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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.
A major aim in ecology is identifying determinants of invasiveness. We performed a meta-analysis of 117 field or experimental-garden studies that measured pair-wise trait differences of a total of 125 invasive and 196 non-invasive plant species in the invasive range of the invasive species. We tested whether invasiveness is associated with performance-related traits (physiology, leaf-area allocation, shoot allocation, growth rate, size and fitness), and whether such associations depend on type of study and on biogeographical or biological factors. Overall, invasive species had significantly higher values than non-invasive species for all six trait categories. More trait differences were significant for invasive vs. native comparisons than for invasive vs. non-invasive alien comparisons. Moreover, for comparisons between invasive species and native species that themselves are invasive elsewhere, no trait differences were significant. Differences in physiology and growth rate were larger in tropical regions than in temperate regions. Trait differences did not depend on whether the invasive alien species originates from Europe, nor did they depend on the test environment. We conclude that invasive alien species had higher values for those traits related to performance than non-invasive species. This suggests that it might become possible to predict future plant invasions from species traits.
Local adaption of the clonal plant Ranunculus reptans to flooding along a small-scale gradient.
(2004)
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
The high potential fitness benefit of phenotypic plasticity tempts us to expect phenotypic plasticity as a frequent adaptation to environmental heterogeneity. Examples of proven adaptive plasticity in plants, however, are scarce and most plastic responses actually may be 'passive' rather than adaptive. This suggests that frequently requirements for the evolution of adaptive plasticity are not met or that such evolution is impeded by constraints. Here we outline requirements and potential constraints for the evolution of adaptive phenotypic plasticity, identify open questions, and propose new research approaches. Important open questions concern the genetic background of plasticity, genetic variation in plasticity, selection for plasticity in natural habitats, and the nature and occurrence of costs and limits of plasticity. Especially promising tools to address these questions are selection gradient analysis, meta-analysis of studies on genotype-by-environment interactions, QTL analysis, cDNA-microarray scanning and quantitative PCR to quantify gene expression, and two-dimensional gel electrophoresis to quantify protein expression. Studying plasticity along the pathway from gene expression to the phenotype and its relationship with fitness will help us to better understand why adaptive plasticity is not more universal, and to more realistically predict the evolution of plastic responses to environmental change
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.
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.
Species diversity promotes the delivery of multiple ecosystem functions (multifunctionality). However, the relative functional importance of rare and common species in driving the biodiversity multifunctionality relationship remains unknown. We studied the relationship between the diversity of rare and common species (according to their local abundances and across nine different trophic groups), and multifunctionality indices derived from 14 ecosystem functions on 150 grasslands across a land use intensity (LUI) gradient. The diversity of above- and below-ground rare species had opposite effects, with rare above-ground species being associated with high levels of multifunctionality, probably because their effects on different functions did not trade off against each other. Conversely, common species were only related to average, not high, levels of multifunctionality, and their functional effects declined with LUI. Apart from the community level effects of diversity, we found significant positive associations between the abundance of individual species and multifunctionality in 6% of the species tested. Species specific functional effects were best predicted by their response to LUI: species that declined in abundance with land use intensification were those associated with higher levels of multifunctionality. Our results highlight the importance of rare species for ecosystem multifunctionality and help guiding future conservation priorities.
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 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.
Supporting species persistence may involve (re)connecting suitable habitats. However, for many declining species habitat suitability and drivers of establishment are poorly known. We addressed this experimentally for a declining flagship species of dry grasslands in Germany, Armeria maritima subsp. elongata. In three regions, we sowed seeds from each of eight source populations back to their origin and to eight apparently suitable, but currently unoccupied, habitats close to the source populations. Overall, seeds germinated and seedlings established equally well in occupied and potential sites indicating that suitable habitats are available, but lack seed input. Germination and establishment varied among sowing sites. Moreover, seeds from populations of lower current connectivity established less well in new sites, and establishment was more variable among seeds from smaller than from larger populations, possibly reflecting genetic consequences of habitat fragmentation. Further, establishment across different new environments differed between seeds from different populations. As this was neither related to a home-away contrast nor to geographic or environmental distance between sites it could not clearly be attributed to local adaptation. To promote long-term persistence within this dry-grassland meta-population context we suggest increasing the density of suitable habitats and supporting dispersal connecting multiple sites, e.g. by promoting sheep transhumance, to increase current populations and their connectivity, and to colonise suitable habitats with material from different sources. We suggest that sowing experiments with characteristic species, including multiple source populations and multiple recipient sites, should be used regularly to inform connecting efforts in plant conservation.
Effects of plant community diversity on ecosystem processes have recently received major attention. In contrast, effects of species richness and functional richness on individual plant performance, and their magnitude relative to effects of community composition, have been largely neglected. Therefore, we examined height, aboveground biomass, and inflorescence production of individual plants of all species present in 82 large plots of the Jena Experiment, a large grassland biodiversity experiment in Germany. These plots differed in species richness (1-60), functional richness (1-4), and community composition. On average, in more species-rich communities, plant individuals grew taller, but weighed less, were less likely to flower, and had fewer inflorescences. In plots containing legumes, non-legumes were higher and weighed more than in plots without legumes. In plots containing grasses, non-grasses were less likely to flower than in plots without grasses. This indicates that legumes positively and grasses negatively affected the performance of other species. Species richness and functional richness effects differed systematically between functional groups. The magnitude of the increase in plant height with increasing species richness was greatest in grasses and was progressively smaller in legumes, small herbs, and tall herbs. Individual aboveground biomass responses to increasing species richness also differed among functional groups and were positive for legumes, less pronouncedly positive for grasses, negative for small herbs, and more pronouncedly negative for tall herbs. Moreover, these effects of species richness differed strongly between species within these functional groups. We conclude that individual plant performance largely depends on the diversity of the surrounding community, and that the direction and magnitude of the effects of species richness and functional richness differs largely between species. Our study suggests that diversity of the surrounding community needs to be taken into account when interpreting drivers of the performance of individual plants.
Understory herbs are an essential part of tropical rain forests, but little is known about factors limiting their reproduction. Many of these herbs are clonal, patchily distributed, and produce large floral displays of nectar-rich 1-d flowers to attract hummingbird pollinators that may transport pollen over long distances. The aim of this study was to investigate the effects of clonality, cross-proximity, and patchy distribution on the reproduction of the hummingbird-pollinated Amazonian herb Heliconia metallica. We experimentally pollinated flowers within populations with self-pollen and with pollen of different diversity, crossed flowers between populations, and added supplemental pollen to ramets growing solitarily or in conspecific patches. Only flowers pollinated early in the morning produced seeds. Selfed flowers produced seeds, but seed number and mass were strongly reduced, suggesting partial sterility and inbreeding depression after selfing. Because of pollen competition, flowers produced more seeds after crosses with several than with single donor plants. Crosses between populations mostly resulted in lower seed production than those within populations, suggesting outbreeding depression. Ramets in patches produced fewer seeds than solitary ramets and were more pollen-limited, possibly due to geitonogamy and biparental inbreeding in patches. We conclude that high rates of geitonogamy due to clonality and pollen limitation due to the short receptivity of flowers and patchy distribution constrain the reproduction of this clonal herb. Even in unfragmented rain forests with highly mobile pollinators, outbreeding depression may be a widespread phenomenon in plant reproduction.