David J. Gowing

Nitrogen deposition threatens species richness of grasslands across Europe.

Evidence from an international survey in the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is reducing plant species richness in acid grasslands. Across the deposition gradient in this region (2-44 kg N ha(-1) yr(-1)) species richness showed a curvilinear response, with greatest reductions in species richness when deposition increased from low levels. This has important implications for conservation policies, suggesting that to protect the most sensitive grasslands resources should be focussed where deposition is currently low. Soil pH is also an important driver of species richness indicating that the acidifying effect of nitrogen deposition may be contributing to species richness reductions. The results of this survey suggest that the impacts of nitrogen deposition can be observed over a large geographical range.

Phylogeny and the hierarchical organization of plant diversity

R. H. Whittakers idea that plant diversity can be divided into a hierarchy of spatial components from alpha at the within-habitat scale through beta for the turnover of species between habitats to gamma along regional gradients implies the underlying existence of alpha, beta, and gamma niches. We explore the hypothesis that the evolution of alpha, beta, and gamma niches is also hierarchical, with traits that define the alpha niche being labile, while those defining beta and gamma niches are conservative. At the alpha level we find support for the hypothesis in the lack of close significant phylogenetic relationship between meadow species that have similar alpha niches. In a second test, alpha niche overlap based on a variety of traits is compared between congeners and noncongeners in several communities; here, too, there is no evidence of a correlation between alpha niche and phylogeny. To test whether beta and gamma niches evolve conservatively, we reconstructed the evolution of relevant traits on evolutionary trees for 14 different clades. Tests against null models revealed a number of instances, including some in island radiations, in which habitat (beta niche) and elevational maximum (an aspect of the gamma niche) showed evolutionary conservatism.

Absence of phylogenetic signal in the niche structure of meadow plant communities

A significant proportion of the global diversity of flowering plants has evolved in recent geological time, probably through adaptive radiation into new niches. However, rapid evolution is at odds with recent research which has suggested that plant ecological traits, including the β- (or habitat) niche, evolve only slowly. We have quantified traits that determine within-habitat α diversity (α niches) in two communities in which species segregate on hydrological gradients. Molecular phylogenetic analysis of these data shows practically no evidence of a correlation between the ecological and evolutionary distances separating species, indicating that hydrological α niches are evolutionarily labile. We propose that contrasting patterns of evolutionary conservatism for α- and β-niches is a general phenomenon necessitated by the hierarchical filtering of species during community assembly. This determines that species must have similar β niches in order to occupy the same habitat, but different α niches in order to coexist.

A fundamental, eco‐hydrological basis for niche segregation in plant communities

• Ecologists still puzzle over how plant species manage to coexist with one another while competing for the same essential resources. The classic answer for animal communities is that species occupy different niches, but how plants do this is more difficult to determine. We previously found niche segregation along fine-scale hydrological gradients in European wet meadows and proposed that the mechanism might be a general one, especially in communities that experience seasonal saturation. • We quantified the hydrological niches of 96 species from eight fynbos communities in the biodiversity hotspot of the Cape Floristic Region, South Africa and 99 species from 18 lowland wet meadow communities in the UK. Niche overlap was computed for all combinations of species. • Despite the extreme functional and phylogenetic differences between the fynbos and wet meadow communities, an identical trade-off (i.e. specialization of species towards tolerance of aeration and/or drying stress) was found to cause segregation along fine-scale hydrological gradients. • This study not only confirms the predicted generality of hydrological niche segregation, but also emphasizes its importance for structuring plant communities. Eco-hydrological niche segregation will have implications for conservation in habitats that face changing hydrology caused by water abstraction and climate change.

Hydrological niches in terrestrial plant communities: a review

1. Despite the fundamental significance of water to plants and the persisting question of how competing species coexist, this is the first review of hydrological niches. We define hydrological niche segregation (HNS) as: (i) partitioning of space on fine-scale soil-moisture gradients, (ii) partitioning of water as a resource, and/or (iii) partitioning of recruitment opportunities among years caused by species specializing on particular patterns of temporal variance of water supply (the storage effect). 2. We propose that there are three types of constraint that lead to the trade-offs that underlie HNS. (i) An edaphic constraint creates a trade-off between the supply to roots of O2 on the one hand vs. water and nutrients on the other. (ii) A biophysical constraint governs gas exchange by leaves, leading to a trade-off between CO2 acquisition vs. water loss. (iii) A structural constraint arising from the physics of water-conducting tissues leads to a safety vs. efficiency trade-off. 3. Significant HNS was found in 43 of 48 field studies across vegetation types ranging from arid to wet, though its role in coexistence remains to be proven in most cases. Temporal partitioning promotes coexistence through the storage effect in arid plant communities, but has yet to be shown elsewhere. In only a few cases is it possible to unequivocally link HNS to a particular trade-off. 4. Synthesis. The field and experimental evidence make it clear that HNS is widespread, though it is less clear what its precise mechanisms or consequences are. HNS mechanisms should be revealed by further study of the constraints and trade-offs that govern how plants obtain and use water and HNS can be mechanistically linked to its consequences with appropriate community models. In a changing climate, such an integrated programme would pay dividends for global change research.

Impact of nitrogen deposition at the species level

In Europe and, increasingly, the rest of the world, the key policy tool for the control of air pollution is the critical load, a level of pollution below which there are no known significant harmful effects on the environment. Critical loads are used to map sensitive regions and habitats, permit individual polluting activities, and frame international negotiations on transboundary air pollution. Despite their fundamental importance in environmental science and policy, there has been no systematic attempt to verify a critical load with field survey data. Here, we use a large dataset of European grasslands along a gradient of nitrogen (N) deposition to show statistically significant declines in the abundance of species from the lowest level of N deposition at which it is possible to identify a change. Approximately 60% of species change points occur at or below the range of the currently established critical load. If this result is found more widely, the underlying principle of no harm in pollution policy may need to be modified to one of informed decisions on how much harm is acceptable. Our results highlight the importance of protecting currently unpolluted areas from new pollution sources, because we cannot rule out ecological impacts from even relatively small increases in reactive N deposition.

Regional trends in soil acidification and exchangeable metal concentrations in relation to acid deposition rates

The deposition of high levels of reactive nitrogen (N) and sulphur (S), or the legacy of that deposition, remain among the worlds most important environmental problems. Although regional impacts of acid deposition in aquatic ecosystems have been well documented, quantitative evidence of wide-scale impacts on terrestrial ecosystems is not common. In this study we analysed surface and subsoil chemistry of 68 acid grassland sites across the UK along a gradient of acid deposition, and statistically related the concentrations of exchangeable soil metals (1 M KCl extraction) to a range of potential drivers. The deposition of N, S or acid deposition was the primary correlate for 8 of 13 exchangeable metals measured in the topsoil and 5 of 14 exchangeable metals in the subsoil. In particular, exchangeable aluminium and lead both show increased levels above a soil pH threshold of about 4.5, strongly related to the deposition flux of acid compounds.

Does Soil Strength Play a Role in Wheat Yield Losses Caused by Soil Drying

Shoot growth in wheat is sensitive to high soil strength, but as high strength and drying tend to occur together it has proved difficult to separate the effects of water stress and mechanical impedance. The results of two field experiments in 2003 and 2004, where soil strength was manipulated by compaction and irrigation, demonstrated that the yield of wheat (Triticum aestivum L.) was sensitive to physical stress in the root zone. We obtained linear relationships between yield and soil strength and between yield and accumulated soil moisture data (accumulation analogous to thermal time), with similar slopes for both seasons. We were unable to detect root-sourced signals of xylem-sap ABA concentration, despite changes in stomatal conductance. When mechanical impedance and matric potential were varied independently in controlled environments, the growth of wheat was sensitive to mechanical impedance, but not to small changes in matric potential. While the response of stomatal conductance to soil drying in the field could be interpreted as evidence of hydraulic signalling, we suggest that the role of high soil strength, in limiting growth rates on moderately dry soil, requires further research.

The impact of nitrogen deposition on acid grasslands in the Atlantic region of Europe

A survey of 153 acid grasslands from the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is changing plant species composition and soil and plant-tissue chemistry. Across the deposition gradient (2-44 kg N ha(-1) yr(-1)) grass richness as a proportion of total species richness increased whereas forb richness decreased. Soil C:N ratio increased, but soil extractable nitrate and ammonium concentrations did not show any relationship with nitrogen deposition. The above-ground tissue nitrogen contents of three plant species were examined: Agrostis capillaris (grass), Galium saxatile (forb) and Rhytidiadelphus squarrosus (bryophyte). The tissue nitrogen content of neither vascular plant species showed any relationship with nitrogen deposition, but there was a weak positive relationship between R. squarrosus nitrogen content and nitrogen deposition. None of the species showed strong relationships between above-ground tissue N:P or C:N and nitrogen deposition, indicating that they are not good indicators of deposition rate.

Soil phosphorus constrains biodiversity across European grasslands

Nutrient pollution presents a serious threat to biodiversity conservation. In terrestrial ecosystems, the deleterious effects of nitrogen pollution are increasingly understood and several mitigating environmental policies have been developed. Compared to nitrogen, the effects of increased phosphorus have received far less attention, although some studies have indicated that phosphorus pollution may be detrimental for biodiversity as well. On the basis of a dataset covering 501 grassland plots throughout Europe, we demonstrate that, independent of the level of atmospheric nitrogen deposition and soil acidity, plant species richness was consistently negatively related to soil phosphorus. We also identified thresholds in soil phosphorus above which biodiversity appears to remain at a constant low level. Our results indicate that nutrient management policies biased toward reducing nitrogen pollution will fail to preserve biodiversity. As soil phosphorus is known to be extremely persistent and we found no evidence for a critical threshold below which no environmental harm is expected, we suggest that agro-environmental schemes should include grasslands that are permanently free from phosphorus fertilization.