Global warming can increase insect pest pressure by enhancing reproductive rates. Whether this translates into yield losses depends on phenological synchronisation of pests with their host plants and natural enemies. Simultaneously, landscape composition may mitigate climate effects by shaping the resource availability for pests and their antagonists. Here, we study the combined effects of temperature and landscape composition on pest abundances, larval parasitism, crop damage and yield, while also considering crop phenology, to identify strategies for sustainable management of oilseed rape (OSR) pests under warming climates. In all, 29 winter OSR crop fields were investigated in different climates (defined by multi-annual mean temperature, MAT) and landscape contexts in Bavaria, Germany. We measured abundances of adult pollen beetles and stem weevil larvae, pollen beetle larval parasitism, bud loss, stem damage and seed yield, and calculated the flowering date from growth stage observations. Landscape parameters (proportion of non-crop and OSR area, change in OSR area relative to the previous year) were calculated at six spatial scales (0.6-5 km). Pollen beetle abundance increased with MAT but to different degrees depending on the landscape context, that is, increased less strongly when OSR proportions were high (1-km scale), interannually constant (5-km scale) or both. In contrast, stem weevil abundance and stem damage did not respond to landscape composition nor MAT. Pollen beetle larval parasitism was overall low, but occasionally exceeded 30% under both low and high MAT and with reduced OSR area (0.6-km scale). Despite high pollen beetle abundance in warm climates, yields were high when OSR flowered early. Thereby, higher temperatures favoured early flowering. Only among late-flowering OSR crop fields yield was higher in cooler than warmer climates. Bud loss responded analogously. Landscape composition did not substantially affect bud loss and yield. Synthesis and applications: Earlier flowering of winter OSR compensates for higher pollen beetle abundance in warmer climates, while interannual continuity of OSR area prevents high pollen beetle abundance in the first place. Thus, regional coordination of crop rotation and crop management promoting early flowering may contribute to sustainable pest management in OSR under current and future climatic conditions.

The massive declines in terrestrial arthropods reported across Europe call for effective methods to monitor and promote biodiversity in human-dominated landscapes. Previous studies vary in their support for the suitability of plants as indicators of arthropod diversity, while the potential of subsets of conservation-relevant plant species to estimate arthropod richness remains to be tested. Moreover, the relative importance of plant species richness compared to other factors driving arthropod richness, such as land-use intensity, habitat amount and landscape configuration, is poorly understood. We conducted a multi-scale field study in Southern Germany, sampling vascular plants and terrestrial arthropods in four local land-use types (forest, grassland, arable field, settlement) across large-scale gradients of climate and land-use intensity. We obtained an extensive arthropod dataset using Malaise traps as sampling method and DNA metabarcoding for species identification. We compared the correlation of three sets of plant species richness (total, red-listed and biotope-indicator plants) with arthropod richness, including different trophic and taxonomic groups. Using mixed-effect models, we assessed the partial effects of plant species richness, habitat amount and landscape configuration on local arthropod richness while controlling for climate and land-use effects, and explored the environmental drivers of plant species richness. Arthropod species numbers of all trophic groups strongly responded to species numbers of plants that indicate protected habitats, while red-listed plant species richness was a key determinant for the richness of red-listed arthropods and butterflies. In most cases, plant species richness and temperature were stronger drivers of arthropod richness than the amount of surrounding semi-natural habitat and landscape configuration. While total plant species richness was highest in settlements, the richness of biotope-indicator plants was highest in forests and positively influenced by the amount of semi-natural habitat and edge density. The number of red-listed plant species increased with edge density at the landscape scale but was independent of habitat amount. We conclude that the richness of conservation-relevant plant species provides a powerful indicator of arthropod diversity, and underpins the potential of such plant lists for monitoring habitat quality. Our study highlights the importance of openings in forests for conservation of plants and arthropods as well as of preserving and restoring fragments of high-quality biotopes in agricultural and urban areas where the amount of semi-natural habitat is limited.

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.

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.

Climate and land use are major determinants of biodiversity, and declines in species richness in cold and human exploited landscapes can be caused by lower rates of biotic interactions. Deadwood fungi and bacteria interact strongly with their hosts due to long‐lasting evolutionary trajectories. However, how rates of biotic interactions (specialization) change with temperature and land‐use intensity are unknown for both microbial groups. We hypothesize a decrease in species richness and specialization of communities with decreasing temperature and increasing land use intensity while controlling for precipitation. We used a full‐factorial nested design to disentangle land use at habitat and landscape scale and temperature spanning an area of 300 × 300 km in Germany. We exposed four deadwood objects representing the main tree species in Central Europe (beech, oak, spruce, pine) in 175 study plots. Overall, we found that fungal and bacterial richness, community composition and specialization were weakly related to temperature and land use. Fungal richness was slightly higher in near‐natural than in urban landscapes. Bacterial richness was positively associated with mean annual temperature, negatively associated with local temperature and highest in grassland habitats. Bacterial richness was positively related to the covariate mean annual precipitation. We found strong effects of host‐tree identity on species richness and community composition. A generally high level of fungal host‐tree specialization might explain the weak response to temperature and land use. Effects of host‐tree identity and specialization were more pronounced in fungi. We suggest that host tree changes caused by land use and climate change will be more important for fungal communities, while changes in climate will affect bacterial communities more directly. Contrasting responses of the two taxonomic groups suggest a reorganization of deadwood microbial communities, which might have further consequences on diversity and decomposition in the Anthropocene.

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 spacefor-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.

AbstractDung 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 (H2′) 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.

SummaryClimate and land-use change are key drivers of environmental degradation in the Anthropocene, but too little is known about their interactive effects on biodiversity and ecosystem services. Long-term data on biodiversity trends are currently lacking. Furthermore, previous ecological studies have rarely considered climate and land use in a joint design, did not achieve variable independence or lost statistical power by not covering the full range of environmental gradients.Here, we introduce a multi-scale space-for-time study design to disentangle effects of climate and land use on biodiversity and ecosystem services. The site selection approach coupled extensive GIS-based exploration and correlation heatmaps with a crossed and nested design covering regional, landscape and local scales. Its implementation in Bavaria (Germany) resulted in a set of study plots that maximizes the potential range and independence of environmental variables at different spatial scales.Stratifying the state of Bavaria into five climate zones and three prevailing land-use types, i.e. near-natural, agriculture and urban, resulted in 60 study regions covering a mean annual temperature gradient of 5.6–9.8 °C and a spatial extent of 380×360 km. Within these regions, we nested 180 study plots located in contrasting local land-use types, i.e. forests, grasslands, arable land or settlement (local climate gradient 4.5–10 °C). This approach achieved low correlations between climate and land-use (proportional cover) at the regional and landscape scale with |r≤0.33| and |r≤0.29|, respectively. Furthermore, using correlation heatmaps for local plot selection reduced potentially confounding relationships between landscape composition and configuration for plots located in forests, arable land and settlements.The suggested design expands upon previous research in covering a significant range of environmental gradients and including a diversity of dominant land-use types at different scales within different climatic contexts. It allows independent assessment of the relative contribution of multi-scale climate and land use on biodiversity and ecosystem services. Understanding potential interdependencies among global change drivers is essential to develop effective restoration and mitigation strategies against biodiversity decline, especially in expectation of future climatic changes. Importantly, this study also provides a baseline for long-term ecological monitoring programs.

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.

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.

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.

Land-use intensification and climate change threaten ecosystem functions. A fundamental, yet often overlooked, function is decomposition of necromass. The direct and indirect anthropogenic effects on decomposition, however, are poorly understood. We measured decomposition of two contrasting types of necromass, rat carrion and bison dung, on 179 study sites in Central Europe across an elevational climate gradient of 168–1122 m a.s.l. and within both local and regional land uses. Local land-use types included forest, grassland, arable fields, and settlements and were embedded in three regional land-use types (near-natural, agricultural, and urban). The effects of insects on decomposition were quantified by experimental exclusion, while controlling for removal by vertebrates. We used generalized additive mixed models to evaluate dung weight loss and carrion decay rate along elevation and across regional and local land-use types. We observed a unimodal relationship of dung decomposition with elevation, where greatest weight loss occurred between 600 and 700 m, but no effects of local temperature, land use, or insects. In contrast to dung, carrion decomposition was continuously faster with both increasing elevation and local temperature. Carrion reached the final decomposition stage six days earlier when insect access was allowed, and this did not depend on land-use effect. Our experiment identified different major drivers of decomposition on each necromass form. The results show that dung and carrion decomposition are rather robust to local and regional land use, but future climate change and decline of insects could alter decomposition processes and the self-regulation of ecosystems.

Contemporary climate change leads to earlier spring phenological events in Europe. In forests, in which overstory strongly regulates the microclimate beneath, it is not clear if further change equally shifts the timing of leaf unfolding for the over- and understory of main deciduous forest species, such as Fagus sylvatica L. (European beech). Furthermore, it is not known yet how this vertical phenological (mis)match—the phenological difference between overstory and understory—affects the remotely sensed satellite signal. To investigate this, we disentangled the start of season (SOS) of overstory F.sylvatica foliage from understory F. sylvatica foliage in forests, within nine quadrants of 5.8 × 5.8 km, stratified over a temperature gradient of 2.5 °C in Bavaria, southeast Germany, in the spring seasons of 2019 and 2020 using time lapse cameras and visual ground observations. We explained SOS dates and vertical phenological (mis)match by canopy temperature and compared these to Sentinel-2 derived SOS in response to canopy temperature. We found that overstory SOS advanced with higher mean April canopy temperature (visual ground observations: −2.86 days per °C; cameras: −2.57 days per °C). However, understory SOS was not significantly affected by canopy temperature. This led to an increase of vertical phenological mismatch with increased canopy temperature (visual ground observations: +3.90 days per °C; cameras: +2.52 days per °C). These results matched Sentinel-2-derived SOS responses, as pixels of higher canopy height advanced more by increased canopy temperature than pixels of lower canopy height. The results may indicate that, with further climate change, spring phenology of F. sylvatica overstory will advance more than F. sylvatica understory, leading to increased vertical phenological mismatch in temperate deciduous forests. This may have major ecological effects, but also methodological consequences for the field of remote sensing, as what the signal senses highly depends on the pixel mean canopy height and the vertical (mis)match.

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.

Interactions between plants and herbivorous invertebrates drive the nutritional quality of resources for higher trophic levels, nutrient cycling and plant-community structure. Thereby, shifts in functional composition of plant communities particularly impact ecosystem processes. However, the current understanding of herbivory is limited concerning climate, land use and plant richness, as comparative studies of different plant functional groups are lacking. This study was conducted on 81 plots covering large climatic and land-use gradients in Bavaria, Germany. We investigated foliar invertebrate herbivory rates (proportional leaf-area loss, following ‘herbivory’) in three major plant functional groups (legumes, non-leguminous forbs, grasses). As drivers we considered multi-annual mean temperature (range: 6.5–10.0 °C), local habitat type (forest, grassland, arable field, settlement), local plant richness (species and family level, ranges: 10–50 species, 5–25 families) and landscape diversity (0.2–3-km scale). Our results largely confirm higher herbivory on legumes than on forbs and grasses. However, herbivory in forests was similar across plant functional groups since herbivory on legumes was low, e.g. lower than on legumes in grasslands. We also observed differential responses of herbivory among plant functional groups in response to plant richness (family level only), but not to landscape diversity. Temperature did not affect overall herbivory, but in grasslands higher temperature decreased herbivory on legumes and increased on forbs and grasses. We conclude that climate, habitat type and family-level plant richness likely assert different effects on herbivory among plant functional groups. This emphasises the importance of functional groups for understanding community-level herbivory and ecosystem functioning.

Using RGB camera data (e.g. webcams, wildlife cameras) has great potential to measure forest phenology over climate gradients, because of its very high temporal resolution, while at the same time being more objective and less time consuming than in situ observations. To make images useful for the purpose of measuring phenological events, such as Start of Season (SOS) and End of Season (EOS), there is need to derive Regions of Interest (ROI) objectively and (semi-)automatically. In order to answer this need, Bothmann et al. (2017) proposed a method which randomly sets a number of pinpricks in the image and calculates how greenness over time from all other pixels correlates to these different pinpricks. Subsequently, ROIs are created by discarding the pixels with low correlation, using multiple thresholds. Despite its advantage of being automated and more objective compared to prevailing expert-based ROIs, and therefore its potential applicability for phenological research using a large amount of cameras, the method has not been reproduced for this purpose so far. Therefore, we assess here how well this method is able to separate foliage of different deciduous species from evergreens and phenologically irrelevant components in time-lapse wildlife camera data and in that way how suitable it is in explaining variation in phenology over a temperature gradient. We used 73 Cuddleback wildlife cameras troughout Bavaria which were installed within nine quadrants of 6*6 kilometers spanning a temperature gradient of 2.5°C. Hourly taken images of deciduous forests in spring, summer and autumn 2019 were analysed. Half of them were facing canopy, and half of them were facing understory. We applied the principles of the method from Bothmann et al. (2017) and assigned the best matching ROI to foliage of Fagus sylvatica or other deciduous species. Within this ROI, mean Green Chromatic Coordinate (GCC), a greenness index, over all pixels within the ROI, was derived per time-stamp. Afterwards, a time-series was calculated on these GCC values and with a suitable combination of curve-fitting techniques, SOS and EOS were derived, expressed in Day of Year (DOY). We compared these SOS and EOS dates with weekly in situ observations of spring and autumn phenology, which were taken in the same quadrants. Despite that Bothmann's method was developed on a single tower-mounted scientific webcam which viewed on canopy from above, while we made use of wildlife cameras at 73 different locations facing either understory perpendicular or canopy from below, it was able to distinguish F. sylvatica and other deciduous foliage from phenologically less relevant information. Time-series derived from these ROIs were able to explain variability in phenology between understory and canopy and over the temperature gradient similarly and supplementary to in situ observations.