Land-use affects pollinator-specific resource availability and pollinator foraging behaviour.

Land-use management is a key factor causing pollinator declines in agricultural grasslands. This decline can not only be directly driven by land-use (e.g., habitat loss) but also be indirectly mediated through a reduction in floral resource abundance and diversity, which might in turn affect pollinator health and foraging. We conducted surveys of the abundance of flowering plant species and behavioural observations of two common generalist pollinator species, namely the bumblebee Bombus lapidarius and the syrphid fly Episyrphus balteatus, in managed grasslands of variable land-use intensity (LUI) to investigate whether land-use affects (1) resource availability of the pollinators, (2) their host plant selection and (3) pollinator foraging behaviour. We have found that the floral composition of plant species that were used as resource by the investigated pollinator species depends on land-use intensity and practices such as mowing or grazing. We have also found that bumblebees, but not syrphid flies, visit different plants depending on LUI or management type. Furthermore, LUI indirectly changed pollinator behaviour via a reduction in plot-level flower diversity and abundance. For example, bumblebees show longer flight durations with decreasing flower cover indicating higher energy expenditure when foraging on land-use intensive plots. Syrphid flies were generally less affected by local land use, showing how different pollinator groups can differently react to land-use change. Overall, we show that land-use can change resource composition, abundance and diversity for pollinators, which can in turn affect pollinator foraging behaviour and potentially contribute to pollinator decline in agricultural grasslands.

Land-use-associated stressors interact to reduce bumblebee health at the individual and colony level.

In agricultural landscapes, bees face a variety of stressors, including insecticides and poor-quality food. Although both stressors individually have been shown to affect bumblebee health negatively, few studies have focused on stressor interactions, a scenario expected in intensively used agricultural landscapes. Using the bumblebee Bombus terrestris, a key pollinator in agricultural landscapes, we conducted a fully factorial laboratory experiment starting at nest initiation. We assessed the effects of food quality and insecticides, alone and in interaction, on health traits at various levels, some of which have been rarely studied. Pollen with a diluted nutrient content (low quality) reduced ovary size and delayed colony development. Wing asymmetry, indicating developmental stress, was increased during insecticide exposure and interactions with poor food, whereas both stressors reduced body size. Both stressors and their interaction changed the workers' chemical profile and reduced worker interactions and the immune response. Our findings suggest that insecticides combined with nutritional stress reduce bumblebee health at the individual and colony levels, thus possibly affecting colony performance, such as development and reproduction, and the stability of plant-pollinator networks. The synergistic effects highlight the need of combining stressors in risk assessments and when studying the complex effects of anthropogenic stressors on health outcomes.

Intra-individual variation in Galium odoratum is affected by experimental drought and shading.

BACKGROUND AND AIMS: Climate-change induced warmer spring temperatures advance tree leaf-out and result in earlier shading of the forest floor. Climate change also leads to more frequent droughts. Forest understorey herbs may respond to these environmental changes by varying traits at different hierarchical levels of organization. While trait mean variation at the inter-individual level in response to environmental changes is well-studied, little is known about how variation at the intra-individual level responds. METHODS: We sampled genets of the forest understorey herb Galium odoratum from 21 populations in three regions in Germany, varying in microclimatic conditions. The genets were transplanted into a common garden, where we applied shading and drought treatments. We measured plant height and leaf length and width, and calculated the coefficient of variation (CV) at different hierarchical levels: intra-population, intra-genet, intra-ramet and intra-shoot. KEY RESULTS: Variance partitioning showed that intra-shoot CV represented most of the total variation, followed by intra-ramet CV. We found significant variation in CV of plant height and leaf width among populations of origin, indicating that CV is at least partly genetically based. The soil temperature at populations' origins correlated negatively with CV in plant height, suggesting adaptation to local conditions. Furthermore, we observed that early shade led to increased intra-ramet CV in leaf length, while drought reduced intra-shoot CV in leaf width. Finally, intra-shoot leaf width mean and CV were independent under control conditions but correlated under drought. CONCLUSIONS: Our experimental results reveal correlations of intra-individual variation with soil temperature, indicating that intra-individual variation can evolve and may be adaptive. Intra-individual variation responded plastically to drought and shading, suggesting functional changes to improve light capture and reduce evapotranspiration. In conclusion, intra-individual variation makes up the majority of total trait variation in this species and can play a key role in plant adaptation to climatic change.

Water Deficit, Nitrogen Availability, and Their Combination Differently Affect Floral Scent Emission in Three Brassicaceae Species.

Floral scent plays a central role in plant-pollinator interactions, as flower visitors can discriminate between scent differences to recognize and forage on rewarding flowers. Changes in scent compositions might therefore lead to recognition mismatches between host plants and flower visitors. An understanding of the phenotypic plasticity of floral scent, especially in crop species, is becoming important because of climate change, e.g., increasing drought periods, and other anthropogenic influences, e.g., nitrogen (N) deposition. We have investigated the effects of the combination of progressive water deficits (dry-down) and N supplementation on floral scent emission in three Brassicaceae species (cultivated vs. wild). Individuals were randomly assigned to one of four treatments: (1) well-watered without N supplementation; (2) well-watered with N supplementation; (3) dry-down without N supplementation; (4) dry-down with N supplementation. We collected scent on day 0, 2, 7, and 14 after the commencement of the watering treatment. All samples were analyzed using gas chromatography coupled with mass spectrometry. We found that the highly cultivated Brassica napus had the lowest overall emission rate; its scent composition was affected by the interaction of watering treatment and N supplementation. Scent bouquets of the cultivated Sinapis alba also changed under these treatments. Scent bouquets of the common weed Sinapis arvensis were affected by watering treatment, but not by time and N supplementation. Furthermore, the influence of treatments on the emission rate of single compounds was highly compound-specific. Nonetheless, our study revealed that especially terpenes were negatively affected by drought-stress.

Land-use stress alters cuticular chemical surface profile and morphology in the bumble bee Bombus lapidarius.

Pollinators and other insects are currently undergoing a massive decline. Several stressors are thought to be of importance in this decline, with those having close relationships to agricultural management and practice seemingly playing key roles. In the present study, we sampled Bombus lapidarius L. workers in grasslands differing in their management intensity and management regime across three different regions along a north-south gradient in Germany. We analyzed the bees with regard to (1) their cuticular hydrocarbon profile (because of its important role in communication in social insects) and amount of scent by using gas chromatography and (2) the size of each individual by using wing distances as a proxy for body size. Our analysis revealed changes related to land-use intensity and temperature in the cuticular scent profile of bumble bees. Decreasing body size and increasing total scent amount were explained by an interaction of land-use intensity and study region, but not by land-use intensity alone. Thus, land-use intensity and temperature influence intracolonial communication and size, both of which can have strong effects on foraging. Land management and climate are therefore probably detrimental for colony maintenance and the reproductive success of bumble bees.

Bimodal activity of diurnal flower visitation at high elevation.

Successful pollination in animal-pollinated plants depends on the temporal overlap between flower presentation and pollinator foraging activity. Variation in the temporal dimension of plant-pollinator networks has been investigated intensely across flowering seasons. However, over the course of a day, the dynamics of plant-pollinator interactions may vary strongly due environmental fluctuations. It is usually assumed there is a unimodal, diurnal, activity pattern, while alternative multimodal types of activity patterns are often neglected and deserve greater investigation. Here, we quantified the daily activity pattern of flower visitors in two different habitats contrasting high elevation meadows versus forests in Southwest China to investigate the role of abiotic conditions in the temporal dynamics of plant-pollinator interactions. We examined diurnal activity patterns for the entire pollinator community. Pollinator groups may differ in their ability to adapt to habitats and abiotic conditions, which might be displayed in their patterns of activity. We hypothesized that (a) pollinator communities show multimodal activity patterns, (b) patterns differ between pollinator groups and habitat types, and (c) abiotic conditions explain observed activity patterns. In total, we collected 4,988 flower visitors belonging to six functional groups. There was a bimodal activity pattern when looking at the entire pollinator community and in five out of six flower visitor groups (exempting solitary bees) regardless of habitat types. Bumblebees, honeybees, dipterans, lepidopterans, and other insects showed activity peaks in the morning and afternoon, whereas solitary bees were most active at midday. Activity of all six pollinator groups increased as solar radiation increased and then decreased after reaching a certain threshold. Our findings suggest that in habitats at higher elevations, a bimodal activity pattern of flower visitation is commonly employed across most pollinator groups that are diurnal foragers. This pattern may be caused by insects avoiding overheating due to elevated temperatures when exposed to high solar radiation at midday.

A meta-analysis of responses in floral traits and flower-visitor interactions to water deficit.

Alterations in water availability and drought events as predicted by climate change scenarios will increasingly impact natural communities with effects already emerging at present. Water deficit leads to increasing physiological stress in plants, likely affecting floral development and causing changes in floral morphology, nectar and pollen production or scent. Understanding how these floral traits are altered by water deficit is necessary to predict changes in plant-pollinator interactions and how communities are impacted in the future. Here we employ a meta-analysis approach to synthesize the current evidence of experimental water deficit on floral traits and plant-pollinator interactions. Furthermore, we explore experimental factors potentially increasing heterogeneity between studies and provide ideas how to enhance comparability between studies. In the end, we highlight future directions and knowledge gaps for floral traits and plant-pollinator interactions under water deficit. Our analysis showed consistent decreases in floral size, number of flowers and nectar volume to reduced water availability. Other floral traits such as the start of flowering or herkogamy showed no consistent pattern. This indicates that effects of reduced water availability differ between specific traits that are potentially involved in different functions such as pollinator attraction or efficiency. We found no general decreasing visitation rates with water deficit for flower-visitor interactions. Furthermore, the comparison of available studies suggests that increased reporting of plant stress severity and including more hydraulic and physiological measurements will improve the comparability across experiments and aid a more mechanistic understanding of plant-pollinator interactions under altered environmental conditions. Overall, our results show that water deficit has the potential to strongly affect plant-pollinator interactions via changes in specific floral traits. Linking these changes to pollination services and pollinator performance is one crucial step for understanding how changing water availability and drought events under climate change will alter plant and pollinator communities.

Local and Landscape Effects on Carrion-Associated Rove Beetle (Coleoptera: Staphylinidae) Communities in German Forests.

Intensification of anthropogenic land use is a major threat to biodiversity and thus to essential ecosystem services provided by insects. Rove beetles (Coleoptera: Staphylinidae), which react sensitively to habitat changes, are species-rich colonizers of vertebrate cadavers and contribute to the important ecosystem service of carrion decomposition. The unveiling of anthropogenic and environmental drivers that modify carrion-associated rove beetle communities should improve our understanding of the plasticity of cadaver decay. We report the presence of 80 rove beetle species on 65 decomposing piglet cadavers at forest sites characterized by a gradient of management intensity across three geographic regions in Germany. Local and landscape drivers were revealed that shape beetle abundance, diversity, and community composition. Forest management and regions affect rove beetle abundance, whereas diversity is influenced by local habitat parameters (soil pH, litter cover) and regions. The community composition of rove beetles changes with management intensification by promoting generalist species. Regarding single species, Philonthus decorus and Anotylus mutator are linked to unmanaged forests and Ontholestes tessellatus to highly used forest stands. The spatial information provided about carrion-associated rove beetle communities in German forests is not only of carrion-ecological but also of forensic entomological interest.

Floral traits are associated with the quality but not quantity of heterospecific stigmatic pollen loads.

BACKGROUND: In flowering communities, plant species commonly share pollinators and therefore plant individuals receive heterospecific pollen (HP). However, the patterns of HP transfers can deviate from patterns of plant-pollinator visitations. Although flower-visitor interactions are known to be mediated by floral traits, e.g. floral size or nectar tube depth, the explanatory power of these traits for HP transfer patterns remains elusive. Here, we have explored pollen transfer patterns at three sites in Southern Germany on three dates (early, mid and late summer). At the plant level, we tested whether flower abundance and floral traits are correlated with HP reception and donation. At the community level, we determined whether flower and bee diversity are correlated with network modularity and whether floral traits explain the module affiliation of plant species. We collected the stigmas of flowering plant species, analysed HP and conspecific pollen (CP) loads and measured floral traits, flower and bee diversity. RESULTS: Our results show that the degree and intensity of HP reception or donation at the plant level do not correlate with floral traits, whereas at the community level, the module affiliation of who is sharing pollen with whom is well-explained by floral traits. Additionally, variation in network modularity between communities is better explained by plant diversity and abundance than by bee diversity and abundance. CONCLUSIONS: Overall, our results indicate that floral traits that are known to mediate flower-visitor interactions can improve our understanding of qualitative HP transfer but only provide limited information about the quantity of HP transfer, which more probably depends on other floral traits, flower-visitor identity or community properties.

Ultraviolet radiation changes plant color.

BACKGROUND: Plant absorption of ultraviolet (UV) radiation can result in multiple deleterious effects to plant tissues. As a result, plants have evolved an array of strategies to protect themselves from UV radiation, particularly in the UV-B range (280-320 nm). A common plant response to UV exposure is investment in phenolic compounds that absorb damaging wavelengths of light. However, the inverse phenomenon - plant reflectance of UV to protect plant tissues - has not previously been explored. In a paired experiment, we expose half of our sample (N = 108) of insect-pollinated plants of the cultivar Zinnia Profusion Series to UV radiation, and protect the other half from all light < 400 nm for 42 days, and measure leaf and flower reflectance using spectroscopy. We compare UV-B reflectance in leaves and flowers at the beginning of the experiment or flowering, and after treatment. RESULTS: We find that plants protected from UV exposure downregulate UV-B reflectance, and that plants exposed to increased levels of UV show trends of increased UV-B reflectance. CONCLUSIONS: Our results indicate that upregulation of UV-B reflecting pigments or structures may be a strategy to protect leaves against highly energetic UV-B radiation.

Bumblebee Behavior on Flowers, but Not Initial Attraction, Is Altered by Short-Term Drought Stress.

Climate change is leading to increasing drought and higher temperatures, both of which reduce soil water levels and consequently water availability for plants. This reduction often induces physiological stress in plants, which in turn can affect floral development and production inducing phenotypic alterations in flowers. Because flower visitors notice and respond to small differences in floral phenotypes, changes in trait expression can alter trait-mediated flower visitor behavior. Temperature is also known to affect floral scent emission and foraging behavior and, therefore, might modulate trait-mediated flower visitor behavior. However, the link between changes in flower visitor behavior and floral traits in the context of increasing drought and temperature is still not fully understood. In a wind-tunnel experiment, we tested the behavior of 66 Bombus terrestris individuals in response to watered and drought-stressed Sinapis arvensis plants and determined whether these responses were modulated by air temperature. Further, we explored whether floral traits and drought treatment were correlated with bumblebee behavior. The initial attractiveness of drought and watered plants did not differ, as the time to first visit was similar. However, bumblebees visited watered plants more often, their visitation rate to flowers was higher on watered plants, and bumblebees stayed for longer, indicating that watered plants were more attractive for foraging. Bumblebee behavior differed between floral trait expressions, mostly independently of treatment, with larger inflorescences and flowers leading to a decrease in the time until the first flower visit and an increase in the number of visits and the flower visitation rate. Temperature modulated bumblebee activity, which was highest at 25°C; the interaction of drought/water treatment and temperature led to higher visitation rate on watered plants at 20°C, possibly as a result of higher nectar production. Thus, bumblebee behavior is influenced by the watered status of plants, and bumblebees can recognize differences in intraspecific phenotypes involving morphological traits and scent emission, despite overall morphological traits and scent emission not being clearly separated between treatments. Our results indicate that plants are able to buffer floral trait expressions against short-term drought events, potentially to maintain pollinator attraction.

Interconnectedness of the Grinnellian and Eltonian Niche in Regional and Local Plant-Pollinator Communities.

Understanding the causes and consequences of coexistence and thus biodiversity is one of the most fundamental endeavors of ecology, which has been addressed by studying species' requirements and impacts - conceptualized as their Grinnellian and Eltonian niches. However, different niche types have been mostly studied in isolation and thus potential covariation between them remains unknown. Here we quantified the realized Grinnellian niche (environmental requirements), the fundamental (morphological phenotype) and realized Eltonian niche (role in networks) of plant and pollinator taxa at a local and regional scale to investigate the interconnectedness of these niche types. We found a strong and scale-independent co-variation of niche types suggesting that taxa specialized in environmental factors are also specialized in their position in trait spaces and their role in bipartite networks. The integration of niche types thus will help to detect the true causes for species distributions, interaction networks, as well as the taxonomic and functional diversity of communities.

A Fast and Robust Way to Estimate Overlap of Niches, and Draw Inference.

The problem of quantifying the overlap of Hutchinsonian niches has received much attention lately, in particular in quantitative ecology, from where it also originates. However, the niche concept has the potential to also be useful in many other application areas, as for example in economics. We are presenting a fully nonparametric, robust solution to this problem, along with exact shortcut formulas based on rank-statistics, and with a rather intuitive probabilistic interpretation. Furthermore, by deriving the asymptotic sampling distribution of the estimators, we are proposing the first asymptotically valid inference method, providing confidence intervals for the niche overlap. The theoretical considerations are supplemented by simulation studies and a real data example.

Covariation and phenotypic integration in chemical communication displays: biosynthetic constraints and eco-evolutionary implications.

Chemical communication is ubiquitous. The identification of conserved structural elements in visual and acoustic communication is well established, but comparable information on chemical communication displays (CCDs) is lacking. We assessed the phenotypic integration of CCDs in a meta-analysis to characterize patterns of covariation in CCDs and identified functional or biosynthetically constrained modules. Poorly integrated plant CCDs (i.e. low covariation between scent compounds) support the notion that plants often utilize one or few key compounds to repel antagonists or to attract pollinators and enemies of herbivores. Animal CCDs (mostly insect pheromones) were usually more integrated than those of plants (i.e. stronger covariation), suggesting that animals communicate via fixed proportions among compounds. Both plant and animal CCDs were composed of modules, which are groups of strongly covarying compounds. Biosynthetic similarity of compounds revealed biosynthetic constraints in the covariation patterns of plant CCDs. We provide a novel perspective on chemical communication and a basis for future investigations on structural properties of CCDs. This will facilitate identifying modules and biosynthetic constraints that may affect the outcome of selection and thus provide a predictive framework for evolutionary trajectories of CCDs in plants and animals.

Time-invariant differences between plant individuals in interactions with arthropods correlate with intraspecific variation in plant phenology, morphology and floral scent.

The basic units of ecological and evolutionary processes are individuals. Network studies aiming to infer mechanisms from complex systems, however, usually focus on interactions between species, not individuals. Accordingly, the structure and underlying mechanisms of individual-based interaction networks remain largely unknown. In a common garden, we recorded all interactions on flowers and leaves of 97 Sinapis arvensis individuals from seedling stage to fruit set and related interindividual differences in interactions to the plant individuals' phenotypes. The plant individuals significantly differed in their quantitative and qualitative interactions with arthropods on flowers and leaves. These differences remained stable over the entire season and thus were time-invariant. Variation in interacting arthropod communities could be explained by a pronounced intraspecific variability in flowering phenology, morphology and flower scent, and translated into variation in reproductive success. Interestingly, plant individuals with a similar composition of flower visitors were also visited by a similar assemblage of interaction partners at leaves. Our results show that the nonuniformity of plant species has pronounced effects in community ecology, potentially with implications for the persistence of communities and populations, and their ability to withstand environmental fluctuations.