Reassessment of the risks of climate change for terrestrial ecosystems.

Forecasting the risks of climate change for species and ecosystems is necessary for developing targeted conservation strategies. Previous risk assessments mapped the exposure of the global land surface to changes in climate. However, this procedure is unlikely to robustly identify priority areas for conservation actions because nonlinear physiological responses and colimitation processes ensure that ecological changes will not map perfectly to the forecast climatic changes. Here, we combine ecophysiological growth models of 135,153 vascular plant species and plant growth-form information to transform ambient and future climatologies into phytoclimates, which describe the ability of climates to support the plant growth forms that characterize terrestrial ecosystems. We forecast that 33% to 68% of the global land surface will experience a significant change in phytoclimate by 2070 under representative concentration pathways RCP 2.6 and RCP 8.5, respectively. Phytoclimates without present-day analogue are forecast to emerge on 0.3-2.2% of the land surface and 0.1-1.3% of currently realized phytoclimates are forecast to disappear. Notably, the geographic pattern of change, disappearance and novelty of phytoclimates differs markedly from the pattern of analogous trends in climates detected by previous studies, thereby defining new priorities for conservation actions and highlighting the limits of using untransformed climate change exposure indices in ecological risk assessments. Our findings suggest that a profound transformation of the biosphere is underway and emphasize the need for a timely adaptation of biodiversity management practices.

Diurnal temperature range as a key predictor of plants' elevation ranges globally.

A prominent hypothesis in ecology is that larger species ranges are found in more variable climates because species develop broader environmental tolerances, predicting a positive range size-temperature variability relationship. However, this overlooks the extreme temperatures that variable climates impose on species, with upper or lower thermal limits more likely to be exceeded. Accordingly, we propose the 'temperature range squeeze' hypothesis, predicting a negative range size-temperature variability relationship. We test these contrasting predictions by relating 88,000 elevation range sizes of vascular plants in 44 mountains to short- and long-term temperature variation. Consistent with our hypothesis, we find that species' range size is negatively correlated with diurnal temperature range. Accurate predictions of short-term temperature variation will become increasingly important for extinction risk assessment in the future.

Differences in mucilage properties and stomatal sensitivity of locally adapted Zea mays in relation with precipitation seasonality and vapour pressure deficit regime of their native environment.

With ongoing climate change and the increase in extreme weather events, especially droughts, the challenge of maintaining food security is becoming ever greater. Locally adapted landraces of crops represent a valuable source of adaptation to stressful environments. In the light of future droughts-both by altered soil water supply and increasing atmospheric water demand (vapor pressure deficit [VPD])-plants need to improve their water efficiency. To do so, plants can enhance their access to soil water by improving rhizosphere hydraulic conductivity via the exudation of mucilage. Furthermore, plants can reduce transpirational water loss via stomatal regulation. Although the role of mucilage and stomata regulation on plant water management have been extensively studied, little is known about a possible coordination between root mucilage properties and stomatal sensitivity as well as abiotic drivers shaping the development of drought resistant trait suits within landraces. Mucilage properties and stomatal sensitivity of eight Mexican landraces of Zea mays in contrast with one inbred line were first quantified under controlled conditions and second related to water demand and supply at their respective site of origin. Mucilage physical properties-namely, viscosity, contact angle, and surface tension-differed between the investigated maize varieties. We found strong influences of precipitation seasonality, thus plant water availability, on mucilage production (R 2 = .88, p < .01) and mucilage viscosity (R 2 = .93, p < .01). Further, stomatal sensitivity to increased atmospheric water demand was related to mucilage viscosity and contact angle, both of which are crucial in determining mucilage's water repellent, thus maladaptive, behavior upon soil drying. The identification of landraces with pre-adapted suitable trait sets with regard to drought resistance is of utmost importance, for example, trait combinations such as exhibited in one of the here investigated landraces. Our results suggest a strong environmental selective force of seasonality in plant water availability on mucilage properties as well as regulatory stomatal effects to avoid mucilage's maladaptive potential upon drying and likely delay critical levels of hydraulic dysfunction. By this, landraces from highly seasonal climates may exhibit beneficial mucilage and stomatal traits to prolong plant functioning under edaphic drought. These findings may help breeders to efficiently screen for local landraces with pre-adaptations to drought to ultimately increase crop yield resistance under future climatic variability.

Constant hydraulic supply and ABA dynamics facilitate the trade-offs in water and carbon.

Carbon-water trade-offs in plants are adjusted through stomatal regulation. Stomatal opening enables carbon uptake and plant growth, whereas plants circumvent drought by closing stomata. The specific effects of leaf position and age on stomatal behavior remain largely unknown, especially under edaphic and atmospheric drought. Here, we compared stomatal conductance (gs ) across the canopy of tomato during soil drying. We measured gas exchange, foliage ABA level and soil-plant hydraulics under increasing vapor pressure deficit (VPD). Our results indicate a strong effect of canopy position on stomatal behavior, especially under hydrated soil conditions and relatively low VPD. In wet soil (soil water potential > -50 kPa), upper canopy leaves had the highest gs (0.727 ± 0.154 mol m-2 s-1) and assimilation rate (A; 23.4 ± 3.9 µmol m-2 s-1) compared to the leaves at a medium height of the canopy (gs : 0.159 ± 0.060 mol m2 s-1; A: 15.9 ± 3.8 µmol m-2 s-1). Under increasing VPD (from 1.8 to 2.6 kPa), gs , A and transpiration were initially impacted by leaf position rather than leaf age. However, under high VPD (2.6 kPa), age effect outweighed position effect. The soil-leaf hydraulic conductance was similar in all leaves. Foliage ABA levels increased with rising VPD in mature leaves at medium height (217.56 ± 85 ng g-1 FW) compared to upper canopy leaves (85.36 ± 34 ng g-1 FW). Under soil drought (< -50 kPa), stomata closed in all leaves resulting in no differences in gs across the canopy. We conclude that constant hydraulic supply and ABA dynamics facilitate preferential stomatal behavior and carbon-water trade-offs across the canopy. These findings are fundamental in understanding variations within the canopy, which helps in engineering future crops, especially in the face of climate change.

Source-specific risk assessment for cadmium in wheat and maize: Towards an enrichment model for China.

Cadmium (Cd) pollution of agricultural soil is of public concern due to its high potential toxicity and mobility. This study aimed to reveal the risk of Cd accumulation in soil and wheat/maize systems, with a specific focus on the source-specific ecological risk, human health risk and Cd enrichment model. For this we investigated more than 6100 paired soil and grain samples with 216 datasets including soil Cd contents, soil pH and grain Cd contents of 85 sites from China. The results showed that mining activities, sewage irrigation, industrial activities and agricultural practices were the critical factors causing Cd accumulation in wheat and maize cultivated sites. Thereinto, mining activities contributed to a higher Cd accumulation risk in the southwest China and Middle Yellow River regions; sewage irrigation influenced the Cd accumulation in the North China Plain. In addition, the investigated sites were classified into different categories by comparing their soil and grain Cd contents with the Chinese soil screening values and food safety values, respectively. Cd enrichment models were developed to predict the Cd levels in wheat and maize grains. The results showed that the models exhibited a good performance for predicting the grain Cd contents among safe and warning sites of wheat (R2 = 0.61 and 0.72, respectively); while the well-fitted model for maize was prone to the overestimated sites (R2 = 0.77). This study will provide national viewpoints for the risk assessments and prediction of Cd accumulation in soil and wheat/maize systems.

Accumulation of potentially toxic elements in Chinese tea (Camellia sinensis): Towards source apportionment and health risk assessment.

Tea (Camellia sinensis) is a popular beverage that is consumed globally. However, a better understanding of potentially toxic elements (PTEs) content in tea leaves and infusion is necessary to minimize risk on human health. Therefore, 249 tea samples (grown in different areas) covering six types of tea were collected in China to investigate the PTEs contents, identify their potential source and assess the health risk associated with drinking tea. PTE contents in tea leaves across six tea types were ND-0.900 (Cd), 0.005-2.133 (As), ND-5.679 (Pb), ND-13.86 (Cr), 1.601-22.93 (Ni), ND-2.048 (Se), 0.109-622.4 (F), 13.02-269.9 (Rb), 1.845-50.88 (Sr), and 2.796-53.23 (Ba) mg/kg. The result of tea infusion showed that 14.3 %-44.1 % (green tea), 14.5 %-46.7 % (black tea), 10.5 %-25.3 % (dark tea), 13.6 %-34.2 % (oolong tea), 16.9 %-40.7 % (yellow tea), and 19.9 %-35.1 % (white tea) of F were released. All tea types, except green tea, exhibited comparatively low leachability of Cd, As, Pb and Cr in tea infusion. The source apportionment revealed that PTEs in tea leaves mainly originated from soil parental materials, while industrial activities, fertilizer application, and manufacturing processes may contribute to exogenous Se, Cd, As, and Cr accumulation. Health risk assessment indicated that F in tea infusion dominated the health risk. Humans may be exposed to a higher health risk by drinking green tea compared to that of other tea types. Nevertheless, the long-term tea consumption is less likely to contribute to pronounced non-carcinogenic and carcinogenic risks. This study confirmed that tea consumption is an important and direct pathway of PTEs uptake in humans. The health risk associated with drinking tea should be of concern.

Chemical properties of key metabolites determine the global distribution of lichens.

In lichen symbioses, fungal secondary metabolites provide UV protection on which lichen algae such as trebouxiophycean green algae-the most prominent group of photobionts in lichen symbioses-sensitively depend. These metabolites differ in their UV absorbance capability and solvability, and thus vary in their propensity of being leached from the lichen body in humid and warm environments, with still unknown implications for the global distribution of lichens. In this study covering more than 10,000 lichenised fungal species, we show that the occurrence of fungal-derived metabolites in combination with their UV absorbance capability and their probability of being leached in warm and humid environments are important eco-evolutionary drivers of global lichen distribution. Fungal-derived UV protection seems to represent an indirect environmental adaptation in which the lichen fungus invests to protect the trebouxiophycean photobiont from high UV radiation in warm and humid climates and, by doing this, secures its carbon source.

The eco-evolutionary significance of rainfall constancy for facultative CAM photosynthesis.

A flexible use of the crassulacean acid metabolism (CAM) has been hypothesised to represent an intermediate stage along a C3 to full CAM evolutionary continuum, when relative contributions of C3 vs CAM metabolism are co-determined by evolutionary history and prevailing environmental constraints. However, evidence for such eco-evolutionary interdependencies is lacking. We studied these interdependencies for the leaf-succulent genus Drosanthemum (Aizoaceae, Southern African Succulent Karoo) by testing for relationships between leaf δ13 C diagnostic for CAM dependence (i.e. contribution of C3 and CAM to net carbon gain), and climatic variables related to temperature and precipitation and their temporal variation. We further quantified the effects of shared phylogenetic ancestry on CAM dependence and its relation to climate. CAM dependence is predicted by rainfall and its temporal variation, with high predictive power of rainfall constancy (temporal entropy). The predictive power of rainfall seasonality and temperature-related variables was negligible. Evolutionary history of the tested clades significantly affected the relationship between rainfall constancy and CAM dependence. We argue that higher CAM dependence might provide an adaptive advantage in increasingly unpredictable rainfall environments when the anatomic exaptation (succulence) is already present. These observations might shed light on the evolution of full CAM.

An operational definition of the biome for global change research.

Biomes are constructs for organising knowledge on the structure and functioning of the world's ecosystems, and serve as useful units for monitoring how the biosphere responds to anthropogenic drivers, including climate change. The current practice of delimiting biomes relies on expert knowledge. Recent studies have questioned the value of such biome maps for comparative ecology and global-change research, partly due to their subjective origin. Here we propose a flexible method for developing biome maps objectively. The method uses range modelling of several thousands of plant species to reveal spatial attractors for different growth-form assemblages that define biomes. The workflow is illustrated using distribution data from 23 500 African plant species. In an example application, we create a biome map for Africa and use the fitted species models to project biome shifts. In a second example, we map gradients of growth-form suitability that can be used to identify sites for comparative ecology. This method provides a flexible framework that (1) allows a range of biome types to be defined according to user needs and (2) enables projections of biome changes that emerge purely from the individualistic responses of plant species to environmental changes.

Data on alpine grassland diversity in Gran Paradiso National Park, Italy.

The diversity of alpine grassland species and their functional traits constitute alpine ecosystem functioning and services that support human-wellbeing. However, alpine grassland diversity is threatened by land use and climate change. Field surveys and monitoring are necessary to understand and preserve such endangered ecosystems. Here we describe data on abundances (percentage cover) of 247 alpine plant species (including mosses and lichens) inside nine 20 m by 20 m plots that were subdivided into 2 m by 2 m subplots. The nine plots are located in Gran Paradiso National Park, Italy. They cover three distinct alpine vegetation subtypes ('pure' natural grassland, sparsely vegetated 'rocky' grassland, and wetland) in each of three valleys (Bardoney, Colle de Nivolet and Levionaz) between 2200 and 2700 m a.s.l., i.e. above the treeline. The vegetation survey was conducted in 2015 at the peak of vegetation development during August. The dataset is provided as supplementary material and associated with the research article "Optimizing sampling effort and information content of biodiversity surveys: a case study of alpine grassland" [1]. See [1] for data interpretation.

The importance of ecological memory for trophic rewilding as an ecosystem restoration approach.

Increasing human pressure on strongly defaunated ecosystems is characteristic of the Anthropocene and calls for proactive restoration approaches that promote self-sustaining, functioning ecosystems. However, the suitability of novel restoration concepts such as trophic rewilding is still under discussion given fragmentary empirical data and limited theory development. Here, we develop a theoretical framework that integrates the concept of 'ecological memory' into trophic rewilding. The ecological memory of an ecosystem is defined as an ecosystem's accumulated abiotic and biotic material and information legacies from past dynamics. By summarising existing knowledge about the ecological effects of megafauna extinction and rewilding across a large range of spatial and temporal scales, we identify two key drivers of ecosystem responses to trophic rewilding: (i) impact potential of (re)introduced megafauna, and (ii) ecological memory characterising the focal ecosystem. The impact potential of (re)introduced megafauna species can be estimated from species properties such as lifetime per capita engineering capacity, population density, home range size and niche overlap with resident species. The importance of ecological memory characterising the focal ecosystem depends on (i) the absolute time since megafauna loss, (ii) the speed of abiotic and biotic turnover, (iii) the strength of species interactions characterising the focal ecosystem, and (iv) the compensatory capacity of surrounding source ecosystems. These properties related to the focal and surrounding ecosystems mediate material and information legacies (its ecological memory) and modulate the net ecosystem impact of (re)introduced megafauna species. We provide practical advice about how to quantify all these properties while highlighting the strong link between ecological memory and historically contingent ecosystem trajectories. With this newly established ecological memory-rewilding framework, we hope to guide future empirical studies that investigate the ecological effects of trophic rewilding and other ecosystem-restoration approaches. The proposed integrated conceptual framework should also assist managers and decision makers to anticipate the possible trajectories of ecosystem dynamics after restoration actions and to weigh plausible alternatives. This will help practitioners to develop adaptive management strategies for trophic rewilding that could facilitate sustainable management of functioning ecosystems in an increasingly human-dominated world.

To replicate, or not to replicate - that is the question: how to tackle nonlinear responses in ecological experiments.

A fundamental challenge in experimental ecology is to capture nonlinearities of ecological responses to interacting environmental drivers. Here, we demonstrate that gradient designs outperform replicated designs for detecting and quantifying nonlinear responses. We report the results of (1) multiple computer simulations and (2) two purpose-designed empirical experiments. The findings consistently revealed that unreplicated sampling at a maximum number of sampling locations maximised prediction success (i.e. the R² to the known truth) irrespective of the amount of stochasticity and the underlying response surfaces, including combinations of two linear, unimodal or saturating drivers. For the two empirical experiments, the same pattern was found, with gradient designs outperforming replicated designs in revealing the response surfaces of underlying drivers. Our findings suggest that a move to gradient designs in ecological experiments could be a major step towards unravelling underlying response patterns to continuous and interacting environmental drivers in a feasible and statistically powerful way.

Ecological and evolutionary legacy of megafauna extinctions.

For hundreds of millions of years, large vertebrates (megafauna) have inhabited most of the ecosystems on our planet. During the late Quaternary, notably during the Late Pleistocene and the early Holocene, Earth experienced a rapid extinction of large, terrestrial vertebrates. While much attention has been paid to understanding the causes of this massive megafauna extinction, less attention has been given to understanding the impacts of loss of megafauna on other organisms with whom they interacted. In this review, we discuss how the loss of megafauna disrupted and reshaped ecological interactions, and explore the ecological consequences of the ongoing decline of large vertebrates. Numerous late Quaternary extinct species of predators, parasites, commensals and mutualistic partners were associated with megafauna and were probably lost due to their strict dependence upon them (co-extinctions). Moreover, many extant species have megafauna-adapted traits that provided evolutionary benefits under past megafauna-rich conditions, but are now of no or limited use (anachronisms). Morphological evolution and behavioural changes allowed some of these species partially to overcome the absence of megafauna. Although the extinction of megafauna led to a number of co-extinction events, several species that likely co-evolved with megafauna established new interactions with humans and their domestic animals. Species that were highly specialized in interactions with megafauna, such as large predators, specialized parasites, and large commensalists (e.g. scavengers, dung beetles), and could not adapt to new hosts or prey were more likely to die out. Partners that were less megafauna dependent persisted because of behavioural plasticity or by shifting their dependency to humans via domestication, facilitation or pathogen spill-over, or through interactions with domestic megafauna. We argue that the ongoing extinction of the extant megafauna in the Anthropocene will catalyse another wave of co-extinctions due to the enormous diversity of key ecological interactions and functional roles provided by the megafauna.

Salt in the wound: The interfering effect of road salt on acidified forest catchments.

Atmospheric acidic depositions have strongly altered the functioning and biodiversity of Central European forest ecosystems. Most impacts occurred until the end of the 20(th) century but the situation substantially improved thereafter caused by legal regulations in the late 1980's to reduce acidifying atmospheric pollution. Since then slow recovery from acidification has been observed in forested catchments and adjacent waters. However, trends of recovery are inconsistent and underlying mechanisms diminishing recovery are still poorly understood. We propose that the input of road salt can significantly affect acidity regime and acidification recovery of forest ecosystems. By comparing the discharge hydro-chemistry and plant community composition of springs fed by forested catchments with and without high levels of salt input over two decades we observed a significant suppression of recovery and elevated levels of nutrient leaching (K(+), Ca(2+) and Mg(2+)) in highly salt contaminated catchments. We show that the pollution of near-surface groundwater (interflow) by road salt application can have lasting effects on ecosystem processes over distances of several hundred metres apart from the salt emitting road.