Camilla Wellstein

Worldwide evidence of a unimodal relationship between productivity and plant species richness

Grassland diversity and ecosystem productivity The relationship between plant species diversity and ecosystem productivity is controversial. The debate concerns whether diversity peaks at intermediate levels of productivity—the so-called humped-back model—or whether there is no clear predictable relationship. Fraser et al. used a large, standardized, and geographically diverse sample of grasslands from six continents to confirm the validity and generality of the humped-back model. Their findings pave the way for a more mechanistic understanding of the factors controlling species diversity. Science, this issue p. 302 The humped-back model of plant species diversity is confirmed by a global grassland survey. The search for predictions of species diversity across environmental gradients has challenged ecologists for decades. The humped-back model (HBM) suggests that plant diversity peaks at intermediate productivity; at low productivity few species can tolerate the environmental stresses, and at high productivity a few highly competitive species dominate. Over time the HBM has become increasingly controversial, and recent studies claim to have refuted it. Here, by using data from coordinated surveys conducted throughout grasslands worldwide and comprising a wide range of site productivities, we provide evidence in support of the HBM pattern at both global and regional extents. The relationships described here provide a foundation for further research into the local, landscape, and historical factors that maintain biodiversity.

European grassland ecosystems: threatened hotspots of biodiversity

Biodiversity is not homogenously distributed over the globe, and ecosystems differ strongly in the number of species they provide. With this special issue we highlight the ecology and endangerment of one of the most diverse ecosystem of Europe: the European grassland ecosystems. The selected 16 contributions describe interactions from below-ground to the atmosphere and focus on (1) effects of abiotic and biotic on species diversity, (2) the impact of various factors along spatial and temporal gradients, (3) the relevance of falling abandoned and eutrophication—including countervailing management strategies like encroachment; and (4) intraspecific effects based on physiology, genetics and intraspecific plasticity. The contributions cover fungi, plants, and invertebrates and highlight effects taking place at the level of ecosystem, species community, species, populations, and also within individuals (physiology and genetics).

Seed bank diversity in mesic grasslands in relation to vegetation type, management and site conditions

Abstract Questions: 1. Do different management types (i.e. hay meadow, silage meadow, meadow-pasture, pasture) have different impact on the size and composition of the seed bank of mesic grassland (Arrhenatheretalia)? 2. How strong is the effect of management on the seed bank in relation to above-ground vegetation, edaphic factors and land-use history? 3. Are there differences in C-S-R plant strategy types and seed longevity under different management regimes? Location: Lahn-Dill Highlands in central-western Germany. Methods: Above-ground vegetation and the soil seed bank of 63 plots (at 21 sites) in mesic grasslands were studied. Differences between management types in quantitative seed bank traits and functional characteristics were tested by ANOVA. The impact of management, above-ground vegetation, site conditions and land-use history on seed bank composition were analysed by partial CCA. Results: Management had no significant impact on species richness and density of the seed bank but significantly influenced their floristic composition and functional characteristics. CCA revealed that even after adjustment for soil chemical parameters and above-ground vegetation management still had significant impact on seed bank composition. ANOVA revealed that silage meadows contained higher proportions of R-strategy compared to hay meadows. In contrast, in hay meadows and meadow-pastures proportions of S-strategy were higher than in silage meadows. Conclusions: The type of grassland management has little impact on quantitative seed bank traits. Management types with a high degree of disturbance lead to an increase of species following a ruderal strategy in the seed bank. Irrespective of management type only a limited proportion of characteristic grassland species is likely to re-establish from the seed bank after disappearance from above-ground vegetation. Nomenclature: Wisskirchen & Haeupler (1998).

Patterns of Clonal Growth Modes Along a Chronosequence of Post-Coppice Forest Regeneration in Beech Forests of Central Italy

Forest coppicing leads to changes in composition of the herbaceous understory through soil disturbance and alteration of the light regime. While the role of seed dispersal traits at the start of succession after coppicing has been extensively studied, the role of persistence traits such as clonal growth and bud banks is not yet sufficiently understood. To gain better understanding of this role, we studied the patterns of clonal growth organs and related clonal traits of species in a series of coppiced beech forests of the Central Apennines (Marches region, Italy) in various stages of recovery after the last coppicing event. We conducted stratified random sampling and established a chronosequence of recovery stages based on stand age (reflecting the number of years since the last coppicing). The beech stands were classified into three age groups (Post-logged, Recovering, and Old-coppice stands) according to the characteristic stages of beech coppice dynamics. Clonal growth organs and the corresponding clonal traits of plants in the forest understory vegetation were assessed with the help of a CLO-PLA1 database. We found no significant change in the proportion of clonal species along the studied chronosequence. In contrast, most of the traits and about the half of the clonal growth organs showed correlation with stand age or preference for a certain habitat (i.e., stage of regeneration). Clonal and bud bank traits proved to play an important role in the persistence of species subjected to forest coppicing cycles in the studied area.

Seed-litter-position drives seedling establishment in grassland species under recurrent drought.

Changes in land use and climate interfere with grassland ecosystem processes. Here I experimentally investigated the combined effects of land-use change related litter cover and contrasting water supply on seedling emergence. In this context, the role of the initial relative position of seeds, i.e. seeds on top of the litter versus seeds beneath the litter in interaction with water supply has not been investigated so far. I hypothesised that facilitative effects of litter on seedling emergence occur when seeds are covered by litter and deteriorate when litter covers the ground and seeds fall on it (seeds on top of the litter). Further, I hypothesised that the importance of seed position for seedling emergence will increase under conditions of recurrent drought. I performed a controlled pot experiment on seedling emergence of three common European grassland species (Pimpinella saxifraga, Leontodon autumnalis, Sanguisorba officinalis) by experimental manipulations of litter and water availability. Seedling emergence under moist conditions showed no significant differences between each litter position compared to the control across species. In contrast, under recurrent drought, seedling emergence was significantly higher below the litter compared to seeds on top of the litter and the control (i.e. no litter). In abandoned land, seedling emergence may be limited when seeds fall on ground-covering litter. In contrast, in grasslands with regular low-intensity land use, seedling emergence may be enhanced when a moderate level of litter covers seeds at the end of the growing season. Protective mechanisms that occur with seeds positioned beneath litter are particularly important under recurrent drought.

Impact of Extreme Drought and Warming on Survival and Growth Characteristics of Different Provenences of Juvenile Quercus pubescens Willd

In the context of climatic changes, the adaptive potential of organisms toward new environmental challenges is becoming of central interest in modern ecology. Here we investigate the response of growth characteristics and survival in juveniles of four European provenances (Bulgaria, Germany, Hungary and Italy) of Quercus pubescens Willd. to extreme drought and warming in a full factorial common garden experiment. Our results show that extreme durations of drought exceed the tolerance of Quercus pubescens of all provenances. Plants of Italian provenance showed the highest capacity of whole-individual survival, and those of Bulgarian provenance showed the highest maintenance of the apex under extreme drought. Bud bank characteristics showed no general differences among provenances but responded to warming manipulations. The warming effect resulted in a reduction of collar buds and biomass. In Italian individuals it caused an increase of the apical budbank. This is suggestive of a better adaptation for height growth under warming. Our results imply that significant local adaptation of growth characteristics and survival occurs in Quercus pubescens. However, our findings do not support the idea that southern provenances are a general predictor of better plant performance.

Diversity and Frequency of Clonal Traits Along Natural and Land-Use Gradients in Grasslands of the Swiss Alps

The frequency of clonal plants in different vegetation types is known to be influenced by environmental and land-use factors. However, the underlying behavior of individual clonal traits or clonal trait diversity has received little attention. Here, we assess for species- and trait-diverse grasslands of the Swiss Alps the relative importance of temperature, soil moisture, land use and species richness on the diversity and frequency of individual compared with all clonal traits. We further analyzed how cover-weighted data alters the relationships found with commonly used presence-absence data. We combined species compositional, land-use and environmental data from 236 28-m2 grassland plots with clonal trait information for 527 species following the Clonal Growth Organ (CGO) classification. Test results are based on linear models, ANOVAs and ANCOVAs. The grassland sites were 84% dominated by clonal species. Drought-prone grasslands harbored the least clonal species. No increase in clonality was detected with decreasing temperature (= altitude). Mown or pastured grasslands had more clonal species than fallows. Certain sets of traits were correlated. Rhizomatous species especially reacted strongly to climatic and land-use gradients and had highest frequencies in cold, moist and disturbed sites. Clonal diversity was strongly dependent on species richness. Cover-weighted and presence-absence based estimates were largely similar. Overall, our data outlined that common clonal traits react differently to natural and land-use gradients as well as differently to the sum of clonal traits. Also, soil moisture was more decisive than temperature (= altitude) for the presence of clonal species. Lastly, the strong correlation between species-richness and clonal trait diversity needs to be accounted for when interpreting the functional role of clonal traits.

Warming and drought do not influence the palatability of Quercus pubescens Willd. leaves of four European provenances

In the context of global warming, the impact of extreme drought events on trees and biotic interactions with herbivore insects is widely unknown. A faster range expansion of insects in a changing climate could lead to mass propagations of pests in forests. Therefore, the aim was to investigate the influence of climatic alterations on leaf palatability. We exposed juvenile Quercus pubescens Willd. individuals of four European provenances (Bulgaria, Germany, Hungary, and Italy) to warming and drought. In addition, we conducted a palatability experiment with the pre-exposed Q. pubescens leaves and the caterpillars of the generalist forest pest Lymantria dispar L. (gypsy moth). Consumed leaf dry material, density of trichomes, and specific leaf area were examined. Surprisingly, neither warming nor drought affected the leaf palatability, but palatability was related to the density of trichomes. The Bulgarian provenance of Q. pubescens, which had the lowest density of trichomes, was most palatable. These findings suggest that global warming and drought might not lead to more frequent infestations of the four tested European Q. pubescens provenances by L. dispar caterpillars in the future.

Unravelling mechanisms of short-term vegetation dynamics in complex coppice forest systems

The silvicultural management of coppicing has been very common in deciduous forests in many European countries. After decades of decline of this practice, socio-economic changes might induce a revival valuing the biomass as a resource. New insights in the ecological processes that regulate plant diversity are relevant for a sustainable forest management. While studies on long-term changes are available, the short-term dynamics of the coppice forest understorey has not yet been explored. In this context, it is interesting to evaluate the species compositional changes, including the processes of species turnover and species impoverishment (nestedness) and to investigate the role of plant functional traits. For this purpose, we resampled a chronosequence of complex coppice beech forests of the Central Apennines (Italy) monitoring the short-time species dynamics of five years (i.e. from 2006 to 2011) in three age classes, i.e. post-logged, recovering and old coppice stands (0–16, 17–31 and > 32 years, respectively). In contrast to our expectation, declining species richness appeared only in the recovering stands, while the landscape scale (between-stand) heterogeneity, except for post-logged and recovering stands in 2011, did not change over five years. Significant temporal nestedness was found in each stage of succession. However, the rate of species turnover and species impoverishment do not significantly differ among the three age classes, indicating their constant importance along the forest regeneration after disturbance. Only in the early stage of forest regeneration after coppicing, species compositional changes are reflected by functional changes with surviving understorey species having clonal regeneration traits. Our results suggest an overall landscape-scale stability (and sustainability) of this coppice forest system. We conclude with management indications, highlighting the importance of maintaining the traditional local approach (coppicing with standards in small 0.5–1.0 ha sized management units with a ca 30-year rotation cycle) where active coppice parcels are interspersed by abandoned stands.

The Status of Quercus pubescens Willd. in Europe

Taxonomic and nomenclatural disagreements are still encountered in the study of Quercus pubescens Willd. in Europe and are discussed here. This includes two current antithetical viewpoints on this taxon, i.e. the acceptance of the huge phenotypic variability among and within its populations within a single species vs. the ranking of these phenotypes as distinct species within the subgenus Quercus (the European white oaks sensu Schwarz).