Cecilia Dupré

Species pool: the concept, its determination and significance for community restoration

. The concept of species pool is reviewed. It is suggested to maintain the terms regional pool and local pool but replace actual pool by community pool. The regional and local pool are considered as selections from the regional and local flora based on ecological similarity. It is also suggested to include in the community pool a selection of species present only as diaspores in the diaspore bank (including diaspores from the seed rain), the selection being based on the same ecological criteria. Four approaches to determine the species pool are discussed: ecological, functional and phytosociological similarity, and an experimental approach. The phytosociological approach appears to be promising. The species pool is elaborated as a fuzzy set in the sense that each species of a community or a local or regional flora is a member of any community, local or regional species pool with different degrees of membership. This membership is defined as a probability of a species to become part of the community pool of a target community which is a function of the ecological (environmen-tal/functional/phytosociological) similarity of the species with the target community; the shortness of the distance of its nearest populations, the frequency/abundance, the dispersal capacity, the actual presence of dispersal mechanisms, the germinability of newly arrived diaspores, and the longevity of seeds (viability) in the diaspore bank. The information on species pools is needed for designing experiments where the number of species in a community is to be manipulated, for instance in restoration management.

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.

Acidification and eutrophication of deciduous forests in northwestern Germany demonstrated by indicator species analysis

A new method of indicator species analysis is suggested that can be used to demonstrate changes in the species composition of forests resulting from atmospheric deposition of acidifying substances and nitrogen. The method is not dependent on an exact re-localization of sites. 2162 published releves of Fagus sylvatica and Quercus robur- Carpinus betulus forests from two study areas in Germany (S Niedersachsen and Nordrhein-Westfalen) were compiled and divided into releves from before 1975 and more recent releves. Weighted averages of Ellenberg indicator values for light (L), moisture (M), reaction (R), and nitrogen (N) were calculated and for each of the four data sets - mesic and dry sites in two areas - N was regressed on R to give a predicted value (Npred) for any given R. The differences between observed values Nobs and Npred were calculated separately for old and new releves (Ndev-old and Ndev-new). The same analysis was applied to model data including 16 different scenarios of acidification and/or eutrophication. The model runs resulted in six different types of regression plots for Ndev and R. All models had in common that the regression lines for Ndev-new lay, completely or largely, above the zero line of no change and above the lines for Ndev-old. Also, in the four data sets, Ndev-new was generally larger than Ndev-old, and particularly so in Nordrhein-Westfalen. The values of Ldev-old and Ldev-new indicate that the differences in Ndev are unlikely to be caused by altered forest management, but must result from atmospheric pollution.

Prediction of occurrence of vascular plants in deciduous forests of South Sweden by means of Ellenberg indicator values

. In this study we present a new method for predicting the occurrences of species using data from deciduous forests in South Sweden. Complete species lists of vascular plants were compiled from 101 stands and from representative sample plots inside the stands. Soil samples from each stand were collected for determination of pH and nitrogen mineralization. Presence-absence data for species were fitted to the values of four environmental variables - soil moisture, soil reaction (pH), soil nitrogen and light - by means of Linear (Multiple) Logistic Regression (LLR), and Gaussian (Multiple) Logistic Regression (GLR). First, these values were estimated by calculating the weighted averages of Ellenberg indicator values. Second, the estimates for reaction and nitrogen were substituted by the real measurements of pH and mineralized NH4+, keeping the Ellenberg estimates for light and moisture. The models were validated by an independent test data set. In general, the models had high predictive abilities. GLR fitted the species occurrences better to the environmental variables than LLR, but had a lower accuracy of prediction of species occurrence in the stands. The use of soil measurements instead of Ellenberg indicator values did not improve the predictive abilities of the models. The environmental conditions in the stand test set were successfully estimated by using species data from the plots. When using the species lists of the stands instead of plot data, a slightly better predictive ability was obtained. The collection of plot data, however, is easier and less time-consuming. The accuracy of prediction differed considerably between species.

Species richness in deciduous forests: Effects of species pools and environmental variables

Abstract The study was conducted in deciduous forests of two Swedish regions, Öland and Uppland. It had two objectives: to (1) test the species pool hypothesis by examining if differences in small-scale species richness are related to differences in large-scale species richness and the size of the regional species pool, and (2) to examine the relationship between species richness and productivity and its scale-dependence. The first data set comprised 36 sites of moderate to high productivity. In each site, we recorded the presence of vascular plant species in nested plots ranging from 0.001 to 1000 m2 and measured several environmental variables. Soil pH and Ellenberg site indicator scores for nitrogen were used as estimators of productivity. The second data set included 24 transects (each with 20 1-m2 plots) on Öland in sites with low to high productivity. Species number, soil pH and relative light intensity were determined in each plot. The forest sites on Öland were more species-rich than the Uppland sites on all spatial scales, although environmental conditions were similar. Small-scale and large-scale species richness were positively correlated. The results present evidence in favour of the species pool hypothesis. In the nested-plots data set, species number was negatively correlated with pH and nitrogen indicator scores, whereas a unimodal relationship between species number and pH was found for the transect data set. These results, as well as previously published data, support the hump-shaped relationship between species richness and productivity in Swedish deciduous forests. Two explanations for the higher species richness of the sites with moderate productivity are given: first, these sites have a higher environmental heterogeneity and second, they have a larger ‘habitat-specific’ species pool. Nomenclature: Karlsson (1997).

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.

Addressing the Impact of Atmospheric Nitrogen Deposition on Western European Grasslands.

There is a growing evidence base demonstrating that atmospheric nitrogen deposition presents a threat to biodiversity and ecosystem function in acid grasslands in Western Europe. Here, we report the findings of a workshop held for European policy makers to assess the perceived importance of reactive nitrogen deposition for grassland conservation, identify areas for policy development in Europe and assess the potential for managing and mitigating the impacts of nitrogen deposition. The importance of nitrogen as a pollutant is already recognized in European legislation, but there is little emphasis in policy on the evaluation of changes in biodiversity due to nitrogen. We assess the potential value of using typical species, as defined in the European Union Habitats Directive, for determining the impact of nitrogen deposition on acid grasslands. Although some species could potentially be used as indicators of nitrogen deposition, many of the typical species do not respond strongly to nitrogen deposition and are unlikely to be useful for identifying impact on an individual site. We also discuss potential mitigation measures and novel ways in which emissions from agriculture could be reduced.

Forest vascular plants as indicators of plant species richness: A data analysis of a flora atlas from northwestern Germany

Abstract Our study had the objective to examine whether the number of forest vascular plants in a forest-poor region may be indicative of total plant species richness and of the number of threatened plant species. We also related forest plant species richness to geological and soil variables. The analysis was based on a regional flora atlas from the Weser-Elbe region in northwestern Germany including incidence data of species in a total of 1109 grid cells (each ca. 2.8 × 2.8 km2). All taxa were classified either as forest or non-forest species. Total species richness in the grid cells ranged from 65 to 597, with a mean value of 308. The number of forest species varied between 20 and 309 (mean 176). Grid cells with or without particular geological units differed in total and forest species richness, with those containing peatland and marshland being particularly species-poor. Indicator value analysis showed that both total and forest species richness in the grid cells were related to soil acidity and nitrogen in a hump-backed manner, with the highest number of species found at moderately low values for nitrogen and at intermediate values of pH. Forest species richness was highly positively correlated with the number of non-forest species and threatened non-forest species. Indicators for high species richness were primarily those species that are confined to closed semi-natural forests with a varied topography and relatively base- and nutrient-rich soils. Grid cells including historically ancient forest exhibited a higher species richness than grid cells lacking ancient forest, indicating the importance of a long habitat continuity for a high phytodiversity. The “habitat coincidence” of high species richness is best explained by similar responses of forest species and species of other habitats to the main environmental gradients. It is suggested that the regional patterns found for the Weser-Elbe region can be transferred also to other forest-poor regions in Central Europe.

Weak effects on plant diversity of two invasive Impatiens species

The study aimed to examine the effect of the invasion of the two congeneric species Impatiens glandulifera and Impatiens parviflora on species richness and composition across a wide range of communities in North-western Germany. We applied a space-for-time substitution approach, comparing invaded plots with adjacent, environmentally similar uninvaded plots, based on the assumption that the latter represent the situation prior to an invasion. Even though the dominance of the invasive species resulted in a lower Shannon diversity in the invaded plots, species richness was not (I. parviflora) or only weakly (I. glandulifera) reduced. Also the community composition of the invaded sites was only marginally different. Invaded and uninvaded plots in general had similar habitat conditions, but both Impatiens species occurred in slightly shadier sites compared to the uninvaded areas, and the plots invaded by I. parviflora tended to have higher nutrient concentrations. These results suggest that dense populations of invading species may often be found at particular microsites. The relatively low impact of Impatiens on the vegetation is most likely caused by the annual life strategy of the species: while I. glandulifera shows large fluctuations in numbers between years and has a later phenological development than most other species in its habitat, I. parviflora establishes preferably in dark and acidic, often disturbed woodlands where it competes with few other species.