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Assessing diversity is among the major tasks in ecology and conservation science. In ecological and conservation studies, epiphytic cryptogams are usually sampled up to accessible heights in forests. Thus, their diversity, especially of canopy specialists, likely is underestimated. If the proportion of those species differs among forest types, plot-based diversity assessments are biased and may result in misleading conservation recommendations. We sampled bryophytes and lichens in 30 forest plots of 20 m x 20 m in three German regions, considering all substrates, and including epiphytic litter fall. First, the sampling of epiphytic species was restricted to the lower 2 m of trees and shrubs. Then, on one representative tree per plot, we additionally recorded epiphytic species in the crown, using tree climbing techniques. Per tree, on average 54% of lichen and 20% of bryophyte species were overlooked if the crown was not been included. After sampling all substrates per plot, including the bark of all shrubs and trees, still 38% of the lichen and 4% of the bryophyte species were overlooked if the tree crown of the sampled tree was not included. The number of overlooked lichen species varied strongly among regions. Furthermore, the number of overlooked bryophyte and lichen species per plot was higher in European beech than in coniferous stands and increased with increasing diameter at breast height of the sampled tree. Thus, our results indicate a bias of comparative studies which might have led to misleading conservation recommendations of plot-based diversity assessments.
We tested whether neighborhood density affects the clonal life history of the stoloniferous plant Ranunculus reptans through selection and genetic drift. After three generations of sexual reproduction of 16 low- and 16 high- density lines, we studied traits related to growth form and reproduction in a common competition free environment. A 7.7% lower branching frequency and slightly longer internodes indicated an evolutionary shift towards a less compact growth form under high neighborhood density, but because stolons grew also more vertically, horizontal spread per ramet was slightly decreased. Neighborhood density had no directional effects on the evolution of allocation to sexual and vegetative reproduction in R. reptans. Variation among replicated high-density lines was significantly lower than among replicated low-density lines in both growth form and reproductive characteristics, indicating less pronounced genetic drift under high neighborhood density. This study demonstrates that a clonal plant can respond to selection imposed by neighborhood density. Moreover, it shows that the effect of random genetic drift increases with decreasing neighborhood density. In a declining species, such as R. reptans in central Europe, this may lower the potential for adaptive evolutionary change and increase extinction risk
Many plant populations are adapted to their local environment. Reciprocal transplant experiments in the field and in the experimental garden allow for studying different aspects of local adaptation. However, usually only one of these approaches is used. We applied both experimental approaches to study the role of spatial scale and soil conditions for local adaptation in the perennial herb Inula hirta. We reciprocally sowed seeds and transplanted juvenile plants among six field sites from two regions and, in the garden, among pots with soil from each field site. We recorded germination percentage, survival percentage, number of stems and plant height in all experiments. We also recorded above- and below-ground biomass, flowering percentage and the number of flower heads in the garden. No population-specific local adaptation was detected in germination, survival, flowering percentages or in the number of flower heads. At the regional scale in the field, however, the performance of local transplants was higher than the performance of foreign transplants by 10% and 7% in the two regions, respectively. Similarly, when grown in the garden in soil from the more basic and nutrient-poorer region, plant height and aboveground biomass of local transplants were higher than the corresponding values for foreign transplants by 31% and 112%, respectively. Congruent evidence for local adaptation from the juvenile-transplant experiments in field and garden suggests that soil conditions represent an important factor of local adaptation in I. hirta. Overall, our results show that both spatial scale and soil conditions play an important role for local adaptation in I. hirta. Moreover, we underline the importance of combining field and garden experiments to reveal factors affecting local adaptation in plants.
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.
1. For managed temperate forests, conservationists and policymakers favour fine-grained uneven-aged (UEA) management over more traditional coarse-grained even-aged (EA) management, based on the assumption that within-stand habitat heterogeneity enhances biodiversity. There is, however, little empirical evidence to support this assumption. We investigated for the first time how differently grained forest management systems affect the biodiversity of multiple above- and below-ground taxa across spatial scales. 2. We sampled 15 taxa of animals, plants, fungi and bacteria within the largest contiguous beech forest landscape of Germany and classified them into functional groups. Selected forest stands have been managed for more than a century at different spatial grains. The EA (coarse-grained management) and UEA (fine-grained) forests are comparable in spatial arrangement, climate and soil conditions. These were compared to forests of a nearby national park that have been unmanaged for at least 20years. We used diversity accumulation curves to compare -diversity for Hill numbers D-0 (species richness), D-1 (Shannon diversity) and D-2 (Simpson diversity) between the management systems. Beta diversity was quantified as multiple-site dissimilarity. 3. Gamma diversity was higher in EA than in UEA forests for at least one of the three Hill numbers for six taxa (up to 77%), while eight showed no difference. Only bacteria showed the opposite pattern. Higher -diversity in EA forests was also found for forest specialists and saproxylic beetles. 4. Between-stand -diversity was higher in EA than in UEA forests for one-third (all species) and half (forest specialists) of all taxa, driven by environmental heterogeneity between age-classes, while -diversity showed no directional response across taxa or for forest specialists. 5. Synthesis and applications. Comparing EA and uneven-aged forest management in Central European beech forests, our results show that a mosaic of different age-classes is more important for regional biodiversity than high within-stand heterogeneity. We suggest reconsidering the current trend of replacing even-aged management in temperate forests. Instead, the variability of stages and stand structures should be increased to promote landscape-scale biodiversity.
Specialisation and diversity of multiple trophic groups are promoted by different forest features
(2019)
While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above- and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation.
Edaphic fauna contributes to important ecosystem functions in grassland soils such as decomposition and nutrient mineralization. Since this functional role is likely to be altered by global change and associated shifts in plant communities, a thorough understanding of large scale drivers on below-ground processes independent of regional differences in soil type or climate is essential. We investigated the relationship between abiotic (soil properties, management practices) and biotic (plant functional group composition, vegetation characteristics, soil fauna abundance) predictors and feeding activity of soil fauna after accounting for sample year and study region. Our study was carried out over a period of two consecutive years in 92 agricultural grasslands in three regions of Germany, spanning a latitudinal gradient of more than 500 km. A structural equation model suggests that feeding activity of soil fauna as measured by the bait-lamina test was positively related to legume and grass species richness in both years. Most probably, a diverse vegetation promotes feeding activity of soil fauna via alterations of both microclimate and resource availability. Feeding activity of soil fauna also increased with earthworm biomass via a pathway over Collembola abundance. The effect of earthworms on the feeding activity in soil may be attributed to their important role as ecosystem engineers. As no additional effects of agricultural management such as fertilization, livestock density or number of cuts on bait consumption were observed, our results suggest that the positive effect of legume and grass species richness on the feeding activity in soil fauna is a general one that will not be overruled by regional differences in management or environmental conditions. We thus suggest that agri-environment schemes aiming at the protection of belowground activity and associated ecosystem functions in temperate grasslands may generally focus on maintaining plant diversity, especially with regard to the potential effects of climate change on future vegetation 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.
1. Plant species persistence in natural communities requires coping with biotic and abiotic challenges. These challenges also depend on plant community composition and diversity. Over time, biodiversity effects have been shown to be strengthened via increasing species complementarity in mixtures. Little is known, however, whether differences in community diversity and composition induce rapid transgenerational phenotypic adaptive differentiation during community assembly. We expect altered plant-plant and other biotic interactions (mutualists or antagonists) in high vs. low diverse communities to affect immediate within-and between-species trait differentiations due to competition for light and nutrients. 2. Three years after the initiation of a large-scale, long-term biodiversity experiment in Jena, Germany, we tested for effects of varying experimental plant community diversity (1-60 plant species; one to four plant functional groups) and composition (with or without legumes and/or grasses) on phenotypic differentiation and variation of the tall herb Knautia arvensis. We measured reproduction at different diversity levels in the Jena Experiment (residents hereafter) and, in an additional common garden experiment without competition, recorded subsequent offspring performance (i.e. growth, reproductive success and susceptibility to powdery mildew) to test for differentiation in phenotypic expression and variability. 3. We observed phenotypic differences among diversity levels with reduced fecundity of K. arvensis residents in more diverse communities. In the next generation grown under common garden conditions, offspring from high-diversity plots showed reduced growth (i.e. height) and lower reproduction (i.e. fewer infructescences), but increased phenotypic trait variability (e.g. in leaf width and powdery mildew presence) and also tended to be less susceptible to powdery mildew infection. 4. Community composition also affected Knautia parents and offspring. In the presence of legumes, resident plants produced more seeds (increased fecundity); however, germination rate of those seeds was reduced at an early seedling stage (reduced fertility). 5. Synthesis. We conclude that rapid transgenerational effects of community diversity and composition on both mean and variation of phenotypic traits among offspring exist. In addition to heritable variation, environmentally induced epigenetic and/or maternal processes matter for early plant community assembly and may also determine future species coexistence and community stability.