Goddert von Oheimb

Community assembly during secondary forest succession in a Chinese subtropical forest

Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30 × 30 m plots (n = 27) from five successional stages ( 1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10 × 10 m quadrant of each plot. In addition, we measured a number of environmen...

The effects of light and soil conditions on the species richness of the ground vegetation of deciduous forests in northern Germany (Schleswig-Holstein)

Abstract This paper reports a study carried out in 91 deciduous forests to investigate the influence of light and soil conditions on the species richness of the ground vegetation (vascular plants, bryophytes, lichens). To examine the relationships between the species richness and environmental factors all data were evaluated using univariate and multivariate (principle component analysis) methods. At all the stands investigated, the conditions of soil and light have a vegetation type specific effect on the species richness of the ground vegetation. In moist forests (alder-ash forests of the alliance Alno-Ulmion ), species richness has a close positive correlation with soil moisture, whilst light conditions and nutrient supply have in the main no effect on species richness. On the other hand, in meso- to eutrophic beech forests (beech forests of the alliance Fagion sylvaticae ), species richness is closely correlated with soil activity and with the base and nitrogen supply. Improving the light conditions for the ground vegetation does not increase the number of typical woodland species as most of these are shade tolerant and have very limited light requirements. Species richness in acidophytic beech and mixed beech–oak forests (beech and mixed beech–oak forests on acid soils of the order Quercetalia roboris ) is affected mainly by the canopy closure and the interior light conditions. Soil moisture, nutrient supply and light availability must therefore be evaluated differently for their effect on the number of species of the ground vegetation and with regard to individual forest communities as they are distinguishable according to their site conditions. If the relation between species richness and environmental factors like soil conditions is analysed not with reference to individual forest communities but rather with reference to the totality of central European forests, relationships may be found which are not helpful for an explanation of the species richness of a certain forest community and thus they are only of restricted usefulness for appropriate recommendations for forest management. The consequences of the results for the effects of silvicultural treatment on deciduous forest ecosystems of the central European lowlands are discussed.

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

Summary 1. Biodiversity–ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25� 8 9 25� 8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ‘ecoscape’ to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev

The effects of windthrow on plant species richness in a Central European beech forest

The effects of soil disturbance caused by the uprooting of a single or a few canopy trees on species richness and composition of vascular plant species and bryophytes were examined in a temperate beech forest (Fagus sylvatica) in northern Germany. We recorded the vegetation in 57 pairs of disturbed and adjacent undisturbed plots and established a chronosequence of mound ages to study the effect of time since microsite formation on plant species richness and composition. We found significant differences in plant species richness and composition between disturbed and adjacent undisturbed plots. Species richness of both vascular plants and bryophytes was higher in the disturbed than in the undisturbed plots, but these differences were more pronounced for bryophytes. We suggest that three main factors are responsible for this differential response. The availability of microsites on the forest floor that are suitable for the recruitment of bryophytes is lower than for vascular plants. Establishment of bryophytes in disturbed microsites is favoured by a greater abundance of propagules in the close vicinity and in the soil of the disturbed microsites, as well as by a greater variety of regeneration strategies in bryophytes than in vascular plants. Time since mound formation was a major factor determining plant species richness and composition. A significant decrease in the mean number of species was found from young mounds to intermediate and old mounds. However, differences were observed between vascular plants and bryophytes in the course of changes through time in species richness and composition. A large number of exclusive and infrequent vascular plant species was observed on young mounds, among them several disturbance specialists. We suggest that the establishment of many vascular plant species was infrequent and short-lived due to unfavourable light conditions and a low abundance of propagules. By contrast, the development of a litter layer was the main reason for the decreased mean number of bryophytes on old mounds. Our study supports the view that groups of species differing in important life history traits exhibit different responses to soil disturbance.

Dispersal of vascular plants by game in northern Germany. Part II: Red deer (Cervus elaphus)

The endozoochorous dispersal of vascular plant species by red deer (Cervus elaphus) was investigated in three forest areas of Schleswig-Holstein and Lower Saxony. The primary aim of this study was to determine of which plant species and in what numbers diaspores are dispersed by red deer, an animal with eminently large home ranges during a single day as well as over the year. A second aim was to examine the significance of endozoochory for plant species composition and diversity in forests. Special emphasis was placed on whether there are differences between the three study sites and in the course of the vegetation period. Freshly deposited faecal pellet groups were collected in the forests on five dates between May and November 2002 and viable seed content was determined by greenhouse germination (seedling emergence method). A total of 28,009 individuals representing 59 vascular plant species were detected in 220 dung samples. Red deer appears thus as an important vector for endozoochorous dispersal of plants. The number of seedlings found in the faecal pellets differs between plant species and research areas as well as in the course of the vegetation period. Dung seed content is largely dominated by one species, Urtica dioica. Grouping the species found in dung according to their habitat preference showed that mainly species that occur in forests as well as in open vegetation and non-forest species were dispersed endozoochorously, while species occurring mainly under a closed canopy were present in the excrements only in low numbers.

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH4+ availability in the O-horizon. We observed only short-term effects (for NO3−, PO43−, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.

Relationships between the vegetation and soil conditions in beech and beech-oak forests of northern Germany

In this study we examine the relationships between the vegetation of beech and beech-oak forest communities (Hordelymo-Fagetum, Galio-Fagetum, Deschampsio-Fagetum, Betulo-Quercetum) and their soil conditions in the lowlands of northern Germany, based on 84 sample plots. In all plots the vegetation was recorded and soil parameters were analysed (thickness of the O- and the A-horizons, pH, S-value, base saturation, C/N, mean Ellenberg moisture indicator value). The vegetation classification according to the traditional Braun-Blanquet approach was compared with the result of a multivariate cluster analysis. Vegetation-site relationships were analysed by means of an indirect gradient analysis (DCA).Both traditional classification methods and the cluster analysis have produced comparable classification results. So far as the species composition is concerned, a similar grouping of sample plots was found in both approaches. Multivariate cluster analysis thus supports the classification found by the Braun-Blanquet method. The result of the DCA shows that the four forest communities mentioned above represent clearly definable ecological units. The main site factor influencing changes in the species composition is a base gradient, which is best expressed by the S-value. In addition, within the series Hordelymo-Fagetum - Galio-Fagetum - Deschampsio-Fagetum the C/N-ratios and the thickness of the organic layers (O-horizon) increase continuously. By contrast, the floristic differences between oligotrophic forest communities (i.e., Deschampsio-Fagetum and Betulo-Quercetum) cannot be explained by a base gradient and increasing C/N-ratios. It is suggested that a different forest management history in some cases (e.g., promotion of Quercus robur by silvicultural treatments) is responsible for differences in the species composition, but on the other hand the result of the DCA indicates that Fagus sylvatica is replaced by Quercus robur with increasing soil moisture (i.e., with the increasing influence of a high groundwater table). Summarizing these results, it can be concluded that the ecological importance of single site factors affecting the species composition changes within the entire site spectrum covered by the beech and beech-oak forests of northern Germany.

Relationship between pH-values and nutrient availability in forest soils - the consequences for the use of ecograms in forest ecology

Summary Measurements of the soil pH are often used to characterize the nutritional conditions for the vegetation at a particular site, for example in ecograms describing the ecological behaviour of a (tree) species. In spite of this use and interpretation of pH measurements, certain prerequisites must be fulfilled when pH-values are to be used as an expression of the nutritional conditions for plants. Our study in deciduous forests of northern Germany shows that in strongly acid soils (pH

Mixed afforestation of young subtropical trees promotes nitrogen acquisition and retention

Summary 1. Afforestation is globally increasing to produce timber and pulp wood, but also to enhance ecosystem services such as carbon sequestration, nutrient retention or groundwater recharge. In China, large areas have been and will be afforested in order to compensate for the negative impacts of former clear-cuttings and to make use of the ecosystem services associated with afforestation. In order to further optimize these services with regard to balanced nutrient (particularly nitrogen) cycles, it is important to know whether the use of mixtures of native tree species in afforestation projects promotes the acquisition and retention of nitrogen compared with the currently established large-scale monocultures. 2. To test the effect of species richness on system N retention and tree sapling N uptake, we conducted a 15 N tracer experiment in a young tree plantation. To this end, saplings of four abundant early successional tree species were planted in monocultures, in two- and four-species mixtures and as single trees. 3. Nitrogen retention increased with higher species richness due to enhanced N pools in sapling biomass. These species richness effects strengthened over time. 4. Species-specific differences in 15 N recoveries over time revealed below-ground niche differentiation with regard to N uptake, which is likely to result in complementary resource use among coexisting species. 5. Synthesis and applications. This study provides evidence that mixed afforestation promotes N retention from the sapling stage. To further improve ecosystem services associated with afforestation, we strongly suggest the use of mixtures of native tree species instead of monocultures. Mixtures of four species may reduce system N losses and thus may lessen groundwater contamination due to N leaching. We encourage further investigations to find optimal species combinations that promote a wide range of ecosystem services related to more closed nutrient cycles and minimized soil erosion. In our study, the plantations’ capability to retain N could be optimized by means of both increasing tree species richness and by choosing the optimal species combinations.

Nitrogen deposition increases susceptibility to drought - experimental evidence with the perennial grass Molinia caerulea (L.) Moench

AimsWe investigated the response of the perennial grass Molinia caerulea (L.) Moench to combined effects of fertilization (N, P) and drought events. We hypothesized that N fertilization increases, and drought decreases productivity, but that N addition strengthens negative effects caused by drought.MethodsWithin a full-factorial 2-year greenhouse experiment we measured biomass productivity and allocation, tissue nutrient concentrations and nitrogen allocation patterns using 15N as a tracer.ResultsN fertilization caused a strong increase in productivity, but effects of drought were almost insignificant. However, we found strongly interrelated, non-additive effects of fertilization and drought, expressed by a strong increase of necrotic tissue. Dead aboveground biomass showed the highest values for N and 15N.ConclusionsAccelerated productivity of aboveground tissue under N fertilization resulted in increased evaporative demands and thus higher drought susceptibility. In addition 15N allocation patterns showed that fertilization-drought treatments disenabled plants’ control of their N allocation. Molinia was unable to withdraw leaf N during the dieback of aboveground tissue. Due to the lack of an adaptive strategy to the combined effects of fertilization and drought, increasing summer drought may weaken the competitive performance of species with traits comparable to those of Molinia in N-fertilized environments.