Impact of herbivores on nitrogen cycling: contrasting effects of small and large species.

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing.

Growth reduction of Sphagnum magellanicum subjected to high nitrogen deposition: the role of amino acid nitrogen concentration.

We tested the relationship between Sphagnum growth and the amount of nitrogen stored in free amino acids in a fertilisation experiment with intact peat monoliths in an open greenhouse in The Netherlands. Three nitrogen deposition scenarios were used: no nitrogen deposition, field conditions and a doubling of the latter, corresponding to 0, 40 and 80 kg N ha(-1 )year(-1). Growth of Sphagnum as expressed by height increment was reduced in the 80 kg N treatment, but showed no correlation with the total nitrogen tissue concentration or with the concentration of individual or pooled free amino acids. The amount of nitrogen stored in free amino acids increased concomitantly with deposition, although it lagged more and more behind the total nitrogen concentration, the latter pointing to the accumulation of unmeasured nitrogen compounds. Asparagine clearly acted as the major storage compound for nitrogen in Sphagnum stem tissue, whereas arginine fulfilled this function to a lesser extent in the capitulum. It appears that nitrogen-induced growth inhibition of Sphagnum is related to acclimation rather than to certain threshold concentrations of amino nitrogen or total nitrogen. We propose that when Sphagnum is exposed to a step increase of nitrogen, its nitrogen metabolism does not adapt fast enough to keep up with the enhanced uptake rate. This imbalance between nitrogen uptake and assimilation may lead to an accumulation of toxic NH(4)(+ )in the cell and a subsequent reduction in growth.

Hydraulic lift in Acacia tortilis trees on an East African savanna.

Recent studies suggest that savanna trees in semi-arid areas can increase understorey plant production. We hypothesized that one of the mechanisms that explains the facilitation between trees and grasses in East African savannas is hydraulic lift (HL). HL in large Acacia tortilis trees was studied during the first 3 months of the dry season during a relatively wet year (1998) and a very dry year (2000). In 1998, we found distinct diel fluctuation in soil water potential (psi(s)), with increasing values during the night and decreasing again the following day. These fluctuations in psi(s )are consistent with other observations of HL and in A. tortilis were found up to 10 m from the tree. In 2000, during a severe drought, psi(s) measurements indicated that HL was largely absent. The finding that HL occurred in wetter years and not in drier years was supported by data obtained on the delta(18)O values in soil, rain and groundwater. The delta(18)O of water extracted from the xylem water of grasses indicated that when they grew near trees they had values similar to those of groundwater. This could be because they either (1) use water from deeper soil layers or (2) use hydraulically lifted water provided by the tree; this was not seen in the same grass species growing outside tree canopies. While our data indicate that HL indeed occurs under Acacia trees, it is also true that psi(s) was consistently lower under trees when compared to outside tree canopies. We believe that this is because tree-grass mixtures take up more water from the upper soil layers than is exuded by the tree each night. This limits the beneficial effect of HL for understorey grasses and suggests that in savannas both facilitation via HL and competition are active processes. The importance of each process may depend upon how wet or dry that particular site or year is.

Agri-environment schemes do not effectively protect biodiversity in Dutch agricultural landscapes.

Roughly 20% of the European Union's farmland is under some form of agri-environment scheme to counteract the negative impacts of modern agriculture on the environment. The associated costs represent about 4% (1.7 billion euros) of the European Union's total expenditure on the Common Agricultural Policy and are expected to rise to 10% in the near future. Although agri-environment schemes have been implemented in various countries for well over a decade, to date no reliable, sufficiently replicated studies have been performed to test whether such measures have the presumed positive effects on biodiversity. Here we present the results of a study evaluating the contribution of agri-environment schemes to the protection of biodiversity in intensively used Dutch agricultural landscapes. We surveyed plants, birds, hover flies and bees on 78 paired fields that either had agri-environment schemes in the form of management agreements or were managed conventionally. Management agreements were not effective in protecting the species richness of the investigated species groups: no positive effects on plant and bird species diversity were found. The four most common wader species were observed even less frequently on fields with management agreements. By contrast, hover flies and bees showed modest increases in species richness on fields with management agreements. Our results indicate that there is a pressing need for a scientifically sound evaluation of agri-environment schemes.

Constraints in the restoration of ecological diversity in grassland and heathland communities.

Species-rich grassland and heathland communities still occur in low-intensity farming systems in many European countries. Gradually, such systems have either been abandoned or more intensively exploited, with a subsequent decrease in species numbers. Until recently, it was thought that restoration of these communities would be straightforward. However, abiotic constraints (with respect to eutrophication and acidification) have hampered restoration more than previously thought. Moreover, very recent research has revealed that biotic constraints can also be important: many plant species are not present in the soil seed bank and their dispersal is limited in the present fragmented landscape.

Nitrogen losses from perennial grass species.

Nitrogen losses from plants may occur through a variety of pathways, but so far, most studies have only quantified losses of nutrients by above-ground litter production. We used 15N pulse labelling to quantify total nitrogen losses from above- and below-ground plant parts. Using this method we were able to include also pathways other than above-ground litter production. To test the hypothesis that species from nutrient-poor habitats lose less nitrogen than species from more fertile soils, six perennial grasses from habitats with a wide range of nutrient availability were investigated: Lolium perenne, Arrhenatherum elatius, Anthoxanthum odoratum, Festuca rubra, F. ovina and Molinia caerulea. The results of an experiment carried out in pots in a green-house at two fertility levels show that statistically significant losses occur through pathways other than above-ground litter production. In the low fertility treatment, most (70%) losses from L. perenne occurred by litter production, but in Ar. elatius, F. rubra, F. ovina and M. caerulea, more than 50% of labelled N losses took place by root turn-over, leaching or exudation from roots. When nutrient supply increased, the 15N losses in above-ground dead material increased in all species and in Ar. elatius, A. odoratum and F. rubra the 15N losses via other pathways decreased. Ranked according to decreasing turnover coefficient the sequence of species was: L. perenne, A. odoratum, F. rubra, F. ovina, Ar. elatius, M. caerulea. These results suggest that species adapted to sites with low availability of nutrients lose less nitrogen (including above- and below-ground losses) than species adapted to more fertile soils.

Root production and root turnover in two dominant species of wet heathlands.

Root biomass production, root length production and root turnover of Erica tetralix and Molinia caerulea were estimated by sequential core sampling and by observations in permanent minirhizotrons in the field. Root biomass production, estimated by core sampling, was 370 (Erica) and 1080 (Molinia) g m-2 yr-1. This was for both species equal to aboveground production. Assuming steady-state conditions for the root system, root biomass turnover rates (yr-1), estimated by core sampling, were 1.72 (Erica) and 1.27 (Molinia). Root length production of both species, estimated by minirhizotron observations, varied significantly with observation depth. Root length turnover rate (yr-1) of both species did not vary significantly with observation depth and averaged 0.92 in Erica and 2.28 in Molinia. Reasons are given for the discrepancy between the results of the two types of turnover measurements. The data suggest that the replacement of Erica by Molinia in a wet heathland, which occurs when nutrient availability increases, leads to an increased flow of carbon and nutrients into the soil-system. Therefore, there may be a positive feedback between dominance of Molinia and nutrient availability.

A comparative study on nutrient cycling in wet heathland ecosystems : I. Litter production and nutrient losses from the plant.

The term relative nutrient requirement is introduced in order to measure and to compare the nutrient losses from different perennial plant populations and the amount of nutrient that they need to absorb to compensate these losses. The relative nutrient requirement (L) is defined as the amount of the growth-limiting nutrient that must be taken up to maintain or replace each unit of biomass during a given time interval (e.g., mgN g-1 biomass year-1). It is derived that in a plant community with two competing perennial plant populations, species1 will become dominant if the relative competition coefficient k 12 (sensu De Wit 1960) exceeds the ratio between the relative nutrient requirements of the two species (L 1/L 2), whereas species 2 will become dominant, if k 12 is below this critical ratio. The above-ground litter production was measured inwet heathland communities dominated by Erica tetralix or by Molinia caeruleain order to estimate N and P losses from theaboveground biomass and to calculate the relative N and P requirements of these species. Molinia lost during one year 63% and 34%, respectively, of the amount of N and P present in the above-ground biomass at the end of the growing season. These losses were in Erica 27% and 31%, respectively. The relative N requirements of the two species show the same difference: 7.5 and 2.6 mg N g-1 yr-1, respectively, in Molinia and in Erica. Also the relative P requirement of Molinia is higher as well as that of Erica (0.18 versus 0.08 mg P g-1 yr-1). The relative competition coefficient of Molinia with respect to Erica (k me ) is equal to unity under unfertilized conditions but increases with increasing nutrient supply. Under nutrient-poor conditions k me is below the critical ratio of the relative nutrient requirements of the two species (L m /L e =2.9 or 2.3), so that Erica will be the dominant species. After an increase in nutrient availability k me increases and exceeds this critical limit which results in Molinia replacing Erica. During the last 20 years this replacement of Erica-dominant communities by monocultures of Molinia has been observed in almost all wet heathlands in The Netherlands along with a strong increase in nitrogen availability.