Jürgen Kreyling

A new generation of climate change experiments : Events, not trends

Intensification of weather extremes is currently emerging as one of the most important facets of climate change. Research on extreme events (“event-focused” in contrast to “trend-focused”) has increased in recent years and, in 2004, accounted for one-fifth of the experimental climate-change studies published. Numerous examples, ranging from microbiology and soil science to biogeography, demonstrate how extreme weather events can accelerate shifts in species composition and distribution, thereby facilitating changes in ecosystem functioning. However, assessing the importance of extreme events for ecological processes poses a major challenge because of the very nature of such events: their effects are out of proportion to their short duration. We propose that extreme events can be characterized by statistical extremity, timing, and abruptness relative to the life cycles of the organisms affected. To test system response to changing magnitude and frequency of weather events, controlled experiments are useful tools. These experiments provide essential insights for science and for societies that must develop coping strategies for such events. Here, we discuss future research needs for climate-change experiments in ecology. For illustration, we describe an experimental plan showing how to meet the challenge posed by changes in the frequency or magnitude of extreme events.

Biodiversity Increases the Resistance of Ecosystem Productivity to Climate Extremes

It remains unclear whether biodiversity buffers ecosystems against climate extremes, which are becoming increasingly frequent worldwide. Early results suggested that the ecosystem productivity of diverse grassland plant communities was more resistant, changing less during drought, and more resilient, recovering more quickly after drought, than that of depauperate communities. However, subsequent experimental tests produced mixed results. Here we use data from 46 experiments that manipulated grassland plant diversity to test whether biodiversity provides resistance during and resilience after climate events. We show that biodiversity increased ecosystem resistance for a broad range of climate events, including wet or dry, moderate or extreme, and brief or prolonged events. Across all studies and climate events, the productivity of low-diversity communities with one or two species changed by approximately 50% during climate events, whereas that of high-diversity communities with 16–32 species was more resistant, changing by only approximately 25%. By a year after each climate event, ecosystem productivity had often fully recovered, or overshot, normal levels of productivity in both high- and low-diversity communities, leading to no detectable dependence of ecosystem resilience on biodiversity. Our results suggest that biodiversity mainly stabilizes ecosystem productivity, and productivity-dependent ecosystem services, by increasing resistance to climate events. Anthropogenic environmental changes that drive biodiversity loss thus seem likely to decrease ecosystem stability, and restoration of biodiversity to increase it, mainly by changing the resistance of ecosystem productivity to climate events.

Stochastic trajectories of succession initiated by extreme climatic events

Deterministic or rule-based succession is expected under homogeneous biotic and abiotic starting conditions. Effects of extreme climatic events such as drought, however, may alter these assembly rules by adding stochastic elements. We monitored the succession of species composition of 30 twin grassland communities with identical biotic and abiotic starting conditions in an initially sown diversity gradient between 1 and 16 species over 13 years. The stochasticity of succession, measured as the synchrony in the development of the species compositions of the twin plots, was strongly altered by the extreme warm and dry summer of 2003. Moreover, it was independent from past and present plant diversity and neighbourhood species compositions. Extreme climatic events can induce stochastic effects in community development and therefore impair predictability even under homogeneous abiotic conditions. Stochastic events may result in lasting shifts of community composition, as well as adverse and unforeseeable effects on the stability of ecological services.

Late frost sensitivity of juvenile Fagus sylvatica L. differs between southern Germany and Bulgaria and depends on preceding air temperature

Fagus sylvatica, the dominant native forest tree species of Central Europe, is sensitive to late frost events. Advanced leaf flushing due to climate warming may lead to more frequent frost damage in the future. Here, we explore local adaptation to late frost events at both continental and regional scales and test how moderate climate warming (+1.5°C) affects late frost sensitivity. Short-term leaf injury and height growth after a late frost event were quantified in a common garden experiment with 2-year-old F. sylvatica seedlings. The fully crossed three-factorial design consisted of a late frost manipulation, a continuous warming manipulation and selected provenances (three provenances from western Bulgaria and three from southern Germany). Late frost led to leaf injury and reduced height growth (−7%). Provenances differed in their late frost sensitivity at the regional scale, and local adaptation was detected. At the larger scale, the Bulgarian provenances showed reduced height growth (−17%), while the German provenances did not exhibit growth reduction. The warming treatment prevented late frost damage, while height growth declined by 19% in the reference temperature treatment. This surprising finding was attributed to advanced leaf maturity in the warming treatment. The impact of late frost events on F. sylvatica in a warmer world will depend on timing. An event that damages leaves immediately after leaf flushing appears negligible a few days earlier or later, thereby complicating projections. Local adaptation to late frost is evident at a regional scale. Management strategies should aim at maximizing genetic diversity to adapt to climate change.

Plant diversity effects on grassland productivity are robust to both nutrient enrichment and drought

Global change drivers are rapidly altering resource availability and biodiversity. While there is consensus that greater biodiversity increases the functioning of ecosystems, the extent to which biodiversity buffers ecosystem productivity in response to changes in resource availability remains unclear. We use data from 16 grassland experiments across North America and Europe that manipulated plant species richness and one of two essential resources—soil nutrients or water—to assess the direction and strength of the interaction between plant diversity and resource alteration on above-ground productivity and net biodiversity, complementarity, and selection effects. Despite strong increases in productivity with nutrient addition and decreases in productivity with drought, we found that resource alterations did not alter biodiversity–ecosystem functioning relationships. Our results suggest that these relationships are largely determined by increases in complementarity effects along plant species richness gradients. Although nutrient addition reduced complementarity effects at high diversity, this appears to be due to high biomass in monocultures under nutrient enrichment. Our results indicate that diversity and the complementarity of species are important regulators of grassland ecosystem productivity, regardless of changes in other drivers of ecosystem function.

Plant responses to climatic extremes: within-species variation equals among-species variation.

Within-species and among-species differences in growth responses to a changing climate have been well documented, yet the relative magnitude of within-species vs. among-species variation has remained largely unexplored. This missing comparison impedes our ability to make general predictions of biodiversity change and to project future species distributions using models. We present a direct comparison of among- versus within-species variation in response to three of the main stresses anticipated with climate change: drought, warming, and frost. Two earlier experiments had experimentally induced (i) summer drought and (ii) spring frost for four common European grass species and their ecotypes from across Europe. To supplement existing data, a third experiment was carried out, to compare variation among species from different functional groups to within-species variation. Here, we simulated (iii) winter warming plus frost for four grasses, two nonleguminous, and two leguminous forbs, in addition to eleven European ecotypes of the widespread grass Arrhenatherum elatius. For each experiment, we measured: (i) C/N ratio and biomass, (ii) chlorophyll content and biomass, and (iii) plant greenness, root (15) N uptake, and live and dead tissue mass. Using coefficients of variation (CVs) for each experiment and response parameter, a total of 156 within- vs. among-species comparisons were conducted, comparing within-species variation in each of four species with among-species variation for each seed origin (five countries). Of the six significant differences, within-species CVs were higher than among-species CVs in four cases. Partitioning of variance within each treatment in two of the three experiments showed that within-species variability (ecotypes) could explain an additional 9% of response variation after accounting for the among-species variation. Our observation that within-species variation was generally as high as among-species variation emphasizes the importance of including both within- and among-species variability in ecological theory (e.g., the insurance hypothesis) and for practical applications (e.g., biodiversity conservation).

Drought Effects in Climate Change Manipulation Experiments: Quantifying the Influence of Ambient Weather Conditions and Rain-out Shelter Artifacts

Extreme drought events challenge ecosystem functioning. Ecological response to drought is studied worldwide in a growing number of field experiments by rain-out shelters. Yet, few meta-analyses face severe challenges in the comparability of studies. This is partly because build-up of drought stress in rain-out shelters is modified by ambient weather conditions. Rain-out shelters can further create confounding effects (radiation, temperature), which may influence plant responses. Yet, a quantification of ecophysiological effects within rain-out shelters under opposing ambient weather conditions and of microclimatological artifacts is missing. Here, we examined phytometers—standardized potted individuals of Plantago lanceolata—under rain-out shelter, rain-out shelter artifact control, and ambient control during opposing outside microclimatological conditions. Furthermore, we tested for artifacts of rain-out shelters on plant responses in a long-term semi-natural grassland experiment. Phytometer plants below the rain-out shelters showed lower stomatal conductance, maximum quantum efficiency, and leaf water potential during warm ambient conditions with high evaporative demand than during cold conditions with low evaporative demand. Plant performance was highly correlated with ambient temperature and vapor pressure deficit (VPD). Rain-out shelter artifacts on plant responses were nonsignificant. Rain-out shelters remain a viable tool for studying ecosystem responses to drought. However, drought manipulations using rain-out shelters are strongly modified by ambient weather conditions. Attributing the results from rain-out shelter studies to drought effects and comparability among studies and study years therefore requires the quantification of the realized drought stress, for example, by relating ecosystem responses to measured microclimatological parameters such as air temperature and VPD.

Multiple facets of biodiversity drive the diversity–stability relationship

A substantial body of evidence has demonstrated that biodiversity stabilizes ecosystem functioning over time in grassland ecosystems. However, the relative importance of different facets of biodiversity underlying the diversity–stability relationship remains unclear. Here we use data from 39 grassland biodiversity experiments and structural equation modelling to investigate the roles of species richness, phylogenetic diversity and both the diversity and community-weighted mean of functional traits representing the ‘fast–slow’ leaf economics spectrum in driving the diversity–stability relationship. We found that high species richness and phylogenetic diversity stabilize biomass production via enhanced asynchrony in the performance of co-occurring species. Contrary to expectations, low phylogenetic diversity enhances ecosystem stability directly, albeit weakly. While the diversity of fast–slow functional traits has a weak effect on ecosystem stability, communities dominated by slow species enhance ecosystem stability by increasing mean biomass production relative to the standard deviation of biomass over time. Our in-depth, integrative assessment of factors influencing the diversity–stability relationship demonstrates a more multicausal relationship than has been previously acknowledged.Analysing data from 39 grassland biodiversity experiments, the authors uncover the direct and indirect contributions to ecosystem stability of taxonomic, phylogenetic and functional trait diversity.

On the influence of provenance to soil quality enhanced stress reaction of young beech trees to summer drought

Abstract Climate projections propose that drought stress will become challenging for establishing trees. The magnitude of stress is dependent on tree species, provenance, and most likely also highly influenced by soil quality. European Beech (Fagus sylvatica) is of major ecological and economical importance in Central European forests. The species has an especially wide physiological and ecological amplitude enabling growth under various soil conditions within its distribution area in Central Europe. We studied the effects of extreme drought on beech saplings (second year) of four climatically distinct provenances growing on different soils (sandy loam and loamy sand) in a full factorial pot experiment. Foliar δ13C, δ15N, C, and N as well as above‐ and belowground growth parameters served as measures for stress level and plant growth. Low‐quality soil enhanced the effect of drought compared with qualitatively better soil for the above‐ and belowground growth parameters, but foliar δ13C values revealed that plant stress was still remarkable in loamy soil. For beeches of one provenance, negative sandy soil effects were clearly smaller than for the others, whereas for another provenance drought effects in sandy soil were sometimes fatal. Foliar δ15N was correlated with plant size during the experiment. Plasticity of beech provenances in their reaction to drought versus control conditions varied clearly. Although a general trend of declining growth under control or drought conditions in sandy soil was found compared to loamy soil, the magnitude of the effect of soil quality was highly provenance specific. Provenances seemed to show adaptations not only to drought but also to soil quality. Accordingly, scientists should integrate information about climatic pre‐adaptation and soil quality within the home range of populations for species distribution modeling and foresters should evaluate soil quality and climatic parameters when choosing donor populations for reforestation projects.