increases across the three levels of response, being greatest with species immigration and/or loss, for example, if plants were to undergo a biome shift, but their associated soil communities did not. Such decoupling is a largely unrecognized, but potentially important regulator of the future global carbon cycle. Synthesis. The framework presented here highlights a need for a new approach to the study of climate change impacts on plant–soil interactions and carbon cycling that integrates this hierarchy of responses, and incorporates the decoupling of above-ground and below-ground networks, across a range of temporal and spatial scales, and ecosystems.

Plant functional group identity and diversity determine biotic resistance to invasion by an exotic grass.

Abstract: Summary 1. Biotic resistance, the ability of species in a community to limit invasion, is central to our understanding of how communities at risk of invasion assemble after disturbances, but it has yet to translate into guiding principles for the restoration of invasion-resistant plant communities. We combined experimental, functional, and modelling approaches to investigate processes of community assembly contributing to biotic resistance to an introduced lineage of Phragmites australis, a model invasive species in North America. We hypothesized that (i) functional group identity would be a good predictor of biotic resistance to P. australis, while species identity effect would be redundant within functional group (ii) mixtures of species would be more invasion resistant than monocultures. 2. We classi! ed 36 resident wetland plants into four functional groups based on eight functional traits. We conducted two competition experiments based on the additive competition design with P. australis and monocultures or mixtures of wetland plants. As an indicator of biotic resistance, we calculated a relative competition index (RCIavg) based on the average performance of P. australis in competition treatment compared with control. To explain diversity effect further, we partitioned it into selection effect and complementarity effect and tested several diversity‐interaction models. 3. In monoculture treatments, RCIavg of wetland plants was signi! cantly different among functional groups, but not within each functional group. We found the highest RCIavg for fast-growing annuals, suggesting priority effect. 4. RCIavg of wetland plants was signi! cantly greater in mixture than in monoculture mainly due to complementarity‐diversity effect among functional groups. In diversity‐interaction models, species interaction patterns in mixtures were described best by interactions between functional groups when ! tted to RCIavg or biomass, implying niche partitioning. 5. Synthesis. Functional group identity and diversity of resident plant communities are good indicators of biotic resistance to invasion by introduced Phragmites australis, suggesting niche preemption (priority effect) and niche partitioning (diversity effect) as underlying mechanisms. Guiding principles to understand and/or manage biological invasion could emerge from advances in community theory and the use of a functional framework. Targeting widely distributed invasive plants in different contexts and scaling up to ! eld situations will facilitate generalization.

Tree effects on grass growth in savannas: competition, facilitation and the stress-gradient hypothesis

Abstract: Summary The stress-gradient hypothesis (SGH) predicts an increasing importance of facilitative mechanisms relative to competition along gradients of increasing environmental stress. Although developed across a variety of ecosystems, the SGH's relevance to the dynamic tree–grass systems of global savannas remains unclear. Here, we present a meta-analysis of empirical studies to explore emergent patterns of tree–grass relationships in global savannas in the context of the SGH. We quantified the net effect of trees on understorey grass production relative to production away from tree canopies along a rainfall gradient in tropical and temperate savannas and compared these findings to the predictions of the SGH. We also analysed soil and plant nutrient concentrations in subcanopy and open-grassland areas to investigate the potential role of nutrients in determining grass production in the presence and absence of trees. Our meta-analysis revealed a shift from net competitive to net facilitative effects of trees on subcanopy grass production with decreasing annual precipitation, consistent with the SGH. We also found a significant difference between sites from Africa and North America, suggesting differences in tree–grass interactions in the savannas of tropical and temperate regions. Nutrient analyses indicate no change in nutrient ratios along the rainfall gradient, but consistent nutrient enrichment under tree canopies. Synthesis. Our results help to resolve questions about the SGH in semi-arid systems, demonstrating that in mixed tree–grass systems, trees facilitate grass growth in drier regions and suppress grass growth in wetter regions. Relationships differ, however, between African and North American sites representing tropical and temperate bioclimates, respectively. The results of this meta-analysis advance our understanding of tree–grass interactions in savannas and contribute a valuable data set to the developing theory behind the SGH.

Biotic plant–soil feedbacks across temporal scales

Abstract: 1. Plant effects on soil biota can result in feedbacks affecting plant performance, with consequences for plant community and ecosystem dynamics on short and long time-scales. In addition, the strength and direction of plant-soil feedbacks depend on temporal shifts in abiotic environmental conditions. 2. We synthesize current knowledge on temporal aspects of plant-soil feedbacks and present new ideas to better understand and predict the effects of plant-soil feedbacks on community and ecosystem properties across temporal scales. 3. Explaining short-term temporal feedback dynamics requires us to better understand mechanistic linkages between plants, soil organisms and locally available resources. On the other hand, we need to refine our understanding of the context-dependency of plant-soil feedbacks, as the strength and direction of feedback interactions are influenced by 'external' temporal ecosystem dynamics, such as variation in soil resource availability after disturbance or during succession. 4. Synthesis. Based on our synthesis of temporal aspects of plant-soil feedbacks, we suggest three main avenues for future research: (i) how plant-soil feedbacks changes with ontogeny, (ii) how plant and soil organism traits drive temporal variation in plant-soil feedbacks and (iii) how environmental changes across temporal scales alter the strength and direction of plant-soil feedbacks.

Consequences of plant–soil feedbacks in invasion

Abstract: 1. Plant species can influence soil biota, which in turn can influence the relative performance of plant species. These plant–soil feedbacks (PSFs) have been hypothesized to affect many community-level dynamics including species coexistence, dominance and invasion. 2. The importance of PSFs in exotic species invasion, although widely hypothesized, has been difficult to determine because invader establishment necessarily precedes invader-mediated PSFs. Here, we combine a spatial simulation model of invasion that incorporates PSFs with a meta-analysis that synthesizes published case studies describing feedbacks between pairs of native and exotic species. 3. While our spatial model confirmed the link between positive soil feedbacks (‘home’ advantage) for exotic species and exotic species spread, results were dependent on the initial abundance of the exotic species and the equivalence of dispersal and life history characteristics between exotic and native species. 4. The meta-analysis of 52 native–exotic pairwise feedback comparisons in 22 studies synthesized measures of native and exotic performance in soils conditioned by native and exotic species. The analysis indicated that the growth responses of native species were often greater in soil conditioned by native species than in soil conditioned exotic species (a ‘home’ advantage). The growth responses of exotic species were variable and not consistently related to species soil-conditioning effects. 5. Synthesis. Overlaying empirical estimates of pairwise PSFs with spatial simulations, we conclude that the empirically measured PSFs between native and exotic plant species are often not consistent with predictions of the spread of exotic species and mono-dominance. This is particularly the case when exotic species are initially rare and share similar dispersal and average fitness characteristics with native species. However, disturbance and other processes that increase the abundance of exotic species as well as the inclusion of species dispersal and life history differences can interact with PSF effects to explain the spread of invasive species

Effects of litter on seedling establishment in natural and semi‐natural grasslands: a meta‐analysis

Abstract: 1. Plant litter is a key component in terrestrial ecosystems. It plays a major role in nutrient cycles and community organization. Land use and climate change may change the accumulation of litter in herbaceous ecosystems and affect plant community dynamics. Additionally, the transfer of seeds containing plant material (i.e. litter) is a widespread technique in grassland restoration.2. Ecosystem responses to litter represent the outcome of interactions, whose sign and strength will depend on many variables (e. g. litter amount, seed size). A previous meta-analysis (from 1999) reported that litter had an overall negative effect on seed germination and seedling establishment in different ecosystems. However, recent studies indicated that this might not be the case in grassland ecosystems.3. We used 914 data from 46 independent studies to analyse the effects of litter on seedling (i) emergence, (ii) survival and (iii) biomass, employing meta-analytical techniques. Each data set was stratified according to methodology, grassland type, irrigation conditions, litter amount and seed size.4. We found an overall neutral effect of litter presence on seedling emergence and survival and a positive effect on seedling biomass. However, whereas for field experiments the response remained neutral, it was positive for common garden studies. In glasshouse experiments, litter effects were negative for emergence and positive for biomass.5. Litter may have a positive effect on seedling recruitment in dry grasslands or under water-limited conditions, or in the presence of low to medium litter amounts ( 500 g m(-2)) will inhibit seedling recruitment. Large seeds showed a more positive response to litter presence with respect to seedling emergence and survival, but not concerning biomass.6. Synthesis. Under dry conditions (e. g. dry grasslands or dry periods) or with low to medium litter amounts, litter presence has a positive effect on seedling establishment. However, climate and land use change may promote litter accumulation and reduce seedling establishment, affecting grasslands composition and ecosystem functions.

Phylogenetic conservatism in plant phenology

Abstract: Summary 1. Phenological events – defined points in the life cycle of a plant or animal – have been regarded as highly plastic traits, reflecting flexible responses to various environmental cues. 2. The ability of a species to track, via shifts in phenological events, the abiotic environment through time might dictate its vulnerability to future climate change. Understanding the predictors and drivers of phenological change is therefore critical. 3. Here, we evaluated evidence for phylogenetic conservatism – the tendency for closely related species to share similar ecological and biological attributes – in phenological traits across flowering plants. We aggregated published and unpublished data on timing of first flower and first leaf, encompassing ~4000 species at 23 sites across the Northern Hemisphere. We reconstructed the phylogeny for the set of included species, first, using the software program Phylomatic, and second, from DNA data. We then quantified phylogenetic conservatism in plant phenology within and across sites. 4. We show that more closely related species tend to flower and leaf at similar times. By contrasting mean flowering times within and across sites, however, we illustrate that it is not the time of year that is conserved, but rather the phenological responses to a common set of abiotic cues. 5. Our findings suggest that species cannot be treated as statistically independent when modelling phenological responses. 6. Synthesis. Closely related species tend to resemble each other in the timing of their life-history events, a likely product of evolutionarily conserved responses to environmental cues. The search for the underlying drivers of phenology must therefore account for species’ shared evolutionary histories.

Pathogen accumulation and long-term dynamics of plant invasions

Abstract: Summary 1. The diversity of pathogens on highly abundant introduced hosts has been positively correlated with time since introduction, geographical range of the introduced species and diversity of invaded habitats. However, little is known about the ecological effects of pathogen accumulation on nonnative invasive plants. 2. Pathogen accumulation on invasive plant species may result from ecological processes such as high plant densities, expanding geographical ranges and pathogen dispersal from the native range, or evolutionary mechanisms such as host range shifts and adaptation of native pathogens to invasive species. 3. Over time pathogen accumulation may cause decline in the density and distribution of invasive plants and facilitate recovery of native species. Alternatively, pathogens might build up on invasive species and then spill back onto co-occurring native species, further exacerbating the effects of invasions. 4. Synthesis. Research efforts should focus on determining the long-term outcomes of pathogen accumulation on invasive species. Such research will require multifaceted approaches including comparative studies of diverse invasive species and habitats, experimental manipulations of hosts and pathogens in nature and controlled environments, and predictive models of host-pathogen interactions within an invasion framework. Results of this research will improve our understanding and ability to predict the outcomes of biological invasions.

Single-trait functional indices outperform multi-trait indices in linking environmental gradients and ecosystem services in a complex landscape

Abstract: Summary Functional traits can be used to describe the composition of communities through indices that seek to explain the factors that drive community assembly, biotic effects on ecosystem processes or both. Appropriately representing functional composition is therefore essential for predicting the consequences of environmental context and management actions for the provisioning of multiple ecosystem services (ESs) in heterogeneous landscapes. Functional indices can be constructed from single or multiple traits; however, it is not clear how they differ in information content or ability to predict biodiversity – ecosystem function relationships in complex landscapes. Here, we compare the utility of analogous single- and multi-trait indices in linking environmental variation and functional composition to ESs in a heterogeneous landscape, relating functional indices based on three plant traits [height, relative growth rate and root density (RD)] to variation in the physical environment and to two ESs (forage production and soil carbon) and their net ES level. Two orthogonal gradients, elevation and soil bulk density (BD), explained significant variation in several dimensions of functional composition comprised of single traits. These traits in turn significantly predicted variation in ESs and their net values. Only one index measured with multiple traits (functional richness) varied with the physical environment, while none predicted variation in ES or net ES levels. One ES, soil carbon, increased with the community-average value of RD, while the other, forage production, was related to the range and community-average value of height. In turn, average RD increased with soil BD while the average and range of height declined with elevation. Due to these environmental patterns, soil carbon and forage production did not covary strongly, leading to moderate net ES levels across the landscape. Synthesis: Single-trait indices of functional composition best linked variation in environmental gradients with productivity and soil carbon. Because the environment–trait functioning relationships were independent of one another, the ESs were independently distributed across the landscape, providing little evidence of synergies or trade-offs. Single- and multi-trait indices contained unique information about functional composition of these communities, and both are likely to have a place in predicting variation in ESs under different scenarios.

Fine root biomass and dynamics in beech forests across a precipitation gradient – is optimal resource partitioning theory applicable to water-limited mature trees?

Abstract: Summary Optimal resource partitioning theory predicts that plants should increase the ratio between water absorbing and transpiring surfaces under short water supply. An increase in fine root mass and surface area relative to leaf area has frequently been found in herbaceous plants, but supporting evidence from mature trees is scarce and several results are contradictory. In 12 mature Fagus sylvatica forests across a precipitation gradient (820–540 mm yr−1), we tested several predictions of the theory by analysing the dependence of standing fine root biomass, fine root production and fine root morphology on mean annual precipitation (MAP), the precipitation of the study year, and stand structural and edaphic variables. The water storage capacity of the soil (WSC) was included as a covariable by comparing pairs of stands on sandy (lower WSC) and loam-richer soils (higher WSC). Fine root biomass, total fine root surface area, fine root production and the fine root : leaf biomass production ratio markedly increased with reduced MAP and precipitation in the study year, while WSC was only a secondary factor and stand structure had no effect. The precipitation effect on fine root biomass and production was more pronounced in stands on sandy soil with lower WSC, which had, at equal precipitation, a higher fine root biomass and productivity than stands on loam-richer soil. The high degree of allocational plasticity in mature F. sylvatica trees contrasts with a low morphological plasticity of the fine roots. On the more extreme sandy soils, a significant decrease in mean fine root diameter and increase in specific root area with decreasing precipitation were found; a similar effect was absent on the loam-richer soils. Synthesis. In support of optimal partitioning theory, mature Fagus sylvatica trees showed a remarkable allocational plasticity as a long-term response to significant precipitation reduction with a large increase in the size and productivity of the fine root system, while only minor adaptive modifications occurred in root morphology. More severe summer droughts in a future warmer climate may substantially alter the above-/below-ground C partitioning of this tree species with major implications for the forest C cycle.

Imperfect detection is the rule rather than the exception in plant distribution studies

Abstract: Summary 1. Imperfect detection can seriously bias conventional estimators of species distributions and species richness. Plant traits, survey-specific conditions and site-specific characteristics may influence plant detection probability. However, the generality of the problems induced by imperfect detection in plants and the magnitude of this challenge for plant distribution studies are currently unknown. 2. We address this question based on data from the Swiss Biodiversity Monitoring, in which vascular plants are surveyed twice in the same year along a 2.5-km transect in 451 1-km 2 quadrats. Overall, 1700 species were recorded. We chose a random sample of 100 species from the 1700 species to determine general detection levels. To examine the relationship of covariates on detection, we chose a stratified random sample of 100 species from 886 species that were detected in at least 18 locations, with 25 each from four life-forms (LF): grass, forb, shrub and tree. Using a Bayesian multispecies site-occupancy model, we estimated occurrence and detection probability of these species and their relation to covariates. 3. Based on the random sample of 100 species, detection probability during the first survey ranged 0.03–0.99 (median 0.74) and during the second survey, 0.03–0.99 (median 0.82). Based on the stratified random sample of 100 species, detection probability during the first survey ranged 0.02–0.99 (median 0.87) and during the second survey, 0.01–1 (median 0.89). Detection probability differed slightly among the four LFs. In 60 species, survey season or elevation had significant effects on detection. We illustrated detection probability maps for Switzerland based on the modelled relationships with environmental covariates. 4. Synthesis. Our findings suggest that even in a standardized monitoring program, imperfect detection of plants may be common. With the absence of a correction for detection errors, maps in plant distribution studies will be confounded with spatial patterns in detection probability. We presume that these problems will be much more widespread in the data sets that are used for conventional plant species distribution modelling. Imperfect detection should be estimated, even in distribution studies of plants and other sessile organisms, to better control detection errors that may compromise the results of species distribution studies.

Central European hardwood trees in a high‐CO2 future: synthesis of an 8‐year forest canopy CO2 enrichment project

Abstract: Rapidly increasing atmospheric CO2 is not only changing the climate system but may also affect the biosphere directly through stimulation of plant growth and ecosystem carbon and nutrient cycling. Although forest ecosystems play a critical role in the global carbon cycle, experimental information on forest responses to rising CO2 is scarce, due to the sheer size of trees. Here, we present a synthesis of the only study world-wide where a diverse set of mature broadleaved trees growing in a natural forest has been exposed to future atmospheric CO2 levels (c. 550ppm) by free-air CO2 enrichment (FACE). We show that litter production, leaf traits and radial growth across the studied hardwood species remained unaffected by elevated CO2 over 8years. CO2 enrichment reduced tree water consumption resulting in detectable soil moisture savings. Soil air CO2 and dissolved inorganic carbon both increased suggesting enhanced below-ground activity. Carbon release to the rhizosphere and/or higher soil moisture primed nitrification and nitrate leaching under elevated CO2; however, the export of dissolved organic carbon remained unaltered.Synthesis. Our findings provide no evidence for carbon-limitation in five central European hardwood trees at current ambient CO2 concentrations. The results of this long-term study challenge the idea of a universal CO2 fertilization effect on forests, as commonly assumed in climate-carbon cycle models.

Uncovering multiscale effects of aridity and biotic interactions on the functional structure of Mediterranean shrublands

Abstract: Summary 1. Habitat filtering (HF, trait convergence) and niche differentiation (ND, trait divergence) are known to impact upon plant community structure. Both processes integrate individual responses to the abiotic environment and biotic interactions. Thus, it is difficult to clearly identify the underlying abiotic and biotic factors that ultimately impact community structure by looking at community-level patterns of trait divergence or convergence alone. 2. We used a functional trait-based and multiscale approach to assess how biotic interactions and aridity determine the functional structure of semi-arid shrublands sampled along a large aridity gradient in Spain. At the regional scale, we investigated functional differences among species (axes of specialization) to identify important traits for community assembly. At the community scale, we evaluated the relative impact of HF and ND on community structure using a null model approach. Finally, at the plant neighbourhood scale, we evaluated the impact of biotic interactions on community structure by investigating the spatial patterns of trait aggregation. 3. The shrub species surveyed can be separated along four axes of specialization based on their above-ground architecture and leaf morphology. Our community scale analysis suggested that the functional structure of semi-arid communities was clearly non-random, HF and ND acting independently on different traits to determine community structure along the aridity gradient. At the plant neighbourhood scale, the spatial distribution of species was also clearly not random, suggesting that competition and facilitation impacted on the observed changes in the functional diversity of shrubland communities along the aridity gradient. 4. Synthesis: Our results demonstrated that HF and ND acted simultaneously on independent traits to jointly determine community structure. Most importantly, our multiscale approach suggested that competition and facilitation interplayed with aridity to determine this structure. Competition appeared to be constant along the aridity gradient and explained the high functional diversity observed in semi-arid shrublands. Facilitation affected subordinate and rare species and, thus, may act to enhance the biodiversity of these ecosystems. Finally, the framework employed in our study allows moving forward from the examination of patterns to the development of mechanistic traitbased approaches to study plant community assembly.