Jasmin Mantilla-Contreras

Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.

Conservation value, management and restoration of Europe's semi-natural open landscapes

Abstract Most semi-natural open habitats in Europe have been traditionally maintained by anthropogenic activities, such as grazing or mowing, preventing the establishment of woody vegetation. These habitats harbour a remarkably rich biodiversity in terms of both plant and animal species, but are also highly threatened, mainly by agricultural intensification and land abandonment. With this Editorial we introduce a Special Issue initiated by the European Dry Grassland Group (EDGG) at the Open Landscapes Conference (Hildesheim, 2013) and the 11th European Dry Grassland Meeting (Kulikovo Pole, 2014). We aim to give a short introduction to the current conservation status, significance and research of semi-natural open habitats in Europe and present the collected articles of the Special Issue. These papers cover a wide range of different semi-natural open habitats, including wood-pastures, heathlands, steppes, semi-dry and dry grasslands across the Palearctic region and address issues related to the assessment methods, threats, management and restoration of these habitats. We conclude that, in order to ensure their conservation and to monitor the changes in open habitats, integrative approaches are needed that take into account not only vegetation records, but also multiple animal taxa, abiotic factors, management practices, ecosystem services and modelling simulations for anticipating possible future scenarios. We also recommend that decision-makers should support actions to conserve open habitats in Europe by addressing such major challenges as the encroachment of woody vegetation. We are convinced that the present Special Issue will contribute to a better understanding of ecosystem functions and support the biodiversity conservation and management of semi-natural open habitats. Izvleček Večino pol naravnih odprtih habitatov v Evropi tradicionalno vzdržuje s svojim delovanjem človek, kot na primer s pašo ali košnjo, in tako preprečuje zaraščanje z lesnato vegetacijo. Ti habitati so biodiverzitetno zelo bogati z rastlinskimi in živalskimi vrstami, vendar tudi močno ogroženi zaradi intenzifikacije kmetijstva in opuščanja obdelave. V uvodu želimo predstaviti posebno številko revije, ki jo je vzpodbudila Evropska skupina za suha travišča (EDGG) na konferenci Open Landscapes (Hildesheim, 2013) in na 11. srečanju skupine (Kulikovo Pole, 2014). Predstaviti želimo trenutni varstveni status, pomen in raziskave pol naravnih odprtih habitatov v Evropi in zbrane članke v posebni številki. Ti članki obravnavajo širok nabor različnih pol naravnih odprtih habitatov, med njimi gozdne pašnike, resave, stepe, pol suha in suha travišča palearktične regije in obravnavajo metode njihove presoje, njihovo ogroženost, gospodarjenje in obnovo. Zaključimo lahko, da moramo poleg vegetacijskih popisov upoštevati tudi številne živalske vrste, abiotske dejavnike, načine gospodarjenja, ekosistemske usluge in modeliranje predvidenih možnih prihodnjih scenarijev, če želimo zagotoviti njihovo ohranjanje in spremljanje sprememb. Predlagamo tudi, da nosilci odločanja podprejo prizadevanja za ohranjanje odprtih habitatov v Evropi z obravnavanjem tako velikih izzivov kot je zaraščanje z lesnato vegetacijo. Prepričani smo, da bo posebna številka prispevala k boljšemu razumevanju ekosistemskih funkcij in omogočila ohranjanje biodiverzitete in gospodarjenje spol naravnimi odprtimi habitati.

The influence of balanced and imbalanced resource supply on biodiversity–functioning relationship across ecosystems

Numerous studies show that increasing species richness leads to higher ecosystem productivity. This effect is often attributed to more efficient portioning of multiple resources in communities with higher numbers of competing species, indicating the role of resource supply and stoichiometry for biodiversity–ecosystem functioning relationships. Here, we merged theory on ecological stoichiometry with a framework of biodiversity–ecosystem functioning to understand how resource use transfers into primary production. We applied a structural equation model to define patterns of diversity–productivity relationships with respect to available resources. Meta-analysis was used to summarize the findings across ecosystem types ranging from aquatic ecosystems to grasslands and forests. As hypothesized, resource supply increased realized productivity and richness, but we found significant differences between ecosystems and study types. Increased richness was associated with increased productivity, although this effect was not seen in experiments. More even communities had lower productivity, indicating that biomass production is often maintained by a few dominant species, and reduced dominance generally reduced ecosystem productivity. This synthesis, which integrates observational and experimental studies in a variety of ecosystems and geographical regions, exposes common patterns and differences in biodiversity–functioning relationships, and increases the mechanistic understanding of changes in ecosystems productivity.

Consistent drivers of plant biodiversity across managed ecosystems.

Ecosystems managed for production of biomass are often characterized by low biodiversity because management aims to optimize single ecosystem functions (i.e. yield) involving deliberate selection of species or cultivars. In consequence, considerable differences in observed plant species richness and productivity remain across systems, and the drivers of these differences have remained poorly resolved so far. In addition, it has remained unclear if species richness feeds back on ecosystem functions such as yield in real-world systems. Here, we establish N = 360 experimental plots across a broad range of managed ecosystems in several European countries, and use structural equation models to unravel potential drivers of plant species richness. We hypothesize that the relationships between productivity, total biomass and observed species richness are affected by management intensity, and that these effects differ between habitat types (dry grasslands, grasslands, and wetlands). We found that local management was an important driver of species richness across systems. Management caused system disturbance, resulting in reduced productivity yet enhanced total biomass. Plant species richness was directly and positively driven by management, with consistently negative effects of total biomass. Productivity effects on richness were positive, negative or neutral. Our study shows that management and total biomass drive plant species richness across real-world managed systems.

Low-intensity Husbandry as a Cost-efficient Way to Preserve Dry Grasslands

Abstract Dry grasslands represent unique ecosystems that are known for high biodiversity. Land use changes have led to a decline in Europe. Shrub encroachment is a major problem in incorrectly or unmanaged areas, and affects not only flora and fauna but also leads to changes in the appearance of the landscape. Our study was conducted on the island of Hiddensee, north-east Germany, where today dry grasslands are endangered by shrub encroachment. We analysed the current land use under consideration of farming costs as well as current agricultural subsidies. Our results show that revenues of low-intensity cattle husbandry only account for 61% of total costs and that farming becomes profitable only with subsidies. Our study illustrates that goats can be a cost-efficient solution for areas with high shrub cover in contrast to manual clearing. The maintenance of dry grasslands is therefore more cost-efficient with grazing animals than with technical-manual labour.

Reading the Leaves’ Palm: Leaf Traits and Herbivory along the Microclimatic Gradient of Forest Layers

Microclimate in different positions on a host plant has strong direct effects on herbivores. But little is known about indirect effects due to changes of leaf traits. We hypothesized that herbivory increases from upper canopy to lower canopy and understory due to a combination of direct and indirect pathways. Furthermore, we hypothesized that herbivory in the understory differs between tree species in accordance with their leaf traits. We investigated herbivory by leaf chewing insects along the vertical gradient of mixed deciduous forest stands on the broad-leaved tree species Fagus sylvatica L. (European beech) with study sites located along a 140 km long transect. Additionally, we studied juvenile Acer pseudoplatanus L. (sycamore maple) and Carpinus betulus L. (hornbeam) individuals within the understory as a reference of leaf traits in the same microclimate. Lowest levels of herbivory were observed in upper canopies, where temperatures were highest. Temperature was the best predictor for insect herbivory across forest layers in our study. However, the direction was opposite to the generally known positive relationship. Herbivory also varied between the three tree species with lowest levels for F. sylvatica. Leaf carbon content was highest for F. sylvatica and probably indicates higher amounts of phenolic defense compounds. We conclude that the effect of temperature must have been indirect, whereby the expected higher herbivory was suppressed due to unfavorable leaf traits (lower nitrogen content, higher toughness and carbon content) of upper canopy leaves compared to the understory.

Environment vs. Plant Ontogeny: Arthropod Herbivory Patterns on European Beech Leaves along the Vertical Gradient of Temperate Forests in Central Germany

Environmental and leaf trait effects on herbivory are supposed to vary among different feeding guilds. Herbivores also show variability in their preferences for plant ontogenetic stages. Along the vertical forest gradient, environmental conditions change, and trees represent juvenile and adult individuals in the understorey and canopy, respectively. This study was conducted in ten forests sites in Central Germany for the enrichment of canopy research in temperate forests. Arthropod herbivory of different feeding traces was surveyed on leaves of Fagus sylvatica Linnaeus (European beech; Fagaceae) in three strata. Effects of microclimate, leaf traits, and plant ontogenetic stage were analyzed as determining parameters for herbivory. The highest herbivory was caused by exophagous feeding traces. Herbivore attack levels varied along the vertical forest gradient for most feeding traces with distinct patterns. If differences of herbivory levels were present, they only occurred between juvenile and adult F. sylvatica individuals, but not between the lower and upper canopy. In contrast, differences of microclimate and important leaf traits were present between the lower and upper canopy. In conclusion, the plant ontogenetic stage had a stronger effect on herbivory than microclimate or leaf traits along the vertical forest gradient.

Effects of dung-pad conditions and density on coprophagous beetle assemblages in a Mediterranean rangeland

Dung beetles highly depend on the ephemeral microhabitat dung which is food resource and larval habitat at the same time. Environmental conditions surrounding a dung pad, such as vegetation structure, have an impact on dung beetle assemblages. We investigated the influence of dung conditions and surrounding habitat characteristics on Mediterranean dung beetle assemblages in a permanently grazed landscape in northern Sardinia. We sampled the dung beetle assemblages of donkey and horse dung in three different vegetation types and assessed species richness and abundance of dung beetles. Species richness was determined by dung and surrounding habitat conditions, whereas abundance was solely affected by dung conditions. However, species richness and abundance decreased with increasing dung density. The effect of dung density on species richness varied depending on vegetation type, with dry grassland exhibiting the highest number of dung beetles species at high dung density. Species composition in dung pads was influenced by abiotic factors with dwellers being negatively affected by increasing dung-pad temperature. Our results underline the importance of diverse vegetation, particularly with respect to the complexity of vegetation which interrelates with the microclimate. Furthermore, our findings illustrate the negative effect of high dung densities on dung beetle assemblages, suggesting that the degree of the intensity of use by grazing animals is important when considering measures for the conservation of dung beetles.

Trait correlation network analysis identifies biomass allocation traits and stem specific length as hub traits in herbaceous perennial plants

1. Correlations among plant traits often reflect important trade‐offs or allometric relationships in biological functions like carbon gain, support, water uptake, and reproduction that are associated with different plant organs. Whether trait correlations can be aggregated to “spectra” or “leading dimensions,” whether these dimensions are consistent across plant organs, spatial scale, and growth forms are still open questions.2. To illustrate the current state of knowledge, we constructed a network of published trait correlations associated with the “leaf economics spectrum,” “biomass allocation dimension,” “seed dimension,” and carbon and nitrogen concentrations. This literature‐based network was compared to a network based on a dataset of 23 traits from 2,530 individuals of 126 plant species from 381 plots in Northwest Europe.3. The observed network comprised more significant correlations than the literature‐based network. Network centrality measures showed that size traits such as the mass of leaf, stem, below‐ground, and reproductive tissues and plant height were the most central traits in the network, confirming the importance of allometric relationships in herbaceous plants. Stem mass and stem‐specific length were “hub” traits correlated with most traits. Environmental selection of hub traits may affect the whole phenotype. In contrast to the literature‐based network, SLA and leaf N were of minor importance. Based on cluster analysis and subsequent PCAs of the resulting trait clusters, we found a “size” module, a “seed” module, two modules representing C and N concentrations in plant organs, and a “partitioning” module representing organ mass fractions. A module representing the plant economics spectrum did not emerge.4. Synthesis. Although we found support for several trait dimensions, the observed trait network deviated significantly from current knowledge, suggesting that previous studies have overlooked trait coordination at the whole‐plant level. Furthermore, network analysis suggests that stem traits have a stronger regulatory role in herbaceous plants than leaf traits.

Experimental Study of Environmental Effects: Leaf Traits of Juvenile Fagus sylvatica, Acer pseudoplatanus, and Carpinus betulus Are Comparable to Leaves of Mature Trees in Upper Canopies

Morphological and functional leaf traits like leaf toughness and nutrient content are essentially influenced by the environment, especially through light and climatic conditions. Varying light conditions have been identified as a significant predictor for the variation of many leaf traits. However, the leaf acclimation to light is suggested to be of secondary importance. The aim of the experimental study was to analyse environmental effects (microclimate and soil moisture), which are present in upper canopies of forest stands, on leaf traits of juvenile Fagus sylvatica L. (European beech; Fagaceae), Acer pseudoplatanus L. (sycamore maple; Sapindaceae), and Carpinus betulus L. (hornbeam; Betulaceae). The experimental design managed to imitate two distinct microclimates causing different temperature and air humidity conditions. Furthermore, the irrigation treatment with different levels of applied water caused distinct soil moisture conditions in the trial pots. As a result of the treatments, leaves of C. betulus showed a tendency of decreased specific leaf area (SLA) caused by the treatment with warmer and drier microclimate. The environmental effect on SLA was even stronger with lower soil moisture conditions. Chlorophyll content showed lower values in treatments with higher soil moisture conditions in both greenhouses for F. sylvatica and A. pseudoplatanus. The trends are in accordance with combined effects of temperature, air humidity, and soil moisture on SLA, and increased leaf chlorophyll content caused by slight drought stress. Plants in the greenhouses were exposed to full sunlight indicating a microclimatic environment comparable to upper canopies in forest stands. The comparable SLA and chlorophyll content between leaves of mature F. sylvatica trees in upper canopies and juvenile trees of the greenhouses suggest similar environmental conditions instead of ontogenetic effects that are responsible for the formation of leaf trait characteristics.