Experimental drought indirectly enhances the individual performance and the abundance of an invasive annual weed.

During environmental change, invasive species may be favored by increased resource input or reduced resource use of the resident community. Plasticity in certain plant traits of invasive species may be one possible mechanism behind their ability to quickly exploit unused resources. We tested whether rainfall manipulations (severe drought, moderate drought, watering) alter the growth and reproductive success of the invasive annual Conyza canadensis, and if it translates into a change in the abundance of the species in a semiarid perennial grassland in Central Hungary. Overall, C. canadensis exhibited greater individual performance and higher abundance in drought plots than in control and watered plots. At individual level, plants showed the strongest response to moderate drought: they grew 2.5-times taller than in control and watered plots, and produced twice and 2.5-times more seeds than in watered and control plots, respectively. Reproductive phenology was advanced in response to rain exclusions. Although severe drought caused 40% mortality, the cumulative performance of C. canadensis, expressed as plot-level aboveground biomass, was consistently greater in severe drought plots than in control and watered plots throughout the 3 years of the study. The higher performance of C. canadensis in drought plots is most likely due to the decreased abundance and, thus, decreased competitive effect of previously dominant perennial grasses caused by the rain exclusions. We conclude that drier summers that suppress perennial grasses will favor this alien annual forb, and phenotypic plasticity in growth and reproduction may contribute to its invasion success.

Fast attrition of springtail communities by experimental drought and richness-decomposition relationships across Europe.

Soil fauna play a fundamental role on key ecosystem functions like organic matter decomposition, although how local assemblages are responding to climate change and whether these changes may have consequences to ecosystem functioning is less clear. Previous studies have revealed that a continued environmental stress may result in poorer communities by filtering out the most sensitive species. However, these experiments have rarely been applied to climate change factors combining multiyear and multisite standardized field treatments across climatically contrasting regions, which has limited drawing general conclusions. Moreover, other facets of biodiversity, such as functional and phylogenetic diversity, potentially more closely linked to ecosystem functioning, have been largely neglected. Here, we report that the abundance, species richness, phylogenetic diversity, and functional richness of springtails (Subclass Collembola), a major group of fungivores and detritivores, decreased within 4 years of experimental drought across six European shrublands. The loss of phylogenetic and functional richness was higher than expected by the loss of species richness, leading to communities of phylogenetically similar species sharing evolutionary conserved traits. Additionally, despite the great climatic differences among study sites, we found that taxonomic, phylogenetic, and functional richness of springtail communities alone were able to explain up to 30% of the variation in annual decomposition rates. Altogether, our results suggest that the forecasted reductions in precipitation associated with climate change may erode springtail communities and likely other drought-sensitive soil invertebrates, thereby retarding litter decomposition and nutrient cycling in ecosystems.

Within-generation and transgenerational plasticity in growth and regeneration of a subordinate annual grass in a rainfall experiment.

Precipitation changes may induce shifts in plant species or life form dominance in ecosystems, making some previously subordinate species abundant. The plasticity of certain plant functional traits of these expanding subordinate species may be one possible mechanism behind their success. In this study, we tested if the subordinate winter annual grass Secale sylvestre shows plasticity in growth and reproduction in response to altered environment associated with field-scale rainfall manipulations (severe drought, moderate drought, and watering) in a semiarid grassland, and whether the maternal environment influences offspring germination or growth in a subsequent pot experiment. Compared to control plots, S. sylvestre plants grew 38% taller, and produced 32% more seeds in severe drought plots, while plants in watered plots were 17% shorter, and had 22% less seeds. Seed mass was greatest in severe drought plots. Plants growing in drought plots had offspring with enhanced juvenile shoot growth compared to the progeny whose mother plants grew in watered plots. These responses are most likely explained by the decreased cover of previously dominant perennial grasses in severe drought plots, which resulted in wetter soil compared to control and watered plots during the peak growth of S. sylvestre. We conclude that the plasticity of this subordinate annual species in response to changing environment may help to gain dominance with recurring droughts that suppress perennial grasses. Our results highlight that exploring both within-generation and transgenerational plasticity of subordinate species may lead to a better prediction of changes in plant species dominance under climate change.

Few multiyear precipitation-reduction experiments find a shift in the productivity-precipitation relationship.

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.

How climate, topography, soils, herbivores, and fire control forest-grassland coexistence in the Eurasian forest-steppe.

Recent advances in ecology and biogeography demonstrate the importance of fire and large herbivores - and challenge the primacy of climate - to our understanding of the distribution, stability, and antiquity of forests and grasslands. Among grassland ecologists, particularly those working in savannas of the seasonally dry tropics, an emerging fire-herbivore paradigm is generally accepted to explain grass dominance in climates and on soils that would otherwise permit development of closed-canopy forests. By contrast, adherents of the climate-soil paradigm, particularly foresters working in the humid tropics or temperate latitudes, tend to view fire and herbivores as disturbances, often human-caused, which damage forests and reset succession. Towards integration of these two paradigms, we developed a series of conceptual models to explain the existence of an extensive temperate forest-grassland mosaic that occurs within a 4.7 million km2 belt spanning from central Europe through eastern Asia. The Eurasian forest-steppe is reminiscent of many regions globally where forests and grasslands occur side-by-side with stark boundaries. Our conceptual models illustrate that if mean climate was the only factor, forests should dominate in humid continental regions and grasslands should prevail in semi-arid regions, but that extensive mosaics would not occur. By contrast, conceptual models that also integrate climate variability, soils, topography, herbivores, and fire depict how these factors collectively expand suitable conditions for forests and grasslands, such that grasslands may occur in more humid regions and forests in more arid regions than predicted by mean climate alone. Furthermore, boundaries between forests and grasslands are reinforced by vegetation-fire, vegetation-herbivore, and vegetation-microclimate feedbacks, which limit tree establishment in grasslands and promote tree survival in forests. Such feedbacks suggest that forests and grasslands of the Eurasian forest-steppe are governed by ecological dynamics that are similar to those hypothesised to maintain boundaries between tropical forests and savannas. Unfortunately, the grasslands of the Eurasian forest-steppe are sometimes misinterpreted as deforested or otherwise degraded vegetation. In fact, the grasslands of this region provide valuable ecosystem services, support a high diversity of plants and animals, and offer critical habitat for endangered large herbivores. We suggest that a better understanding of the fundamental ecological controls that permit forest-grassland coexistence could help us prioritise conservation and restoration of the Eurasian forest-steppe for biodiversity, climate adaptation, and pastoral livelihoods. Currently, these goals are being undermined by tree-planting campaigns that view the open grasslands as opportunities for afforestation. Improved understanding of the interactive roles of climate variability, soils, topography, fire, and herbivores will help scientists and policymakers recognise the antiquity of the grasslands of the Eurasian forest-steppe.

The response of litter decomposition to extreme drought modified by plant species, plant part, and soil depth in a temperate grassland.

Plant litter decomposition is a key ecosystem process in carbon and nutrient cycling, and is heavily affected by changing climate. While the direct effects of drought on decomposition are widely studied, in order to better predict the overall drought effect, indirect effects associated with various drought-induced changes in ecosystems should also be quantified. We studied the effect of an extreme (5-month) experimental drought on decomposition, and if this effect varies with two dominant perennial grasses, plant parts (leaves vs. roots), and soil depths (0-5 cm vs. 10-15 cm) in a semi-arid temperate grassland. After 12 months, the average litter mass loss was 43.5% in the control plots, while only 25.7% in the drought plots. Overall, mass loss was greater for leaves (44.3%) compared to roots (24.9%), and for Festuca vaginata (38.6%) compared to Stipa borysthenica (30.5%). This variation was consistent with the observed differences in nitrogen and lignin content between plant parts and species. Mass loss was greater for deep soil (42.8%) than for shallow soil (26.4%). Collectively, these differences in decomposition between the two species, plant parts, and soil depths were similar in magnitude to direct drought effect. Drought induces multiple changes in ecosystems, and our results highlight that these changes may in turn modify decomposition. We conclude that for a reliable estimate of decomposition rates in an altered climate, not only direct but also indirect climatic effects should be considered, such as those arising from changing species dominance, root-to-shoot ratio, and rooting depth.

Field experiments underestimate aboveground biomass response to drought.

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.

Beyond the Forest-Grassland Dichotomy: The Gradient-Like Organization of Habitats in Forest-Steppes.

Featuring a transitional zone between closed forests and treeless steppes, forest-steppes cover vast areas, and have outstanding conservation importance. The components of this mosaic ecosystem can conveniently be classified into two basic types, forests and grasslands. However, this dichotomic classification may not fit reality as habitat organization can be much more complex. In this study, our aim was to find out if the main habitat types can be grouped into two distinct habitat categories (which would support the dichotomic description), or a different paradigm better fits this complex ecosystem. We selected six main habitats of sandy forest-steppes, and, using 176 relevés, we compared their vegetation based on species composition (NMDS ordination, number of common species of the studied habitats), relative ecological indicator values (mean indicators for temperature, soil moisture, and light availability), and functional species groups (life-form categories, geoelement types, and phytosociological preference groups). According to the species composition, we found a well-defined gradient, with the following habitat order: large forest patches, medium forest patches, small forest patches, north-facing edges, south-facing edges, and grasslands. A considerable number of species were shared among all habitats, while the number of species restricted to certain habitat types was also numerous, especially for north-facing edges. The total (i.e., pooled) number of species peaked near the middle of the gradient, in north-facing edges. The relative ecological indicator values and functional species groups showed mostly gradual changes from the large forest patches to the grasslands. Our results indicate that the widely used dichotomic categorization of forest-steppe habitats into forest and grassland patches is too simplistic, potentially resulting in a considerable loss of information. We suggest that forest-steppe vegetation better fits the gradient-based paradigm of landscape structure, which is able to reflect continuous variations.

Effects of single and repeated drought on soil microarthropods in a semi-arid ecosystem depend more on timing and duration than drought severity.

Soil moisture is one of the most important factors affecting soil biota. In arid and semi-arid ecosystems, soil mesofauna is adapted to temporary drought events, but, until now, we have had a limited understanding of the impacts of the different magnitudes and frequencies of drought predicted to occur according to future climate change scenarios. The present study focuses on how springtails and mites respond to simulated repeated drought events of different magnitudes in a field experiment in a Hungarian semi-arid sand steppe. Changes in soil arthropod activities were monitored with soil trapping over two years in a sandy soil. In the first year (2014), we applied an extreme drought pretreatment, and in the consecutive year, we applied less devastating treatments (severe drought, moderate drought, water addition) to these sites. In the first year, the extreme drought pretreatment tended to have a negative effect (either significantly or not significantly) on the capture of all Collembola groups, whereas all mite groups increased in activity density. However, in the consecutive year, between the extreme drought and control treatments, we only detected differences in soil microbial biomass. In the cases of severe drought, moderate drought and water addition, we did not find considerable changes across the microarthropods, except in the case of epedaphic Collembola. In the cases of the water addition and drought treatments, the duration and timing of the manipulation seemed to be more important for soil mesofauna than their severity (i.e., the level of soil moisture decrease). We suggest that in these extreme habitats, soil mesofauna are able to survive extreme conditions, and their populations recover rapidly, but they may not be able to cope with very long drought periods.

Mean annual precipitation predicts primary production resistance and resilience to extreme drought.

Extreme drought is increasing in frequency and intensity in many regions globally, with uncertain consequences for the resistance and resilience of ecosystem functions, including primary production. Primary production resistance, the capacity to withstand change during extreme drought, and resilience, the degree to which production recovers, vary among and within ecosystem types, obscuring generalized patterns of ecological stability. Theory and many observations suggest forest production is more resistant but less resilient than grassland production to extreme drought; however, studies of production sensitivity to precipitation variability indicate that the processes controlling resistance and resilience may be influenced more by mean annual precipitation (MAP) than ecosystem type. Here, we conducted a global meta-analysis to investigate primary production resistance and resilience to extreme drought in 64 forests and grasslands across a broad MAP gradient. We found resistance to extreme drought was predicted by MAP; however, grasslands (positive) and forests (negative) exhibited opposing resilience relationships with MAP. Our findings indicate that common plant physiological mechanisms may determine grassland and forest resistance to extreme drought, whereas differences among plant residents in turnover time, plant architecture, and drought adaptive strategies likely underlie divergent resilience patterns. The low resistance and resilience of dry grasslands suggests that these ecosystems are the most vulnerable to extreme drought - a vulnerability that is expected to compound as extreme drought frequency increases in the future.

Shrubland primary production and soil respiration diverge along European climate gradient.

Above- and belowground carbon (C) stores of terrestrial ecosystems are vulnerable to environmental change. Ecosystem C balances in response to environmental changes have been quantified at individual sites, but the magnitudes and directions of these responses along environmental gradients remain uncertain. Here we show the responses of ecosystem C to 8-12 years of experimental drought and night-time warming across an aridity gradient spanning seven European shrublands using indices of C assimilation (aboveground net primary production: aNPP) and soil C efflux (soil respiration: Rs). The changes of aNPP and Rs in response to drought indicated that wet systems had an overall risk of increased loss of C but drier systems did not. Warming had no consistent effect on aNPP across the climate gradient, but suppressed Rs more at the drier sites. Our findings suggest that above- and belowground C fluxes can decouple, and provide no evidence of acclimation to environmental change at a decadal timescale. aNPP and Rs especially differed in their sensitivity to drought and warming, with belowground processes being more sensitive to environmental change.

Increased sensitivity to climate change in disturbed ecosystems.

Human domination of the biosphere includes changes to disturbance regimes, which push many ecosystems towards early-successional states. Ecological theory predicts that early-successional ecosystems are more sensitive to perturbations than mature systems, but little evidence supports this relationship for the perturbation of climate change. Here we show that vegetation (abundance, species richness and species composition) across seven European shrublands is quite resistant to moderate experimental warming and drought, and responsiveness is associated with the dynamic state of the ecosystem, with recently disturbed sites responding to treatments. Furthermore, most of these responses are not rapid (2-5 years) but emerge over a longer term (7-14 years). These results suggest that successional state influences the sensitivity of ecosystems to climate change, and that ecosystems recovering from disturbances may be sensitive to even modest climatic changes. A research bias towards undisturbed ecosystems might thus lead to an underestimation of the impacts of climate change.