Distance functions of carabids in crop fields depend on functional traits, crop type and adjacent habitat: a synthesis.

Natural pest and weed regulation are essential for agricultural production, but the spatial distribution of natural enemies within crop fields and its drivers are mostly unknown. Using 28 datasets comprising 1204 study sites across eight Western and Central European countries, we performed a quantitative synthesis of carabid richness, activity densities and functional traits in relation to field edges (i.e. distance functions). We show that distance functions of carabids strongly depend on carabid functional traits, crop type and, to a lesser extent, adjacent non-crop habitats. Richness of both carnivores and granivores, and activity densities of small and granivorous species decreased towards field interiors, whereas the densities of large species increased. We found strong distance decays in maize and vegetables whereas richness and densities remained more stable in cereals, oilseed crops and legumes. We conclude that carabid assemblages in agricultural landscapes are driven by the complex interplay of crop types, adjacent non-crop habitats and further landscape parameters with great potential for targeted agroecological management. In particular, our synthesis indicates that a higher edge-interior ratio can counter the distance decay of carabid richness per field and thus likely benefits natural pest and weed regulation, hence contributing to agricultural sustainability.

Copper Uptake and Its Effects on Two Riparian Plant Species, the Native Urtica dioica, and the Invasive Fallopia japonica.

Copper accumulating in stream sediments can be transported to adjacent riparian habitats by flooding. Although being an essential element for plants, copper is toxic at high concentrations and restricts, among other things, plant growth. Besides copper, invasive plants, such as Fallopia japonica, which are known to be tolerant toward heavy metals, modify riparian habitats. If the tolerance of F. japonica is higher compared to native plants, this could accelerate invasion under high heavy metal stress. Therefore, we aimed to compare the effect of copper on two common riparian plants, the invasive F. japonica and the native Urtica dioica. We performed a pot experiment with a gradient from 0 to 2430 mg kg-1 of soil copper. We hypothesized that (i) negative effects on plant growth increase with increasing soil copper concentrations with F. japonica being less affected and (ii) accumulating higher amounts of copper in plant tissues compared to U. dioica. In support of our first hypothesis, growth (height, leaf number) and biomass (above- and belowground) of F. japonica were impacted at the 810 mg kg-1 treatment, while the growth of U. dioica was already impacted at 270 mg kg-1. Due to 100% mortality of plants, the 2430 mg kg-1 treatment was omitted from the analysis. In contrast, chlorophyll content slightly increased with increasing copper treatment for both species. While U. dioica accumulated more copper in total, the copper uptake by F. japonica increased more strongly after exposure compared to the control. In the 810 mg kg-1 treatment, copper concentrations in F. japonica were up to 2238% higher than in the control but only up to 634% higher in U. dioica. Our results indicate that F. japonica might be able to more efficiently detoxify internal copper concentrations controlling heavy metal effects compared to the native species. This could give F. japonica a competitive advantage particularly in polluted areas, facilitating its invasion success.

Urbanization increases fluctuating asymmetry and affects behavioral traits of a common grasshopper.

Urbanization has a major impact on biodiversity. For many organisms, the urbanization process means environmental stress caused by fragmentation and increased temperatures in cities and atmospheric, soil, light, and noise pollution. Such environmental stress can influence both the morphology and behavior of animals. Hence, individuals might be selected for survival-facilitating traits under high pressures in urban areas. The specific impact of urbanization on insect behavior is still largely unexplored. We studied the impact of urbanization on one of the most common grasshopper species in Germany, Chorthippus biguttulus, by comparing morphological and behavioral traits of individuals sampled from grasslands with low, medium, and high urbanization levels. We first investigated whether urbanization as a stressor affected body size and fluctuating asymmetry in the locomotor organs. Next, we examined whether urbanization induced changes in the individuals' boldness and activity. Our results showed that fluctuating asymmetry of grasshoppers' locomotory organs increased more than twofold with urbanization level. Further, individuals' boldness and walking activity increased from areas with low to high urbanization levels. Our results indicate strong responses of grasshoppers in terms of morphology and behavior to the urban environment. To compensate for urbanization effects on arthropod populations, management strategies need to be developed that maintain ecological processes and reduce environmental stress in urban areas.

Native pedunculate oaks support more biodiversity than non-native oaks, but non-native oaks are healthier than native oaks: A study on street and park trees of a city.

Trees in cities provide multiple ecosystem services. However, simultaneously ensuring healthy trees with high habitat diversity can be challenging in a harsh urban environment. We compared health, microhabitats, and bat activities between native (Quercus robur L.) and non-native (Quercus rubra L.) oaks growing in different urban habitats (street vs. park) in Karlsruhe, southwestern Germany. We randomly selected 167 oak trees with a diameter at breast height (DBH) >20 cm across the city from Urban Tree Registrar. We performed tree health assessment, dendrometric, and microhabitat inventory. We recorded the four-day bat activities on 45 native and non-native oaks with acoustic loggers installed on the trees. We found that non-native oaks were healthier than native oaks but provided less abundance and richness of microhabitats. Tree size (positive effect) and pruning (negative effect) strongly influence microhabitat richness and abundance. In addition, park trees hosted significantly more microhabitats than street trees. We recorded the activities of 9 bat species from 4 genera. Pipistrellus bats were more active in park trees than street trees. Long-eared bats (Plecotus) were more active near the native than non-native oaks. Bats are likely favored by microhabitats such as fork split, lightning scar, and woodpecker "flute" that are more common in less healthy trees. We conclude that non-native red oak can be planted alongside streets, where the conditions are harsher than in parks to better adapt to climatic changes and stay healthy with less maintenance. The preservation of native pedunculate oak trees, especially within parks, is paramount for urban biodiversity conservation because of their potential to provide microhabitats and supporting bats.

Structural and functional development of twelve newly established floodplain pond mesocosms.

Ecosystems are complex structures with interacting abiotic and biotic processes evolving with ongoing succession. However, limited knowledge exists on the very initial phase of ecosystem development and colonization. Here, we report results of a comprehensive ecosystem development monitoring for twelve floodplain pond mesocosms (FPM; 23.5 m × 7.5 m × 1.5 m each) located in south-western Germany. In total, 20 abiotic and biotic parameters, including structural and functional variables, were monitored for 21 months after establishment of the FPMs. The results showed evolving ecosystem development and primary succession in all FPMs, with fluctuating abiotic conditions over time. Principal component analyses and redundancy analyses revealed season and succession time (i.e., time since ecosystem establishment) to be significant drivers of changes in environmental conditions. Initial colonization of both aquatic (i.e., water bodies) and terrestrial (i.e., riparian land areas) parts of the pond ecosystems occurred within the first month, with subsequent season-specific increases in richness and abundance for aquatic and terrestrial taxa over the entire study period. Abiotic environmental conditions and aquatic and terrestrial communities showed increasing interpond variations over time, that is, increasing heterogeneity among the FPMs due to natural environmental divergence. However, both functional variables assessed (i.e., aquatic and terrestrial litter decomposition) showed opposite patterns as litter decomposition rates slightly decreased over time and interpond differences converged with successional ecosystem developments. Overall, our results provide rare insights into the abiotic and biotic conditions and processes during the initial stages of freshwater ecosystem formation, as well as into structural and functional developments of the aquatic and terrestrial environment of newly established pond ecosystems.

Disentangling the role of management, vegetation structure, and plant quality for Orthoptera in lowland meadows.

Seminatural grasslands provide habitats for various species and are important for biodiversity conservation. The understanding of the diverse responses of species and traits to different grassland management methods is therefore urgently needed. We disentangled the role of grassland management (fertilization and irrigation), vegetation structure (biomass, sward height) and plant quality (protein and fiber content) for Orthoptera communities in lowland hay meadows in Germany. We found vegetation structure to be the most important environmental category in explaining community structure of Orthoptera (species richness, total individuals, functional diversity and species composition). Intensively used meadows (fertilized, irrigated, high plant biomass) were characterized by assemblages with few species, low functional diversity, and low conservation value. Thereby, the relatively moderate fertilizer inputs in our study system of up to ∼75 kg N/ha/year reduced functional diversity of Orthoptera, while this negative effect of fertilization was not detectable when solely considering taxonomic aspects. We found strong support for a prominent role of plant quality in shaping Orthoptera communities and especially the trait composition. Our findings demonstrate the usefulness of considering both taxonomic and functional components (functional diversity) in biodiversity research and we suggest a stronger involvement of plant quality measures in Orthoptera studies.

Dominance of cropland reduces the pollen deposition from bumble bees.

Intensive agricultural landscapes can be hostile for bees due to a lack of floral and nesting resources, and due to management-related stress such as pesticide use and soil tillage. This threatens the pollination services that bees deliver to insect-pollinated crops. We studied the effects of farming intensity (organic vs. conventional, number of insecticide applications) and availability of semi-natural habitats at the field and landscape scale on pollinator visits and pollen delivery to pumpkin in Germany. We found that wild bumble bees were the key pollinators of pumpkin in terms of pollen delivery, despite fivefold higher visitation frequency of honey bees. Critically, we observed that the area of cropland had stronger effects on bees' pollen deposition than the area of seminatural habitats. Specifically, a 10% increase of the proportion of cropland reduced pollen delivery by 7%. Pumpkin provides a striking example for a key role of wild pollinators in crop pollination even at high numerical dominance of honey bees. In addition, our findings suggest that habitat conversion to agricultural land is a driver of deteriorating pollination. This underlines the importance to maintain sufficient areas of non-crop habitats in agricultural landscapes.

Effectiveness of corridors varies among phytosociological plant groups and dispersal syndromes.

In agricultural landscapes, semi-natural habitats are scarce and remaining habitat patches are largely isolated. However, linear landscape elements might facilitate dispersal of plant species through the agricultural landscape matrix. We investigated the following research questions: 1. are open linear landscape elements (LLE) effective corridors for dispersal of vascular plant species? 2. Which plant species, with respect to phytosociological group and dispersal-distance class, do use LLE as corridors? 3. To which extent is floristic similarity of communities influenced by dispersal through corridors? Field work was carried out in agricultural landscapes of Northwest Germany. We sampled 50 vegetation relevés on open linear landscape elements i.e. field margins (incl. road verges) and ditches, in eight 1-km2 study areas. Then, we calculated Jaccard similarities of all plot pairs within study areas using either all species or only species of certain phytosociological groups or dispersal-distance classes. We assessed the isolation of the plots from each other using both Euclidean distance and resistance distance along LLE. Resistance distance reflected the degree of connectivity of the LLE network between the plots. A stronger effect on Jaccard similarity of resistance distance compared to Euclidean distance would indicate corridor dispersal of plants through LLE. Relationships between Jaccard similarity and the two isolation measures were analysed with Generalised Linear Mixed Models. Resistance distance of LLE had a stronger negative effect on Jaccard similarity than Euclidean distance in field margins, but not in ditches. This was found for species of 'meadows and pastures' and short to medium dispersal distance. In plot pairs that were highly connected by LLE, the models suggested that roughly 20% of all species occurred in both plots due to dispersal through LLE. Other species groups did not respond more strongly to resistance distance than to Euclidean distance. We conclude that linear landscape elements in agricultural landscapes are effective corridors for dispersal of plant species that are confined to semi-natural habitats, such as traditional grasslands, and lack mechanisms of long-distance dispersal.

Sensitivity of commercial pumpkin yield to potential decline among different groups of pollinating bees.

The yield of animal-pollinated crops is threatened by bee declines, but its precise sensitivity is poorly known. We therefore determined the yield dependence of Hokkaido pumpkin in Germany on insect pollination by quantifying: (i) the relationship between pollen receipt and fruit set and (ii) the cumulative pollen deposition of each pollinator group. We found that approximately 2500 pollen grains per flower were needed to maximize fruit set. At the measured rates of flower visitation, we estimated that bumblebees (21 visits/flower lifetime, 864 grains/visit) or honeybees (123 visits, 260 grains) could individually achieve maximum crop yield, whereas halictid bees are ineffective (11 visits, 16 grains). The pollinator fauna was capable of delivering 20 times the necessary amount of pollen. We therefore estimate that pumpkin yield was not pollination-limited in our study region and that it is currently fairly resilient to single declines of honeybees or wild bumblebees.

Impacts of invasive plants on resident animals across ecosystems, taxa, and feeding types: a global assessment.

As drivers of global change, biological invasions have fundamental ecological consequences. However, it remains unclear how invasive plant effects on resident animals vary across ecosystems, animal classes, and functional groups. We performed a comprehensive meta-analysis covering 198 field and laboratory studies reporting a total of 3624 observations of invasive plant effects on animals. Invasive plants had reducing (56%) or neutral (44%) effects on animal abundance, diversity, fitness, and ecosystem function across different ecosystems, animal classes, and feeding types while we could not find any increasing effect. Most importantly, we found that invasive plants reduced overall animal abundance, diversity and fitness. However, this significant overall effect was contingent on ecosystems, taxa, and feeding types of animals. Decreasing effects of invasive plants were most evident in riparian ecosystems, possibly because frequent disturbance facilitates more intense plant invasions compared to other ecosystem types. In accordance with their immediate reliance on plants for food, invasive plant effects were strongest on herbivores. Regarding taxonomic groups, birds and insects were most strongly affected. In insects, this may be explained by their high frequency of herbivory, while birds demonstrate that invasive plant effects can also cascade up to secondary consumers. Since data on impacts of invasive plants are rather limited for many animal groups in most ecosystems, we argue for overcoming gaps in knowledge and for a more differentiated discussion on effects of invasive plant on native fauna.

Effects of a Major Tree Invader on Urban Woodland Arthropods.

Biological invasions are a major threat to biodiversity; however, the degree of impact can vary depending on the ecosystem and taxa. Here, we test whether a top invader at a global scale, the tree Robinia pseudoacacia (black locust or false acacia), which is known to profoundly change site conditions, significantly affects urban animal diversity. As a first multi-taxon study of this kind, we analyzed the effects of Robinia dominance on 18 arthropod taxa by pairwise comparisons of woodlands in Berlin, Germany, that were dominated by R. pseudoacacia or the native pioneer tree Betula pendula. As a negative effect, abundances of five arthropod taxa decreased (Chilopoda, Formicidae, Diptera, Heteroptera, Hymenoptera); 13 others were not affected. Woodland type affected species composition of carabids and functional groups in spiders, but surprisingly did not decrease alpha and beta diversity of carabid and spider assemblages or the number of endangered species. Tree invasion thus did not induce biotic homogenization at the habitat scale. We detected no positive effects of alien dominance. Our results illustrate that invasions by a major tree invader can induce species turnover in ground-dwelling arthropods, but do not necessarily reduce arthropod species abundances or diversity and might thus contribute to the conservation of epigeal invertebrates in urban settings. Considering the context of invasion impacts thus helps to set priorities in managing biological invasions and can illustrate the potential of novel ecosystems to maintain urban biodiversity.

Effects of traditional flood irrigation on invertebrates in lowland meadows.

Lowland meadow irrigation used to be widespread in Central Europe, but has largely been abandoned during the 20th century. As a result of agri-environment schemes and nature conservation efforts, meadow irrigation is now being re-established in some European regions. In the absence of natural flood events, irrigation is expected to favour fauna typical of lowland wet meadows. We analysed the effects of traditional flood irrigation on diversity, densities and species composition of three invertebrate indicator taxa in lowland meadows in Germany. Unexpectedly, alpha diversity (species richness and Simpson diversity) and beta diversity (multivariate homogeneity of group dispersions) of orthopterans, carabids, and spiders were not significantly different between irrigated and non-irrigated meadows. However, spider densities were significantly higher in irrigated meadows. Furthermore, irrigation and elevated humidity affected species composition and shifted assemblages towards moisture-dependent species. The number of species of conservation concern, however, did not differ between irrigated and non-irrigated meadows. More variable and intensive (higher duration and/or frequency) flooding regimes might provide stronger conservation benefits, additional species and enhance habitat heterogeneity on a landscape scale.

Simulating small-scale climate change effects-lessons from a short-term field manipulation experiment on grassland arthropods.

Climate change is expected to cause major consequences on biodiversity. Understanding species-specific reactions, such as species shifts, species declines, and changes in population dynamics is a key issue to quantify large-scale impacts of climate change on biotic communities. As it is often impossible or at least impracticable to conduct large-scale experiments on biotic responses to climate change, studies at a smaller scale may be a useful alternative. In our study, we therefore tested responses of grassland arthropods (carabid beetles, spiders, grasshoppers) to simulated climate change in terms of species activity densities and diversity. We conducted a controlled field experiment by changing water and microclimatic conditions at a small scale (16 m(2) ). Roof constructions were used to increase drought-like conditions, whereas water supply was enhanced by irrigation. In all, 2 038 carabid beetles (36 species), 4 893 spiders (65 species), and 303 Orthoptera (4 species) were caught using pitfall traps from May to August, 2010. During our experiment, we created an artificial small-scale climate change; and statistics revealed that these changes had short-term effects on the total number of individuals and Simpson diversity of the studied arthropod groups. Moreover, our results showed that certain species might react very quickly to climate change in terms of activity densities, which in turn might influence diversity due to shifts in abundance patterns. Finally, we devised methodological improvements that may further enhance the validity of future studies.