Long-term effects of nitrogen deposition on vegetation in a deciduous forest near Munich,Germany

Question: What are the main reasons for changes in the spatial distribution of vegetation types during the last four decades? Location: Isolated small deciduous forest; surrounded by farmland in the northeast of Munich (Germany). Methods: Based on sequential vegetation mapping from the last four decades the spatial development of the vegetation was analysed. Additionally, environmental parameters (soil parameters, PAR, N-deposition) have been analysed to describe the different vegetation types. Results: By linking the vegetation types to environmental parameters, it was possible to identify N-deposition as the most important factor for the changes. In the 1960s to 1980s the replacement of vegetation types adapted to N-poor conditions by N-rich vegetation was very fast. A vegetation type containing species signifying soil impoverishment vanished totally, another vegetation type indicating nutrient poor conditions decreased dramatically. However, since 1985 up to now the decrease of N-poor vegetation types has slowed, but is still ongoing. As a reason for the decreased rate of replacement, we stressed changes in the vertical structure: From 1961 to 1985 both N-deposition as well as changes in vertical vegetation structure seem to be important. Since 1985 up to now only minor changes in vertical structure could be found; changes are mainly due to N-availability. Conclusion: In this paper, the limitations of different methods to detect vegetation changes are discussed. We focus on the potentials of historical vegetation data and vegetation maps. It is shown that valuable information on N-induced vegetation changes can be retrieved from historical vegetation data.     

The interrelationship between productivity,plant species richness and livestock diet: a question of scale?

Question: What relationship exists between productivity, plant species richness and livestock diet? Are the results dependent on scale? Location: A sheep‐grazed Koelerio‐Corynephoretea sandy habitat of the northern upper Rhine (Germany) as a low productivity model system. Methods: The investigation was carried out for three years at a fine scale (2 m²) and for two years at a broad scale (79 m²). Productivity was measured by means of weighed above‐ground phytomass for fine scale and colour‐infrared (CIR) aerial photographs of the same system for fine and broad scales. For both scales, total numbers of vascular plant species and numbers of endangered vascular plant species were extracted from current vegetation relevés. Additionally, we obtained data on livestock diet (grazed phytomass, crude protein content). Results: Statistical analyses show an influence of the year on all variables; relationships between variables are not significant in every year. Species richness and number of endangered species are negatively related to productivity at fine scale while crude protein content and grazed phytomass are positively related to productivity. At the broad scale the diversity‐productivity relationship shows a ‘hump’ with highest species numbers in middle pioneer stages; numbers of endangered species are highest in all pioneer stages. Conclusions: We found a strong impact of scale and year on the diversity‐productivity relationship. It is inappropriate to analyse only small plots (2 m²), and it is necessary to study different years. This vegetation complex is dependent on grazing impact; thus there is an inversely proportional relationship between nature conservation value (high diversity) and livestock nutrition.     

Prediction of herbage yield in grassland: How well do Ellenberg N‐values perform?

Question: How useful are Ellenberg N‐values for predicting the herbage yield of Central European grasslands in comparison to approaches based on ordination scores of plant species composition or on soil parameters? Location: Central Germany (11°00′‐11°37’E, 50°21‐50°34’N, 500–840 m a.s.l.). Methods: Based on data from a field survey in 2001, the following models were constructed for predicting herbage yield in montane Central European grasslands: (1) Linear regression of mean Ellenberg N‐, R‐ and F‐values; (2) Linear regression of ordination scores derived from Non‐metric Multidimensional Scaling (NMDS) of vegetation data; and (3) Multiple linear regression (MLR) of soil variables. Models were evaluated by cross‐validation and validation with additional data collected in 2002. Results: Best predictions were obtained with models based on species composition. Ellenberg N‐values and NMDS scores performed equally well and better than models based on Ellenberg R‐ or F‐values. Predictions based on soil variables were least accurate. When tested with data from 2002, models based on Ellenberg N‐values or on NMDS scores accurately predicted productivity rank order of sites, but not the actual herbage yield of particular sites. Conclusions: Mean Ellenberg N‐values, which are easy to calculate, are as accurate as ordination scores in predicting herbage yield from plant species composition. In contrast, models based on soil variables may be useful for generating hypotheses about the factors limiting herbage yield, but not for prediction. We support the view that Ellenberg N‐values should be called productivity values rather than nitrogen values.     

The impact of site conditions and seed dispersal on restoration success in alluvial meadows

Abstract. We studied the restoration success of flood plain meadows in the northern Upper Rhine valley, where between 1988 and 1992, 35 ha of arable land was converted into grassland and subsequently managed for nature conservation. Remnant populations of typical alluvial meadow species were found in old meadows and along drainage ditches that dissect the whole area. We analysed the site conditions and phytosociological relevés in old and new meadows. Small differences in site parameters between old and new meadows contrasted with a clear floristic differentiation between the two meadow types. The vegetation of old meadows was much more differentiated along prevailing environmental gradients than the vegetation of new meadows. Despite the favourable site conditions for the re‐establishment of species‐rich meadows on the former arable land, restoration success was limited to the vicinity of remnant stands. In contrast to old meadows, indicator species of new grassland were still typical species of regularly disturbed ruderal and arable habitats, often capable of building up a persistent seed bank. The precise mapping of 23 target species revealed that even wind dispersal predominantly leads to re‐establishment in the close circumference of parent plants. We found no indication that regular flooding, hay‐making and autumnal grazing had an impact on recolonization of newly created grassland. Even under favourable conditions for the re‐establishment of target species, restoration success in alluvial meadows proved to be strongly dispersal limited. We discuss the implications of our findings for future restoration management in grasslands.     

Dispersal,diversity and distribution patterns in pioneer vegetation: The role of river‐floodplain connectivity

Question: Are there changes in dispersal patterns in floodplain pioneer vegetation with effects on seedling number, species richness and species composition along a gradient of declining river‐floodplain connectivity? Location: Middle Elbe river floodplain, Germany. Methods: An experiment with five treatments was set up along a gradient of declining river‐floodplain connectivity, partitioning seedlings into three groups: (1) emerging solely from water dispersed seeds, (2) from wind/animal dispersed seeds and (3) from the soil diaspore bank. Two controls were established: without any manipulation and exclusion of all seeds. The results were compared with those of vegetation and soil sampling to evaluate the representativeness of the experimental sites in terms of species composition, diversity, seedling number and soil parameters. Results: Water dispersal and the soil diaspore bank were the major dispersal strategies shaping floodplain pioneer vegetation at the Middle Elbe river. The number of seedlings, species richness and the variation in species composition in these habitats depend on the degree of connectivity. The seedling number and species richness is highest in sites of permanent or almost permanent exchange with the main channel, where water dispersal additionally contributes to the number of seedlings grown from the soil seed bank. Conclusion: The results underline the importance of river‐floodplain ecotones as sink habitats for water‐dispersed seeds. Considering the strongly reduced river‐floodplain interactions due to dykes and other engineering structures, management strategies are necessary to improve connectivity and the renewal of fluvial land forms.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Assessing soil seed bank persistence in flood‐meadows: The search for reliable traits

Abstract. To assess seed bank persistence of target species in endangered flood‐meadows (alliances Cnidion and Molinion), we investigated established vegetation and soil seed bank of 46 plots for 3 yr and 2 yr, respectively. As traits of seed persistence we calculated various continuous indices that refer to the frequency and abundance of species in above‐ground vegetation and at different soil depths. Furthermore, we tested the significance and soundness of easily observed traits such as maximum seed density per plot and seed attributes (mass, size and shape) as predictors of soil seed bank features. In linear regression, SAI, the seed accumulation index, showed the best agreement (R²= 0.64) with the seed longevity index that was derived from the database by Thompson et al. (1997) for a set of 115 species. The second best predictor (R²= 0.39) of the seed longevity index was maximum seed density per plot in the lower soil layer (5–10 cm). Depth distribution indices and seed attributes showed weaker but still significant relations. The dynamic character of flood‐meadows was reflected by a large proportion of species with a strong tendency to accumulate seeds in the soil relative to their importance in above‐ground vegetation. Most of these species have a ruderal strategy, exploiting gaps after flood disturbances, while the dominants of flood‐meadows tended to have short‐lived seed banks. Compared to other grassland types, a relatively large proportion of rare and endangered target species can be expected to form long‐term persistent seed banks. However, only under marginal conditions that facilitate seed survival in the soil (e.g. fallow) are these persistent seed banks likely to contribute to restoration. We conclude that easily observed traits of persistence such as seed weight, size and shape do not meet the accuracy needed in scientific and practical applications. Thus, there is a crucial demand for further seed bank studies in poorly investigated habitats and of rare species.     

The impact of flooding regime on the soil seed bank of flood‐meadows

Abstract. We assessed the significance of flooding for the floristic composition of seed banks in flood‐meadows of the northern valley of the Upper Rhine. We compared three hydrological compartments of the alluvial plain, consisting of the regularly flooded land between the river and low summer dykes (functional flood‐plain), the occasionally flooded land between summer dykes and high winter dykes (hybrid floodplain) and the land behind the winter dykes, which is now only submerged by ascending groundwater (fossil flood‐plain). Due to their different flooding regime, the three compartments should differ with respect to the prevailing conditions of diaspore input. The seed density of soil samples increased with the duration of flooding in the three compartments, while species richness and the proportion of species not occurring in the vegetation was constant. The increase in seed density can be largely attributed to an increase of disturbance indicators, which are present in the above‐ground vegetation and capable of forming a long‐term persistent seed bank. No effects of flooding on the composition of seed banks in the three flood‐plain compartments were found. The differences in seed bank composition can be largely explained by corresponding differences in above‐ground vegetation and former and present‐day meadow management. Seeds of species absent from above‐ground vegetation can be attributed to the local species pool present in the immediate vicinity of the study plots. We discuss consequences of the results for the restoration of species‐rich flood‐plain meadows.     

Effects of topsoil removal,seed transfer with plant material and moderate grazing on restoration of riparian fen grasslands

Question: How do moderate grazing, topsoil removal and hay transfer affect species diversity and abundance on a eutrophic fen grassland site? Location: Northern Germany. Method: A three-factorial field experiment with the factors grazing, topsoil removal and hay transfer of diaspore-rich material was established in 2001. Soil nutrients and seed bank were analysed at the beginning of the experiment, species composition and vegetation development was monitored for four years (2002–2005). Results: Topsoil removal had a significant effect on the abundance of different plant species groups: resident vegetation of agricultural grasslands was suppressed, while clonal reed species were facilitated in recolonising the area. The establishment of regionally rare and endangered species of nutrient-poor fens and wet meadows introduced with hay was achieved mainly on plots with topsoil removal, with the exception of Rhinan-thus angustifolius, which also established on plots with intact topsoil. Effects of grazing after four years of experiments were of minor influence on species composition. Conclusion: The establishment of target plant species of nutrient-poor fens is most successful when both an adequate number of viable diaspores and suitable sites for germination and establishment are available. In our experiment this was achieved by the combination of topsoil removal and hay transfer. We recommend this combination, together with continuous management (grazing/cutting), for further restoration in fen grasslands.