Carl Beierkuhnlein

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.

Ecosystem effects of biodiversity manipulations in European grasslands.

We present a multisite analysis of the relationship between plant diversity and ecosystem functioning within the European BIODEPTH network of plant-diversity manipulation experiments. We report results of the analysis of 11 variables addressing several aspects of key ecosystem processes like biomass production, resource use (space, light, and nitrogen), and decomposition, measured across three years in plots of varying plant species richness at eight different European grassland field sites. Differences among sites explained substantial and significant amounts of the variation of most of the ecosystem processes examined. However, against this background of geographic variation, all the aspects of plant diversity and composition we examined (i.e., both numbers and types of species and functional groups) produced significant, mostly positive impacts on ecosystem processes. Analyses using the additive partitioning method revealed that complementarity effects (greater net yields than predicted from monocultures due to resource partitioning, positive interactions, etc.) were stronger and more consistent than selection effects (the covariance between monoculture yield and change in yield in mixtures) caused by dominance of species with particular traits. In general, communities with a higher diversity of species and functional groups were more productive and utilized resources more completely by intercepting more light, taking up more nitrogen, and occupying more of the available space. Diversity had significant effects through both increased vegetation cover and greater nitrogen retention by plants when this resource was more abundant through N2 fixation by legumes. However, additional positive diversity effects remained even after controlling for differences in vegetation cover and for the presence of legumes in communities. Diversity effects were stronger on above- than belowground processes. In particular, clear diversity effects on decomposition were only observed at one of the eight sites. The ecosystem effects of plant diversity also varied between sites and years. In general, diversity effects were lowest in the first year and stronger later in the experiment, indicating that they were not transitional due to community establishment. These analyses of our complete ecosystem process data set largely reinforce our previous results, and those from comparable biodiversity experiments, and extend the generality of diversity–ecosystem functioning relationships to multiple sites, years, and processes.

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.

Precipitation manipulation experiments: challenges and recommendations for the future

Climatic changes, including altered precipitation regimes, will affect key ecosystem processes, such as plant productivity and biodiversity for many terrestrial ecosystems. Past and ongoing precipitation experiments have been conducted to quantify these potential changes. An analysis of these experiments indicates that they have provided important information on how water regulates ecosystem processes. However, they do not adequately represent global biomes nor forecasted precipitation scenarios and their potential contribution to advance our understanding of ecosystem responses to precipitation changes is therefore limited, as is their potential value for the development and testing of ecosystem models. This highlights the need for new precipitation experiments in biomes and ambient climatic conditions hitherto poorly studied applying relevant complex scenarios including changes in precipitation frequency and amplitude, seasonality, extremity and interactions with other global change drivers. A systematic and holistic approach to investigate how soil and plant community characteristics change with altered precipitation regimes and the consequent effects on ecosystem processes and functioning within these experiments will greatly increase their value to the climate change and ecosystem research communities. Experiments should specifically test how changes in precipitation leading to exceedance of biological thresholds affect ecosystem resilience and acclimation.

Inventory, differentiation, and proportional diversity: a consistent terminology for quantifying species diversity

Almost half a century after Whittaker (Ecol Monogr 30:279–338, 1960) proposed his influential diversity concept, it is time for a critical reappraisal. Although the terms alpha, beta and gamma diversity introduced by Whittaker have become general textbook knowledge, the concept suffers from several drawbacks. First, alpha and gamma diversity share the same characteristics and are differentiated only by the scale at which they are applied. However, as scale is relative––depending on the organism(s) or ecosystems investigated––this is not a meaningful ecological criterion. Alpha and gamma diversity can instead be grouped together under the term “inventory diversity.” Out of the three levels proposed by Whittaker, beta diversity is the one which receives the most contradictory comments regarding its usefulness (“key concept” vs. “abstruse concept”). Obviously beta diversity means different things to different people. Apart from the large variety of methods used to investigate it, the main reason for this may be different underlying data characteristics. A literature review reveals that the multitude of measures used to assess beta diversity can be sorted into two conceptually different groups. The first group directly takes species distinction into account and compares the similarity of sites (similarity indices, slope of the distance decay relationship, length of the ordination axis, and sum of squares of a species matrix). The second group relates species richness (or other summary diversity measures) of two (or more) different scales to each other (additive and multiplicative partitioning). Due to that important distinction, we suggest that beta diversity should be split into two levels, “differentiation diversity” (first group) and “proportional diversity” (second group). Thus, we propose to use the terms “inventory diversity” for within-sample diversity, “differentiation diversity” for compositional similarity between samples, and “proportional diversity” for the comparison of inventory diversity across spatial and temporal scales.

Coordinated distributed experiments: an emerging tool for testing global hypotheses in ecology and environmental science

There is a growing realization among scientists and policy makers that an increased understanding of todays environmental issues requires international collaboration and data synthesis. Meta-analyses have served this role in ecology for more than a decade, but the different experimental methodologies researchers use can limit the strength of the meta-analytic approach. Considering the global nature of many environmental issues, a new collaborative approach, which we call coordinated distributed experiments (CDEs), is needed that will control for both spatial and temporal scale, and that encompasses large geographic ranges. Ecological CDEs, involving standardized, controlled protocols, have the potential to advance our understanding of general principles in ecology and environmental science.

Worldwide evidence of a unimodal relationship between productivity and plant species richness

Grassland diversity and ecosystem productivity The relationship between plant species diversity and ecosystem productivity is controversial. The debate concerns whether diversity peaks at intermediate levels of productivity—the so-called humped-back model—or whether there is no clear predictable relationship. Fraser et al. used a large, standardized, and geographically diverse sample of grasslands from six continents to confirm the validity and generality of the humped-back model. Their findings pave the way for a more mechanistic understanding of the factors controlling species diversity. Science, this issue p. 302 The humped-back model of plant species diversity is confirmed by a global grassland survey. The search for predictions of species diversity across environmental gradients has challenged ecologists for decades. The humped-back model (HBM) suggests that plant diversity peaks at intermediate productivity; at low productivity few species can tolerate the environmental stresses, and at high productivity a few highly competitive species dominate. Over time the HBM has become increasingly controversial, and recent studies claim to have refuted it. Here, by using data from coordinated surveys conducted throughout grasslands worldwide and comprising a wide range of site productivities, we provide evidence in support of the HBM pattern at both global and regional extents. The relationships described here provide a foundation for further research into the local, landscape, and historical factors that maintain biodiversity.

Effects of Extreme Weather Events on Plant Productivity and Tissue Die-Back are Modified by Community Composition

Extreme weather events are expected to increase in frequency and magnitude due to climate change. Their effects on vegetation are widely unknown. Here, experimental grassland and heath communities in Central Europe were exposed either to a simulated single drought or to a prolonged heavy rainfall event. The magnitude of manipulations imitated the local 100-year weather extreme according to extreme value statistics. Overall productivity of both plant communities remained stable in the face of drought and heavy rainfall, despite significant effects on tissue die-back. Grassland communities were more resistant against the extreme weather events than heath communities. Furthermore, effects of extreme weather events on community tissue die-back were modified by functional diversity, even though conclusiveness in this part is limited by the fact that only one species composition was available per diversity level within this case study. More diverse grassland communities exhibited less tissue die-back than less complex grassland communities. On the other side, more diverse heath communities were more vulnerable to extreme weather events compared to less complex heath communities. Furthermore, legumes did not effectively contribute to the buffering against extreme weather events in both vegetation types. Tissue die-back proved a strong stress response in plant communities exposed to 100-year extreme weather events, even though one important ecosystem function, namely productivity, remained surprisingly stable in this experiment. Theories and concepts on biodiversity and ecosystem functioning (insurance hypothesis, redundancy hypothesis) may have to be revisited when extreme weather conditions are considered.

Climate change effects on Chikungunya transmission in Europe: geospatial analysis of vector’s climatic suitability and virus’ temperature requirements

BackgroundChikungunya was, from the European perspective, considered to be a travel-related tropical mosquito-borne disease prior to the first European outbreak in Northern Italy in 2007. This was followed by cases of autochthonous transmission reported in South-eastern France in 2010. Both events occurred after the introduction, establishment and expansion of the Chikungunya-competent and highly invasive disease vector Aedes albopictus (Asian tiger mosquito) in Europe. In order to assess whether these outbreaks are indicative of the beginning of a trend or one-off events, there is a need to further examine the factors driving the potential transmission of Chikungunya in Europe. The climatic suitability, both now and in the future, is an essential starting point for such an analysis.MethodsThe climatic suitability for Chikungunya outbreaks was determined by using bioclimatic factors that influence, both vector and, pathogen. Climatic suitability for the European distribution of the vector Aedes albopictus was based upon previous correlative environmental niche models. Climatic risk classes were derived by combining climatic suitability for the vector with known temperature requirements for pathogen transmission, obtained from outbreak regions. In addition, the longest potential intra-annual season for Chikungunya transmission was estimated for regions with expected vector occurrences.In order to analyse spatio-temporal trends for risk exposure and season of transmission in Europe, climate change impacts are projected for three time-frames (2011–2040, 2041–2070 and 2071–2100) and two climate scenarios (A1B and B1) from the Intergovernmental Panel on Climate Change (IPCC). These climatic projections are based on regional climate model COSMO-CLM, which builds on the global model ECHAM5.ResultsEuropean areas with current and future climatic suitability of Chikungunya transmission are identified. An increase in risk is projected for Western Europe (e.g. France and Benelux-States) in the first half of the 21st century and from mid-century onwards for central parts of Europe (e.g. Germany). Interestingly, the southernmost parts of Europe do not generally provide suitable conditions in these projections. Nevertheless, many Mediterranean regions will persist to be climatically suitable for transmission. Overall, the highest risk of transmission by the end of the 21st century was projected for France, Northern Italy and the Pannonian Basin (East-Central Europe). This general tendency is depicted in both, the A1B and B1 climate change scenarios.ConclusionIn order to guide preparedness for further outbreaks, it is crucial to anticipate risk as to identify areas where specific public health measures, such as surveillance and vector control, can be implemented. However, public health practitioners need to be aware that climate is only one factor driving the transmission of vector-borne disease.

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.