The plant root economics space in relation to nutrient limitation in Eurasian herbaceous plant communities.

Plant species occupy distinct niches along a nitrogen-to-phosphorus (N:P) gradient, yet there is no general framework for belowground nutrient acquisition traits in relation to N or P limitation. We retrieved several belowground traits from databases, placed them in the "root economics space" framework, and linked these to a dataset of 991 plots in Eurasian herbaceous plant communities, containing plant species composition, aboveground community biomass and tissue N and P concentrations. Our results support that under increasing N:P ratio, belowground nutrient acquisition strategies shift from "fast" to "slow" and from "do-it-yourself" to "outsourcing", with alternative "do-it-yourself" to "outsourcing" strategies at both ends of the spectrum. Species' mycorrhizal capacity patterns conflicted with root economics space predictions based on root diameter, suggesting evolutionary development of alternative strategies under P limitation. Further insight into belowground strategies along nutrient stoichiometry is crucial for understanding the high abundance of threatened plant species under P limitation.

Spatially explicit removal strategies increase the efficiency of invasive plant species control.

Effective management strategies are needed to control expansion of invasive alien plant species and attenuate economic and ecological impacts. While previous theoretical studies have assessed optimal control strategies that balance economic costs and ecological benefits, less attention has been paid to the ways in which the spatial characteristics of individual patches may mediate the effectiveness of management strategies. We developed a spatially explicit cellular automaton model for invasive species spread, and compared the effectiveness of seven control strategies. These control strategies used different criteria to prioritize the removal of invasive species patches from the landscape. The different criteria were related to patch size, patch geometry, and patch position within the landscape. Effectiveness of strategies was assessed for both seed dispersing and clonally expanding plant species. We found that, for seed-dispersing species, removal of small patches and removal of patches that are isolated within the landscape comprised relatively effective control strategies. For clonally expanding species, removal of patches based on their degree of isolation and their geometrical properties comprised relatively effective control strategies. Subsequently, we parameterized the model to mimic the observed spatial distribution of the invasive species Antigonon leptopus on St. Eustatius (northern Caribbean). This species expands clonally and also disperses via seeds, and model simulations showed that removal strategies focusing on smaller patches that are more isolated in the landscape would be most effective and could increase the effectiveness of a 10-yr control strategy by 30-90%, as compared to random removal of patches. Our study emphasizes the potential for invasive plant species management to utilize recent advances in remote sensing, which enable mapping of invasive species at the high spatial resolution needed to quantify patch geometries. The presented results highlight how this spatial information can be used in the design of more effective invasive species control strategies.

Low investment in sexual reproduction threatens plants adapted to phosphorus limitation.

Plant species diversity in Eurasian wetlands and grasslands depends not only on productivity but also on the relative availability of nutrients, particularly of nitrogen and phosphorus. Here we show that the impacts of nitrogen:phosphorus stoichiometry on plant species richness can be explained by selected plant life-history traits, notably by plant investments in growth versus reproduction. In 599 Eurasian sites with herbaceous vegetation we examined the relationship between the local nutrient conditions and community-mean life-history traits. We found that compared with plants in nitrogen-limited communities, plants in phosphorus-limited communities invest little in sexual reproduction (for example, less investment in seed, shorter flowering period, longer lifespan) and have conservative leaf economy traits (that is, a low specific leaf area and a high leaf dry-matter content). Endangered species were more frequent in phosphorus-limited ecosystems and they too invested little in sexual reproduction. The results provide new insight into how plant adaptations to nutrient conditions can drive the distribution of plant species in natural ecosystems and can account for the vulnerability of endangered species.

Plant species occurrence patterns in Eurasian grasslands reflect adaptation to nutrient ratios.

Previous studies of Eurasian grasslands have suggested that nutrient ratios, rather than absolute nutrient availabilities and associated productivity, may be driving plant species richness patterns. However, the underlying assumption that species occupy distinct niches along nutrient ratio gradients remains to be tested. We analysed plant community composition and nutrient status of 644 Eurasian wet grassland plots. The importance of nutrient ratios driving variation in species composition was analysed using ordination methods (DCA and CCA). Subsequently, we assessed the niche position and width along the most important nutrient ratio gradient [N:P] for each species. We found that the N:P ratio explained part of the variation in species composition independent from conventional explanatory variables. The N:P ratio explained less variation than soil moisture or pH, but more than productivity or the availability of N and P separately, highlighting its importance for grassland species composition. Species occupied distinct niches along the N:P gradient, and species' niche widths decreased toward extreme nutrient limitation. After correcting for niche position, there was no overall difference in niche width between endangered and non-endangered species. Surprisingly, endangered species with niche optima at the extreme P-limited end of the gradient had broader niches than their non-endangered counterparts. As species occupied distinct niches along a nutrient ratio gradient, future grassland conservation efforts may benefit from targeting changes in nutrient ratios, i.e. the balance between N and P, rather than only focussing on a general reduction in nutrient availability. However, what management interventions can be used for this purpose remains unclear.

Climate forcing due to optimization of maximal leaf conductance in subtropical vegetation under rising CO2.

Plant physiological adaptation to the global rise in atmospheric CO(2) concentration (CO(2)) is identified as a crucial climatic forcing. To optimize functioning under rising CO(2), plants reduce the diffusive stomatal conductance of their leaves (g(s)) dynamically by closing stomata and structurally by growing leaves with altered stomatal densities and pore sizes. The structural adaptations reduce maximal stomatal conductance (g(smax)) and constrain the dynamic responses of g(s). Here, we develop and validate models that simulate structural stomatal adaptations based on diffusion of CO(2) and water vapor through stomata, photosynthesis, and optimization of carbon gain under the constraint of a plant physiological cost of water loss. We propose that the ongoing optimization of g(smax) is eventually limited by species-specific limits to phenotypic plasticity. Our model reproduces observed structural stomatal adaptations and predicts that adaptation will continue beyond double CO(2). Owing to their distinct stomatal dimensions, angiosperms reach their phenotypic response limits on average at 740 ppm and conifers on average at 1,250 ppm CO(2). Further, our simulations predict that doubling today's CO(2) will decrease the annual transpiration flux of subtropical vegetation in Florida by ≈60 W·m(-2). We conclude that plant adaptation to rising CO(2) is altering the freshwater cycle and climate and will continue to do so throughout this century.

Microbial community composition in the dung of five sympatric European herbivore species.

The dung microbiome is a complex system that is highly influenced by species and diet. This study characterized the dung bacterial and fungal communities of five herbivore species inhabiting the National Park Zuid-Kennemerland, the Netherlands. The five selected herbivore species were rabbit (Oryctolagus cuniculus L.), cow (Bos taurus L.), horse (Equus ferus caballus L.), fallow deer (Dama dama L.), and European bison (Bison bonasus L.). We explored the effects of distinct digestive physiology (ruminants vs. non-ruminants) and diverse dietary preferences on the microbial community composition of herbivore dung. Firmicutes and Bacteroidetes were dominant bacterial phyla in the dung of all five herbivore species, and Ascomycota was the predominant fungal phylum. Verrucomicrobiota and Mucoromycota were more present in horse dung and Proteobacteria were more abundant in rabbit dung than the three ruminant dung types. There were few significant differences in the microbial community structure among the three ruminant dung types. The alpha and beta diversity of dung microbial communities significantly differed between ruminants and non-ruminants, especially in bacterial communities. Based on MetaCyc pathways, we found that the primary functions of bacteria in herbivore dung were focused on biosynthesis, various super pathways, and degradation, with a few differences between ruminant and non-ruminant dung. FUNGuild analysis showed that horse dung had more saprotrophic fungi, while the fungi in fallow deer dung had more symbiotrophic properties, with the fungal functions of bison, cow, and rabbit dung somewhere in between. There was also a correlation between microbial community and nutrient composition of the substrate in herbivore dung. Understanding the dung microbial community composition of these herbivore species can enrich the database of mammalian gut microbiomes for studying the mechanisms of microbial community variation while preparing for exploring a new perspective to study the impact of herbivores on ecosystems through dung deposition.

A global analysis of how human infrastructure squeezes sandy coasts.

Coastal ecosystems provide vital services, but human disturbance causes massive losses. Remaining ecosystems are squeezed between rising seas and human infrastructure development. While shoreline retreat is intensively studied, coastal congestion through infrastructure remains unquantified. Here we analyse 235,469 transects worldwide to show that infrastructure occurs at a median distance of 392 meter from sandy shorelines. Moreover, we find that 33% of sandy shores harbour less than 100 m of infrastructure-free space, and that 23-30% of this space may be lost by 2100 due to rising sea levels. Further analyses show that population density and gross domestic product explain 35-39% of observed squeeze variation, emphasizing the intensifying pressure imposed as countries develop and populations grow. Encouragingly, we find that nature reserves relieve squeezing by 4-7 times. Yet, at present only 16% of world's sandy shores have a protected status. We therefore advocate the incorporation of nature protection into spatial planning policies.

Wetscapes: Restoring and maintaining peatland landscapes for sustainable futures.

Peatlands are among the world's most carbon-dense ecosystems and hotspots of carbon storage. Although peatland drainage causes strong carbon emissions, land subsidence, fires and biodiversity loss, drainage-based agriculture and forestry on peatland is still expanding on a global scale. To maintain and restore their vital carbon sequestration and storage function and to reach the goals of the Paris Agreement, rewetting and restoration of all drained and degraded peatlands is urgently required. However, socio-economic conditions and hydrological constraints hitherto prevent rewetting and restoration on large scale, which calls for rethinking landscape use. We here argue that creating integrated wetscapes (wet peatland landscapes), including nature preserve cores, buffer zones and paludiculture areas (for wet productive land use), will enable sustainable and complementary land-use functions on the landscape level. As such, transforming landscapes into wetscapes presents an inevitable, novel, ecologically and socio-economically sound alternative for drainage-based peatland use.

Endangered plants persist under phosphorus limitation.

Nitrogen enrichment is widely thought to be responsible for the loss of plant species from temperate terrestrial ecosystems. This view is based on field surveys and controlled experiments showing that species richness correlates negatively with high productivity and nitrogen enrichment. However, as the type of nutrient limitation has never been examined on a large geographical scale the causality of these relationships is uncertain. We investigated species richness in herbaceous terrestrial ecosystems, sampled along a transect through temperate Eurasia that represented a gradient of declining levels of atmospheric nitrogen deposition--from approximately 50 kg ha(-1) yr(-1) in western Europe to natural background values of less than 5 kg ha(-1) yr(-1) in Siberia. Here we show that many more endangered plant species persist under phosphorus-limited than under nitrogen-limited conditions, and we conclude that enhanced phosphorus is more likely to be the cause of species loss than nitrogen enrichment. Our results highlight the need for a better understanding of the mechanisms of phosphorus enrichment, and for a stronger focus on conservation management to reduce phosphorus availability.

Phosphorus fertilization is eradicating the niche of northern Eurasia's threatened plant species.

The greater bioavailability of nitrogen (N), phosphorus (P) and potassium (K) in the Anthropocene has strongly impacted terrestrial plant communities. In northwest Europe, because high N deposition is considered the main driver of plant diversity loss, European Union (EU) legislation to reduce N deposition is expected to promote plant species recovery. However, this expectation is simplistic: it ignores the role of other macronutrients. Analysing the relationship between plant species pools and species stoichiometric niches along nutrient gradients across northern Eurasia's herbaceous ecosystems, we found that both absolute and relative P availability are more critical than N or K availability. This result is consistent with stoichiometric niche theory, and with findings from studies of hyperdiverse forests and shrublands at lower latitudes. We show that ecosystems with low absolute and relative P availability harbour a unique set of threatened species that have narrower nutrient-based niche widths than non-threatened species. Such ecosystems represent a conservation priority, but may be further threatened by latent effects of relative P enrichment arising from reduction of N availability without simultaneous reduction of P. The narrow focus of EU legislation on reducing N, but not P, may therefore inadvertently increase the threat to many of Europe's already threatened plant species. An EU Phosphate Directive is needed.

Source and sink activity of Holcus lanatus in response to absolute and relative supply of nitrogen and phosphorus.

Mineral nutrients influence photosynthesis and tissue formation; a shift from nitrogen (N)-limited to phosphorus (P)-limited growth induced by high N deposition may change plant growth in terms of physiology and morphology. This experiment showed that absolute and relative N and P supply affected net photosynthesis (source activity) and biomass formation (sink activity), and the relationship between source and sink activities of Holcus lanatus L. under various nutrient treatments. H. lanatus was grown at three N:P ratios (5, 15, 45) with two absolute supply levels of N and P. Between N:P 5 at low level and N:P 45 at high level, and between N:P 45 at low level and N:P 5 at high level, there was a nine-fold difference in N and P supply. Maximum light-saturated net photosynthesis rate (Amax), specific leaf area (SLA), leaf area, and shoot and root biomass were determined during and after the growth process. Amax was minimal at N:P 5 and increased only with increasing absolute N supply. Neither SLA nor leaf area were affected by N:P; increasing absolute P supply significantly increased leaf area. Shoot and root biomass were minimal at N:P 45 and increased dramatically with increasing absolute P supply. Plant biomass was not correlated with Amax. Our results highlight that H. lanatus growth is predominantly controlled by P supply and to a lesser extent by N, whereas net photosynthesis exerted no apparent control on growth under these sink-limited growth conditions. Our findings contribute to understanding of plant growth under sink-limited conditions.

Resource contrast in patterned peatlands increases along a climatic gradient.

Spatial patterning of ecosystems can be explained by several mechanisms. One approach to disentangling the influence of these mechanisms is to study a patterned ecosystem along a gradient of environmental conditions. This study focused on hummock-hollow patterning of peatlands. Previous models predicted that patterning in drainage-dominated peatlands is driven by a peat-accumulation mechanism, reflected by higher nutrient availability in hollows relative to hummocks. Alternatively, patterning in evapotranspiration (ET)-dominated peatlands may be driven by a nutrient-accumulation mechanism, reflected by reversed nutrient distribution, namely, higher nutrient availability in hummocks relative to hollows. Here, we tested these predictions by comparing nutrient distributions among patterned peatlands in maritime (Scotland), humid temperate (Sweden), and humid continental (Siberia) climates. The areas comprise a climatic gradient from very wet and drainage-dominated (Scotland) to less wet and ET-dominated (Siberia) peatlands. Nutrient distribution was quantified as resource contrast, a measure for hummock-hollow difference in nutrient availability. We tested the hypothesis that the climatic gradient shows a trend in the resource contrast; from negative (highest nutrient availability in hollows) in Scotland to positive (highest nutrient availability in hummocks) in Siberia. The resource contrasts as measured in vegetation indeed showed a trend along the climatic gradient: contrasts were negative to slightly positive in Scotland, positive in Sweden, and strongly positive in Siberia. This finding corroborates the main prediction of previous models. Our results, however, also provided indications for further model development. The low concentrations of nutrients in the water suggest that existing models could be improved by considering both the dissolved and adsorbed phase and explicit inclusion of both nutrient-uptake and nutrient-storage processes. Our study suggests that future climate change may affect the ecosystem functioning of patterned peatlands by altering the contribution of pattern-forming mechanisms to redistribution of water and nutrients within these systems.

The impact of climate change on flow conditions and wetland ecosystems in the Lower Biebrza River (Poland).

Water plays a key role in the functioning of wetlands and a shortage or contamination of it leads to changes in habitat conditions and degradation of ecosystems. This article scrutinizes the impact of climate change on the hydrological characteristics of floods (maximum flow, duration, volume) in the River Biebrza wetlands (North-East Poland). We analysed the trends in duration and volume of flood and maximum discharges in the historical period 1970-2000 and predicted these for the future periods 2020-2050 and 2070-2100, respectively. Next we assessed the impact on the wetland ecosystems. The basis of our assessments consists of statistical analyses of hydrographs and calculations by the Soil and Water Assessment Tool hydrological model and considering nine bias-corrected climate models. The results indicate that both volume and duration of winter floods will keep increasing continuously under Representative Concentration Pathways 4.5 and 8.5. The reduction in peak annual floods is expected to decline slightly in both scenarios. On the other hand, the analysis of trends in mean and standard deviation revealed negligible tendencies in the datasets for summer and winter hydrological seasons within the three time frames analysed (1970-2000; 2020-2050; 2070-2100). We foresee several future implications for the floodplain ecosystems. Shifts in transversal ecosystem zonation parallel to the river will likely take place with more highly productive flood tolerant vegetation types. Nutrient availability and algal blooms during spring inundations will likely increase. Slowdown of organic matter turnover later in summer will lead to a higher peat accumulation rate. Logistical problems with summer mowing and removal of bushes in winter may enhance shrub encroachment.

Can sand nourishment material affect dune vegetation through nutrient addition?

In the Netherlands it is common to nourish the coastline with sand from the seabed. Foredunes are replenished with sand from the beach and can be transported further into the dune area. We investigated whether nourishment material alters the phosphorus (P) content of dune soil and the nitrogen (N):P ratio of dune vegetation in two areas: a mega sand nourishment with fixed foredunes (SE) and a traditional sand nourishment with dynamic foredunes (NWC). Four zones were considered: beach (zone 1), frontal foredunes (zone 2), foredunes crest (zone 3) and inner dunes (zone 4). We estimated the characteristics of fine (< 250-µm) and coarse (250-2000 µm) sand. Total P, P speciation and available P of SE and NWC were similar until zone 4. Zone 1-3 consisted mainly of coarse sand, whereas the sand in zone 4 was finer with higher amounts at NWC. Iron (Fe) bound P was comparable for fine and coarse sand in zone 1-3, but high contents were present in zone 4. In zone 1-3, calcium (Ca) bound P was mainly found in the fine fraction, which was abundant in the coarse fraction of zone 4. After a period of 4 years, the effect of dynamic dunes on P fractions and dune plant species was not apparent yet, although inblowing sand mainly consisted of fine sand with high contents of Ca-bound P. This may change over time, especially in dynamic dunes with higher eolian activity of fine sand. Consequently, pH buffering of the soil may increase because of a higher Ca­carbonate content, which leads to decreased solubility of Ca-bound P and low P availability for the vegetation. Both low P availability and high buffering capacity are known environmental factors that facilitate endangered dune plant species.

Nutrients and hydrology indicate the driving mechanisms of peatland surface patterning.

Peatland surface patterning motivates studies that identify underlying structuring mechanisms. Theoretical studies so far suggest that different mechanisms may drive similar types of patterning. The long time span associated with peatland surface pattern formation, however, limits possibilities for empirically testing model predictions by field manipulations. Here, we present a model that describes spatial interactions between vegetation, nutrients, hydrology, and peat. We used this model to study pattern formation as driven by three different mechanisms: peat accumulation, water ponding, and nutrient accumulation. By on-and-off switching of each mechanism, we created a full-factorial design to see how these mechanisms affected surface patterning (pattern of vegetation and peat height) and underlying patterns in nutrients and hydrology. Results revealed that different combinations of structuring mechanisms lead to similar types of peatland surface patterning but contrasting underlying patterns in nutrients and hydrology. These contrasting underlying patterns suggest that the presence or absence of the structuring mechanisms can be identified by relatively simple short-term field measurements of nutrients and hydrology, meaning that longer-term field manipulations can be circumvented. Therefore, this study provides promising avenues for future empirical studies on peatland patterning.

How nitrogen and sulphur addition, and a single drought event affect root phosphatase activity in Phalaris arundinacea.

Conservation and restoration of fens and fen meadows often aim to reduce soil nutrients, mainly nitrogen (N) and phosphorus (P). The biogeochemistry of P has received much attention as P-enrichment is expected to negatively impact on species diversity in wetlands. It is known that N, sulphur (S) and hydrological conditions affect the biogeochemistry of P, yet their interactive effects on P-dynamics are largely unknown. Additionally, in Europe, climate change has been predicted to lead to increases in summer drought. We performed a greenhouse experiment to elucidate the interactive effects of N, S and a single drought event on the P-availability for Phalaris arundinacea. Additionally, the response of plant phosphatase activity to these factors was measured over the two year experimental period. In contrast to results from earlier experiments, our treatments hardly affected soil P-availability. This may be explained by the higher pH in our soils, hampering the formation of Fe-P or Fe-Al complexes. Addition of S, however, decreased the plants N:P ratio, indicating an effect of S on the N:P stoichiometry and an effect on the plant's P-demand. Phosphatase activity increased significantly after addition of S, but was not affected by the addition of N or a single drought event. Root phosphatase activity was also positively related to plant tissue N and P concentrations, plant N and P uptake, and plant aboveground biomass, suggesting that the phosphatase enzyme influences P-biogeochemistry. Our results demonstrated that it is difficult to predict the effects of wetland restoration, since the involved mechanisms are not fully understood. Short-term and long-term effects on root phosphatase activity may differ considerably. Additionally, the addition of S can lead to unexpected effects on the biogeochemistry of P. Our results showed that natural resource managers should be careful when restoring degraded fens or preventing desiccation of fen ecosystems.

Herbivore dung quality affects plant community diversity.

Nutrient availability is important for plant community composition and diversity, but most studies focus on inorganic nutrients. Far less is known about the impact of nutrients in organic forms such as herbivore dung. Here we show that dung of 11 European herbivore species varies widely in nitrogen (N) and phosphorus (P) concentrations, as well as in C:N:P ratios. We demonstrate that variation in dung quality of five herbivore species influences the diversity and composition of a mesocosm plant community. The impact of dung quality was at least as strong as, or stronger than, the effect of manipulating the quantity of dung by a factor six. Our study supports the hypothesis that both nutrient quantity and nutrient imbalances are important controlling factors for plant species diversity, and stresses the important role of herbivores on plant communities, not only via selective foraging, but also via stoichiometric variation of nutrients in their dung.

Ecotoxicological risk of trace element mobility in coastal semiartificial depositional areas near the mouth of the river Rhine, the Netherlands.

Artificial sand replenishments are globally used as innovative coastal protection measures. In these replenishments elevated porewater concentrations of trace elements are found. The present study investigated possible ecotoxicological risks at 2 intertidal depositional sites, the Sand Engine as a recent innovative Dutch coastal management project and a semiartificial tidal flat. Using the sediment quality triad approach, we considered 3 major lines of evidence: geochemical characterization, toxicity characterization using bioassays with the estuarine amphipod Corophium volutator, and ecological field survey. In both depositional areas C. volutator is at risk: moderate (Sand Engine) and low (tidal flat). For tidal flat, the bioavailability of trace elements differs between the field site and the laboratory. Contamination from arsenic and copper is present, but the low survival rate of C. volutator from the bioassay suggests the presence of additional contaminations. The highly morphological dynamic environment of Sand Engine creates a less favorable habitat for C. volutator, where local spots with stagnant water can temporarily create hypoxic conditions and sulfate becomes reduced. The dynamic system mobilizes especially arsenic, triggering adverse ecotoxic effects at low original sediment concentrations. To conclude, the sediment quality triad approach shows that a semiartificial tidal flat is preferred over a highly dynamic coastal management project like the Sand Engine. The Sand Engine concept does not provide suitable conditions for macrobenthos species like C. volutator; therefore, limiting the nature development goal set together with the coastal protection goal. Assessing each line of evidence from the approach together with additional measurements established more precise and realistic conclusions, showing that evaluating the contributions of this method is necessary to understand the causes of risk in a site-specific manner. Environ Toxicol Chem 2018;37:2933-2946. © 2018 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals, Inc. on behalf of SETAC.