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Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species-rich semi-natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on-going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades.
Are gastropods, rather than ants, important dispersers of seeds of myrmecochorous forest herbs?
(2012)
Seed dispersal by ants (myrmecochory) is widespread, and seed adaptations to myrmecochory are common, especially in the form of fatty appendices (elaiosomes). In a recent study, slugs were identified as seed dispersers of myrmecochores in a central European beech forest. Here we used 105 beech forest sites to test whether myrmecochore presence and abundance is related to ant or gastropod abundance and whether experimentally exposed seeds are removed by gastropods. Myrmecochorous plant cover was positively related to gastropod abundance but was negatively related to ant abundance. Gastropods were responsible for most seed removal and elaiosome damage, whereas insects (and rodents) played minor roles. These gastropod effects on seeds were independent of region or forest management. We suggest that terrestrial gastropods can generally act as seed dispersers of myrmecochorous plants and even substitute myrmecochory, especially where ants are absent or uncommon.
Carrion plays an essential role in shaping the structure and functioning of ecosystems and has far‐reaching implications for biodiversity conservation. The change in availability and type of carcasses throughout ecosystems can involve negative effects for scavenging communities. To address this issue, there have been recent conservation management measures of carrion provision in natural systems. However, the optimal conditions under which exposing carcasses to optimize conservation outcomes are still limited. Here, we used camera traps throughout elevational and vegetational gradients to monitor the consumption of 48 deer carcasses over a study period of six years by evaluating 270,279 photographs resulting out of 15,373 trap nights. We detected 17 species visiting carcass deployments, including five endangered species. Our results show that large carcasses, the winter season, and a heterogeneous surrounding habitat enhanced the frequency of carcass visits and the species richness of scavenger assemblages. Contrary to our expectations, carcass species, condition (fresh/frozen), and provision schedule (continuous vs single exposure) did not influence scavenging frequency or diversity. The carcass visitation frequency increased with carcass mass and lower temperatures. The effect of large carcasses was especially pronounced for mesopredators and the Eurasian lynx (Lynx lynx ). Lynx were not too influenced in its carrion acquisition by the season, but exclusively preferred remote habitats containing higher forest cover. Birds of prey, mesopredators, and top predators were also positively influenced by the visiting rate of ravens (Corvus corax ), whereas no biotic or abiotic preferences were found for wild boars (Sus scrofa ). This study provides evidence that any ungulate species of carrion, either in a fresh or in previously frozen condition, attracts a high diversity of scavengers especially during winter, thereby supporting earlier work that carcass provisions may support scavenger communities and endangered species.
Management intensity modifies soil properties, e.g., organic carbon (C-org) concentrations and soil pH with potential feedbacks on plant diversity. These changes might influence microbial P concentrations (P-mic) in soil representing an important component of the Pcycle. Our objectives were to elucidate whether abiotic and biotic variables controlling P-mic concentrations in soil are the same for forests and grasslands, and to assess the effect of region and management on P-mic concentrations in forest and grassland soils as mediated by the controlling variables. In three regions of Germany, Schwabische Alb, Hanich-Dun, and Schorfheide-Chorin, we studied forest and grassland plots (each n=150) differing in plant diversity and land-use intensity. In contrast to controls of microbial biomass carbon (C-mic), P-mic was strongly influenced by soil pH, which in turn affected phosphorus (P) availability and thus microbial Puptake in forest and grassland soils. Furthermore, P-mic concentrations in forest and grassland soils increased with increasing plant diversity. Using structural equation models, we could show that soil C-org is the profound driver of plant diversity effects on P-mic in grasslands. For both forest and grassland, we found regional differences in P-mic attributable to differing environmental conditions (pH, soil moisture). Forest management and tree species showed no effect on P-mic due to a lack of effects on controlling variables (e.g., C-org). We also did not find management effects in grassland soils which might be caused by either compensation of differently directed effects across sites or by legacy effects of former fertilization constraining the relevance of actual practices. We conclude that variables controlling P-mic or C-mic in soil differ in part and that regional differences in controlling variables are more important for P-mic in soil than those induced by management.
Factors controlling decomposition rates of fine root litter in temperate forests and grasslands
(2014)
Fine root decomposition contributes significantly to element cycling in terrestrial ecosystems. However, studies on root decomposition rates and on the factors that potentially influence them are fewer than those on leaf litter decomposition. To study the effects of region and land use intensity on fine root decomposition, we established a large scale study in three German regions with different climate regimes and soil properties. Methods In 150 forest and 150 grassland sites we deployed litterbags (100 mu m mesh size) with standardized litter consisting of fine roots from European beech in forests and from a lowland mesophilous hay meadow in grasslands. In the central study region, we compared decomposition rates of this standardized litter with root litter collected on-site to separate the effect of litter quality from environmental factors.
Standardized herbaceous roots in grassland soils decomposed on average significantly faster (24 +/- 6 % mass loss after 12 months, mean +/- SD) than beech roots in forest soils (12 +/- 4 %; p < 0.001). Fine root decomposition varied among the three study regions. Land use intensity, in particular N addition, decreased fine root decomposition in grasslands. The initial lignin:N ratio explained 15 % of the variance in grasslands and 11 % in forests. Soil moisture, soil temperature, and C:N ratios of soils together explained 34 % of the variance of the fine root mass loss in grasslands, and 24 % in forests.
Grasslands, which have higher fine root biomass and root turnover compared to forests, also have higher rates of root decomposition. Our results further show that at the regional scale fine root decomposition is influenced by environmental variables such as soil moisture, soil temperature and soil nutrient content. Additional variation is explained by root litter quality.
Intransitive competition is widespread in plant communities and maintains their species richness
(2015)
Intransitive competition networks, those in which there is no single best competitor, may ensure species coexistence. However, their frequency and importance in maintaining diversity in real-world ecosystems remain unclear. We used two large data sets from drylands and agricultural grasslands to assess: (1) the generality of intransitive competition, (2) intransitivity-richness relationships and (3) effects of two major drivers of biodiversity loss (aridity and land-use intensification) on intransitivity and species richness. Intransitive competition occurred in >65% of sites and was associated with higher species richness. Intransitivity increased with aridity, partly buffering its negative effects on diversity, but was decreased by intensive land use, enhancing its negative effects on diversity. These contrasting responses likely arise because intransitivity is promoted by temporal heterogeneity, which is enhanced by aridity but may decline with land-use intensity. We show that intransitivity is widespread in nature and increases diversity, but it can be lost with environmental homogenisation.
Recent declines in biodiversity have given new urgency to questions about the relationship between land-use change, biodiversity and ecosystem processes. Despite the existence of a large body of research on the effects of land use on species richness, it is unclear whether the effects of land use on species richness are principally direct or indirect, mediated by concomitant changes in ecosystem processes. Therefore, we compared the direct effects of land use (fertilization, mowing and grazing) on species richness with indirect ones (mediated via grassland productivity) for grasslands in central Europe. We measured the richness and above-ground biomass in 150 grassland plots in 3 regions of Germany (the so-called Biodiversity Exploratories). We used univariate and structural equation models to examine direct and indirect land-use effects. The direct effects of mowing (-0.37, effect size) and grazing (0.04) intensity on species richness were stronger compared with the indirect effects of mowing (-0.04) and grazing (-0.01). However, the strong negative effect of fertilization (-0.23) on species richness was mainly indirect, mediated by increased productivity compared with the weak direct negative effect (-0.07). Differences between regions in land-use effects showed five times weaker negative effects of mowing (-0.13) in the region with organic soils (Schorfheide-Chorin), strong overall negative effects of grazing (-0.29) for the region with organic soils opposed to a similar strong positive effect (0.30) in the Hainich-Dun region, whereas the Schwabische Alb region displayed a five times weaker positive effect (0.06) only. Further, fertilization effects on species richness were positive (0.03) for the region with organic soils compared to up to 25 times stronger negative effects in the other two regions. Synthesis. Our results clearly show the importance of studying both direct and indirect effects of land-use intensity. They demonstrate the indirect nature, via productivity, of the negative effect of fertilization intensity on plant species richness in the real-world context of management-induced gradients of intensity of fertilization, mowing and grazing. Finally, they highlight that careful consideration of regional environments is necessary before attempting to generalize land-use effects on species diversity.
The relationship of different types of grassland use with plant species richness and composition ( functional groups of herbs, legumes, and grasses) has so far been studied at small regional scales or comprising only few components of land use. We comprehensively studied the relationship between abandonment, fertilization, mowing intensity, and grazing by different livestock types on plant diversity and composition of 1514 grassland sites in three regions in North-East, Central and South-West Germany. We further considered environmental site conditions including soil type and topographical situation. Fertilized grasslands showed clearly reduced plant species diversity (-15% plant species richness, -0.1 Shannon diversity on fertilized grasslands plots of 16m(2)) and changed composition (-3% proportion of herb species), grazing had the second largest effects and mowing the smallest ones. Among the grazed sites, the ones grazed by sheep had higher than average species richness (+27%), and the cattle grazed ones lower (-42%). Further, these general results were strongly modulated by interactions between the different components of land use and by regional context: land-use effects differed largely in size and sometimes even in direction between regions. This highlights the importance of comparing different regions and to involve a large number of plots
Intensive land use is a major cause of biodiversity loss, but most studies comparing the response of multiple taxa rely on simple diversity measures while analyses of other community attributes are only recently gaining attention. Species-abundance distributions (SADs) are a community attribute that can be used to study changes in the overall abundance structure of species groups, and whether these changes are driven by abundant or rare species. We evaluated the effect of grassland management intensity for three land-use modes (fertilization, mowing, grazing) and their combination on species richness and SADs for three belowground (arbuscular mycorrhizal fungi, prokaryotes and insect larvae) and seven aboveground groups (vascular plants, bryophytes and lichens; arthropod herbivores; arthropod pollinators; bats and birds). Three descriptors of SADs were evaluated: general shape (abundance decay rate), proportion of rare species (rarity) and proportional abundance of the commonest species (dominance). Across groups, taxonomic richness was largely unaffected by land-use intensity and only decreased with increasing mowing intensity. Of the three SAD descriptors, abundance decay rate became steeper with increasing combined land-use intensity across groups. This reflected a decrease in rarity among plants, herbivores and vertebrates. Effects of fertilization on the three descriptors were similar to the combined land-use intensity effects. Mowing intensity only affected the SAD descriptors of insect larvae and vertebrates, while grazing intensity produced a range of effects on different descriptors in distinct groups. Overall, belowground groups had more even abundance distribtitions than aboveground groups. Strong differences among aboveground groups and between above- and belowground groups indicate that no single taxonomic group can serve as an indicator for effects in other groups. In the past, the use of SADs has been hampered by concerns over theoretical models underlying specific forms of SADs. Our study shows that SAD descriptors that are not connected to a particular model are suitable to assess the effect of land use on community structure.
Intensive land use is a driving force for biodiversity decline in many ecosystems. In semi-natural grasslands, land-use activities such as mowing, grazing and fertilization affect the diversity of plants and arthropods, but the combined effects of different drivers and the chain of effects are largely unknown. In this study we used structural equation modelling to analyse how the arthropod communities in managed grasslands respond to land use and whether these responses are mediated through changes in resource diversity or resource quantity (biomass). Plants were considered resources for herbivores which themselves were considered resources for predators. Plant and arthropod (herbivores and predators) communities were sampled on 141 meadows, pastures and mown pastures within three regions in Germany in 2008 and 2009. Increasing land-use intensity generally increased plant biomass and decreased plant diversity, mainly through increasing fertilization. Herbivore diversity decreased together with plant diversity but showed no response to changes in plant biomass. Hence, land-use effects on herbivore diversity were mediated through resource diversity rather than quantity. Land-use effects on predator diversity were mediated by both herbivore diversity (resource diversity) and herbivore quantity (herbivore biomass), but indirect effects through resource quantity were stronger. Our findings highlight the importance of assessing both direct and indirect effects of land-use intensity and mode on different trophic levels. In addition to the overall effects, there were subtle differences between the different regions, pointing to the importance of regional land-use specificities. Our study underlines the commonly observed strong effect of grassland land use on biodiversity. It also highlights that mechanistic approaches help us to understand how different land-use modes affect biodiversity.
Many bryophytes may depend on epizoochorous dispersal by large mammals for maintaining their populations and colonizing new habitats. On dry grassland we investigated the transport of bryophyte diaspores by sheep, which are known to be effective epizoochorous dispersal vectors of seed plants. Twelve sheep of two breeds were examined by collecting gametophyte fragments from their fleece and hooves. Furthermore, microscopic diaspores from the soil adhering to the hooves were grown in a greenhouse. Species frequency and composition were compared to bryophyte vegetation on the pasture. Considerable amounts of unspecialized gametophyte fragments from 16 moss species, 40% of the species growing on the pasture, were transported by sheep, especially on the belly and the tail. Additionally, we provide first evidence of epizoochorous dispersal of thallus fragments from fruticose lichens. Sheep breeds differed in species spectrum and frequency, i.e. sheep with dense, curly fleece carried more fragments and larger species than those with smooth and fine hair. Among the dispersed bryophytes pleurocarps, certain taxa, sizes (small species) and life forms (mats) were overrepresented compared to the vegetation of the pasture, while large species, acrocarps, wefts and turfs were underrepresented, reflecting fragmentation and adhesion features of the species. In the hooves mostly acrocarpous colonist species were transported. We conclude that sheep are important dispersal vectors with the potential of long-range dispersal for bryophytes, especially for species without sexual reproduction, and that epizoochorous dispersal of microscopic diaspores might be underestimated in its importance so far.
The Middle Spotted Woodpecker (Dendrocopos medius) is the bird species which Germany has the greatest global responsibility to protect. It is an umbrella species for the entire assemblage of animals associated with mature broadleaved trees, especially oak. Even though well studied in small to medium scale stands, the validity of habitat suitability analysis for this species in larger forests has not previously been proved. Aim of this study was to test suitability of permanent forest inventory plots for modelling its distribution in a 17,000 ha forest landscape and to derive habitat threshold values as a basis for formulating management guidelines. Based on 150 randomly selected 12.5 ha plots we identified mean age and basal area of oaks as the most important habitat factors using a backward selection logistic model. Internal validation showed an AUC of 0.89 and a R-2(N) of 0.58. Determination of thresholds using maximally selected rank statistics found higher probability of occurrence in stands with a mean age >95 years. Above that age the probability increased again in stands with more than 6.4 m(2) basal area oak/ha. Our results show that widely available forest inventory data can serve as a valuable basis for monitoring the Middle Spotted Woodpecker, either within the framework of the Natura 2000 Network, or more generally in integrated forest management with the aim of providing suitable habitats for the entire assemblage of species on old deciduous trees, especially oak.
Impact of Land-Use intensity and productivity on bryophyte diversity in agricultural grasslands
(2012)
While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m(2)) and central Germany (6.4 species per 16 m(2)) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.
Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands
(2014)
Disturbance is supposed to play an important role for biodiversity and ecosystem stability as described by the intermediate disturbance hypothesis (IDH), which predicts highest species richness at intermediate levels of disturbances. In this study, we tested the effects of artificial soil disturbances on diversity of annual and perennial vascular plants and bryophytes in a field experiment in 86 agricultural grasslands differing in land use in two regions of Germany. On each grassland, we implemented four treatments: three treatments differing in application time of soil disturbances and one control. One year after experimental disturbance, we recorded vegetation and measured biomass productivity and bare ground. We analysed the disturbance response taking effects of region and land-use-accompanied disturbance regimes into account.
Region and land-use type strongly determined plant species richness. Experimental disturbances had small positive effects on the species richness of annuals, but none on perennials or bryophytes. Bare ground was positively related to species richness of bryophytes. However, exceeding the creation of 12% bare ground further disturbance had a detrimental effect on bryophyte species richness, which corresponds to the IDH. As biomass productivity was unaffected by disturbance our results indicate that the disturbance effect on species richness of annuals was not due to decreased overall productivity, but rather due to short-term lowered inter- and intraspecific competition at the newly created microsites.
Generally, our results highlight the importance of soil disturbances for species richness of annual plants and bryophytes in agricultural grasslands. However, most grasslands were disturbed naturally or by land-use practices and our additional experimental soil disturbances only had a small short-term effect. Overall, total plant diversity in grasslands seemed to be more limited by the availability of propagules rather than by suitable microsites for germination. Thus, nature conservation efforts to increase grassland diversity should focus on overcoming propagule limitation, for instance by additional sowing of seeds, while the creation of additional open patches by disturbance might only be appropriate where natural disturbances are scarce.
We studied the effect of three major forest management types (unmanaged beech, selection beech, and age class forests) and stand variables (SMId, soil pH, proportion of conifers, litter cover, deadwood cover, rock cover and cumulative cover of woody trees and shrubs) on bryophyte species richness in 1050 forest plots in three regions in Germany. In addition, we analysed the species richness of four ecological guilds of bryophytes according to their colonized substrates (deadwood, rock, soil, bark) and the number of woodland indicator bryophyte species. Beech selection forests turned out to be the most species rich management type, whereas unmanaged beech forests revealed even lower species numbers than age-class forests. Increasing conifer proportion increased bryophyte species richness but not the number of woodland indicator bryophyte species. The richness of the four ecological guilds mainly responded to the abundance of their respective substrate. We conclude that the permanent availability of suitable substrates is most important for bryophyte species richness in forests, which is not stringently linked to management type. Therefore, managed age-class forests and selection forests may even exceed unmanaged forests in bryophyte species richness due to higher substrate supply and therefore represent important habitats for bryophytes. Typical woodland indicator bryophytes and their species richness were negatively affected by SMId (management intensity) and therefore better indicate forest integrity than the species richness of all bryophytes. Nature conservation efforts should focus on the reduction of management intensity. Moreover, maintaining and increasing a variability of substrates and habitats, such as coarse woody debris, increasing structural heterogeneity by retaining patches with groups of old, mature to over-mature trees in managed forests, maintaining forest climate conditions by silvicultural methods that assure stand continuity, e.g. by selection cutting rather than clear cutting and shelterwood logging might promote bryophyte diversity and in particular the one of woodland indicator bryophytes.
Epixylic bryophytes are important components of forest vegetation but are currently endangered by increment of wood harvest and intensive forest management. In this paper we present a study about the relationship between forest management, deadwood abundance, deadwood attributes and species richness of epixylic bryophytes on 30 plots comprising three forest types (managed coniferous, managed deciduous and unmanaged deciduous forests) in three regions in Germany. Additionally we analyzed the relations between deadwood attributes (wood species, decay, deadwood type, size) and bryophytes on deadwood items (n = 799) and calculated species interaction networks of wood species and bryophytes. Overall, species richness of epixylic bryophytes was positively related to deadwood abundance and diversity. The mean deadwood abundance was lowest in unmanaged forests (9.7 m(3) ha(-1)) compared with 15.0 m(3) ha(-1) in managed deciduous and 25.1 m(3) ha(-1) in managed coniferous forests. Accordingly, epixylic bryophyte species richness per plot increased from 7 species per 400 m(2) in unmanaged, 10 in managed deciduous and 16 in managed coniferous forests. The interaction network provided evidence of importance of tree-species diversity for bryophyte diversity and the relevance of particular wood species for rare bryophytes.
Generally, the results demonstrate a considerable lack of deadwood in all forest types, even in unmanaged forests. Species richness of epixylic bryophytes was strongly limited by available substrates within the observed deadwood abundance ranging up to only 60 m(3) ha(-1). Altogether, this suggests a high demand to increase both abundance and diversity of deadwood in forests.
The Hazel Grouse Bonasa bonasia is strongly affected by forest dynamics, and populations in many areas within Europe are declining. As a result of the 'wilding' concept implemented in the National Park Bavarian Forest, this area is one of the refuges for the species in Germany. Even though the effects of prevailing processes make the situation there particularly interesting, no recent investigation about habitat selection in the rapidly changing environment of the national park has been undertaken. We modelled the species-habitat relationship to derive the important habitat features in the national park as well as factors and critical threshold for monitoring, and to evaluate the predictive power of models based on field surveys compared to an analysis of infrared aerial photographs. We conducted our surveys on 49 plots of 25 ha each where Hazel Grouse was recorded and on an equally sized set of plots with no grouse occurrence, and used this dataset to build a predictive habitat-suitability model using logistic regression with backward stepwise variable selection. Habitat heterogeneity, stand structure, presence of mountain ash and willow, root plates, forest aisles, and young broadleaf stands proved to be predictive habitat variables. After internal validation via bootstrapping, our model shows an AUC value of 0.91 and a correct classification rate of 87%. Considering the methodological difficulties attached to backward selection, we applied Bayesian model averaging as an alternative. This multi-model approach also yielded similar results. To derive simple thresholds for important predictors as a basis for management decisions, we alternatively ran tree-based modelling, which also leads to a very similar selection of predictors. Performance of our different survey approaches was assessed by comparing two independent models with a model including both data resources: one constructed only from field survey data, the other based on data derived from aerial photographs. Models based on field data seem to perform slightly better than those based on aerial photography, but models using both predictor datasets provided the highest predictive accuracy.
Although genetic diversity is one of the key components of biodiversity, its drivers are still not fully understood. While it is known that genetic diversity is affected both by environmental parameters as well as habitat history, these factors are not often tested together. Therefore, we analyzed 14 microsatellite loci in Abax parallelepipedus, a flightless, forest dwelling ground beetle, from 88 plots in two study regions in Germany. We modeled the effects of historical and environmental variables on allelic richness, and found for one of the regions, the Schorfheide-Chorin, a significant effect of the depth of the litter layer, which is a main component of habitat quality, and of the sampling effort, which serves as an inverse proxy for local population size. For the other region, the Schwabische Alb, none of the potential drivers showed a significant effect on allelic richness. We conclude that the genetic diversity in our study species is being driven by current local population sizes via environmental variables and not by historical processes in the studied regions. This is also supported by lack of genetic differentiation between local populations sampled from ancient and from recent woodlands. We suggest that the potential effects of former fragmentation and recolonization processes have been mitigated by the large and stable local populations of Abax parallelepipedus in combination with the proximity of the ancient and recent woodlands in the studied landscapes.