Earlier and more uniform spring green-up linked to lower insect richness and biomass in temperate forests.

Urbanization and agricultural intensification are considered the main causes of recent insect decline in temperate Europe, while direct climate warming effects are still ambiguous. Nonetheless, higher temperatures advance spring leaf emergence, which in turn may directly or indirectly affect insects. We therefore investigated how Sentinel-2-derived start of season (SOS) and its spatial variability (SV-SOS) are affected by spring temperature and whether these green-up variables can explain insect biomass and richness across a climate and land-use gradient in southern Germany. We found that the effects of both spring green-up variables on insect biomass and richness differed between land-use types, but were strongest in forests. Here, insect richness and biomass were higher with later green-up (SOS) and higher SV-SOS. In turn, higher spring temperatures advanced SOS, while SV-SOS was lower at warmer sites. We conclude that with a warming climate, insect biomass and richness in forests may be affected negatively due to earlier and more uniform green-up. Promising adaptation strategies should therefore focus on spatial variability in green-up in forests, thus plant species and structural diversity.

Plant species composition and local habitat conditions as primary determinants of terrestrial arthropod assemblages.

Arthropods respond to vegetation in multiple ways since plants provide habitat and food resources and indicate local abiotic conditions. However, the relative importance of these factors for arthropod assemblages is less well understood. We aimed to disentangle the effects of plant species composition and environmental drivers on arthropod taxonomic composition and to assess which aspects of vegetation contribute to the relationships between plant and arthropod assemblages. In a multi-scale field study in Southern Germany, we sampled vascular plants and terrestrial arthropods in typical habitats of temperate landscapes. We compared independent and shared effects of vegetation and abiotic predictors on arthropod composition distinguishing between four large orders (Lepidoptera, Coleoptera, Hymenoptera, Diptera), and five functional groups (herbivores, pollinators, predators, parasitoids, detritivores). Across all investigated groups, plant species composition explained the major fraction of variation in arthropod composition, while land-cover composition was another important predictor. Moreover, the local habitat conditions depicted by the indicator values of the plant communities were more important for arthropod composition than trophic relationships between certain plant and arthropod species. Among trophic groups, predators showed the strongest response to plant species composition, while responses of herbivores and pollinators were stronger than those of parasitoids and detritivores. Our results highlight the relevance of plant community composition for terrestrial arthropod assemblages across multiple taxa and trophic levels and emphasize the value of plants as a proxy for characterizing habitat conditions that are hardly accessible to direct environmental measurements.

Low precipitation due to climate change consistently reduces multifunctionality of urban grasslands in mesocosms.

Urban grasslands are crucial for biodiversity and ecosystem services in cities, while little is known about their multifunctionality under climate change. Thus, we investigated the effects of simulated climate change, i.e., increased [CO2] and temperature, and reduced precipitation, on individual functions and overall multifunctionality in mesocosm grasslands sown with forbs and grasses in four different proportions aiming at mimicking road verge grassland patches. Climate change scenarios RCP2.6 (control) and RCP8.5 (worst-case) were simulated in walk-in climate chambers of an ecotron facility, and watering was manipulated for normal vs. reduced precipitation. We measured eight indicator variables of ecosystem functions based on below- and aboveground characteristics. The young grassland communities responded to higher [CO2] and warmer conditions with increased vegetation cover, height, flower production, and soil respiration. Lower precipitation affected carbon cycling in the ecosystem by reducing biomass production and soil respiration. In turn, the water regulation capacity of the grasslands depended on precipitation interacting with climate change scenario, given the enhanced water efficiency resulting from increased [CO2] under RCP8.5. Multifunctionality was negatively affected by reduced precipitation, especially under RCP2.6. Trade-offs arose among single functions that performed best in either grass- or forb-dominated grasslands. Grasslands with an even ratio of plant functional types coped better with climate change and thus are good options for increasing the benefits of urban green infrastructure. Overall, the study provides experimental evidence of the effects of climate change on the functionality of urban ecosystems. Designing the composition of urban grasslands based on ecological theory may increase their resilience to global change.

Dung-visiting beetle diversity is mainly affected by land use, while community specialization is driven by climate.

Dung beetles are important actors in the self-regulation of ecosystems by driving nutrient cycling, bioturbation, and pest suppression. Urbanization and the sprawl of agricultural areas, however, destroy natural habitats and may threaten dung beetle diversity. In addition, climate change may cause shifts in geographical distribution and community composition. We used a space-for-time approach to test the effects of land use and climate on α-diversity, local community specialization (H 2') on dung resources, and γ-diversity of dung-visiting beetles. For this, we used pitfall traps baited with four different dung types at 115 study sites, distributed over a spatial extent of 300 km × 300 km and 1000 m in elevation. Study sites were established in four local land-use types: forests, grasslands, arable sites, and settlements, embedded in near-natural, agricultural, or urban landscapes. Our results show that abundance and species density of dung-visiting beetles were negatively affected by agricultural land use at both spatial scales, whereas γ-diversity at the local scale was negatively affected by settlements and on a landscape scale equally by agricultural and urban land use. Increasing precipitation diminished dung-visiting beetle abundance, and higher temperatures reduced community specialization on dung types and γ-diversity. These results indicate that intensive land use and high temperatures may cause a loss in dung-visiting beetle diversity and alter community networks. A decrease in dung-visiting beetle diversity may disturb decomposition processes at both local and landscape scales and alter ecosystem functioning, which may lead to drastic ecological and economic damage.

Plant richness, land use and temperature differently shape invertebrate leaf-chewing herbivory on plant functional groups.

Higher temperatures can increase metabolic rates and carbon demands of invertebrate herbivores, which may shift leaf-chewing herbivory among plant functional groups differing in C:N (carbon:nitrogen) ratios. Biotic factors influencing herbivore species richness may modulate these temperature effects. Yet, systematic studies comparing leaf-chewing herbivory among plant functional groups in different habitats and landscapes along temperature gradients are lacking. This study was conducted on 80 plots covering large gradients of temperature, plant richness and land use in Bavaria, Germany. We investigated proportional leaf area loss by chewing invertebrates ('herbivory') in three plant functional groups on open herbaceous vegetation. As potential drivers, we considered local mean temperature (range 8.4-18.8 °C), multi-annual mean temperature (range 6.5-10.0 °C), local plant richness (species and family level, ranges 10-51 species, 5-25 families), adjacent habitat type (forest, grassland, arable field, settlement), proportion of grassland and landscape diversity (0.2-3 km scale). We observed differential responses of leaf-chewing herbivory among plant functional groups in response to plant richness (family level only) and habitat type, but not to grassland proportion, landscape diversity and temperature-except for multi-annual mean temperature influencing herbivory on grassland plots. Three-way interactions of plant functional group, temperature and predictors of plant richness or land use did not substantially impact herbivory. We conclude that abiotic and biotic factors can assert different effects on leaf-chewing herbivory among plant functional groups. At present, effects of plant richness and habitat type outweigh effects of temperature and landscape-scale land use on herbivory among legumes, forbs and grasses.

Interactive effects of climate and land use on pollinator diversity differ among taxa and scales.

Changes in climate and land use are major threats to pollinating insects, an essential functional group. Here, we unravel the largely unknown interactive effects of both threats on seven pollinator taxa using a multiscale space-for-time approach across large climate and land-use gradients in a temperate region. Pollinator community composition, regional gamma diversity, and community dissimilarity (beta diversity) of pollinator taxa were shaped by climate-land-use interactions, while local alpha diversity was solely explained by their additive effects. Pollinator diversity increased with reduced land-use intensity (forest < grassland < arable land < urban) and high flowering-plant diversity at different spatial scales, and higher temperatures homogenized pollinator communities across regions. Our study reveals declines in pollinator diversity with land-use intensity at multiple spatial scales and regional community homogenization in warmer and drier climates. Management options at several scales are highlighted to mitigate impacts of climate change on pollinators and their ecosystem services.

Landscape diversity and local temperature, but not climate, affect arthropod predation among habitat types.

Arthropod predators are important for ecosystem functioning by providing top-down regulation of insect herbivores. As predator communities and activity are influenced by biotic and abiotic factors on different spatial scales, the strength of top-down regulation ('arthropod predation') is also likely to vary. Understanding the combined effects of potential drivers on arthropod predation is urgently needed with regard to anthropogenic climate and land-use change. In a large-scale study, we recorded arthropod predation rates using artificial caterpillars on 113 plots of open herbaceous vegetation embedded in contrasting habitat types (forest, grassland, arable field, settlement) along climate and land-use gradients in Bavaria, Germany. As potential drivers we included habitat characteristics (habitat type, plant species richness, local mean temperature and mean relative humidity during artificial caterpillar exposure), landscape diversity (0.5-3.0-km, six scales), climate (multi-annual mean temperature, 'MAT') and interactive effects of habitat type with other drivers. We observed no substantial differences in arthropod predation rates between the studied habitat types, related to plant species richness and across the Bavarian-wide climatic gradient, but predation was limited when local mean temperatures were low and tended to decrease towards higher relative humidity. Arthropod predation rates increased towards more diverse landscapes at a 2-km scale. Interactive effects of habitat type with local weather conditions, plant species richness, landscape diversity and MAT were not observed. We conclude that landscape diversity favours high arthropod predation rates in open herbaceous vegetation independent of the dominant habitat in the vicinity. This finding may be harnessed to improve top-down control of herbivores, e.g. agricultural pests, but further research is needed for more specific recommendations on landscape management. The absence of MAT effects suggests that high predation rates may occur independent of moderate increases of MAT in the near future.

Modelling the Relative Abundance of Roe Deer (Capreolus capreolus L.) along a Climate and Land-Use Gradient.

European roe deer (Capreolus capreolus L.) are important given their economic, recreational and ecological value. However, uncontrolled roe deer numbers can result in negative impacts on forest regeneration and agricultural crops, disease transmission and occurrences of deer-vehicle collisions. Information on the abundance and distribution is needed for effective management. We combined distance sampling (DS) of roe deer dung pellet groups with multiple variables to develop a density surface model (DSM) in the federal state of Bavaria in south-eastern Germany. We used the estimates of pellet group density as a proxy for roe deer relative abundance. We extrapolated our best DSM, conducted a quantitative evaluation and contrasted relative abundance along climate and land-use gradients. Relative abundance of roe deer was influenced by a combination of habitat type, climate and wildlife management variables, which differed between seasons and which reflected changes in food and shelter availability. At the landscape scale, the highest abundance was observed in agriculture-dominated areas and the lowest in urban areas. Higher abundance was also observed in areas with intermediate temperatures compared to the warmest areas. Our results provide information on possible future changes in the distribution of relative abundance due to changes in climate and land-use.

Relationship of insect biomass and richness with land use along a climate gradient.

Recently reported insect declines have raised both political and social concern. Although the declines have been attributed to land use and climate change, supporting evidence suffers from low taxonomic resolution, short time series, a focus on local scales, and the collinearity of the identified drivers. In this study, we conducted a systematic assessment of insect populations in southern Germany, which showed that differences in insect biomass and richness are highly context dependent. We found the largest difference in biomass between semi-natural and urban environments (-42%), whereas differences in total richness (-29%) and the richness of threatened species (-56%) were largest from semi-natural to agricultural environments. These results point to urbanization and agriculture as major drivers of decline. We also found that richness and biomass increase monotonously with increasing temperature, independent of habitat. The contrasting patterns of insect biomass and richness question the use of these indicators as mutual surrogates. Our study provides support for the implementation of more comprehensive measures aimed at habitat restoration in order to halt insect declines.