Population synchrony indicates functional connectivity in a threatened sedentary butterfly.

Dispersal is a key influence on species' persistence, particularly in the context of habitat fragmentation and environmental change. Previously, residual population synchrony has been demonstrated to be an effective proxy for dispersal in mobile butterflies (Powney et al. 2012). Here, we highlight the utility and limitations of population synchrony as an indicator of functional connectivity and persistence, at a range of spatial scales, in a specialist, sedentary butterfly. While at the local scale, population synchrony is likely indicative of dispersal in the pearl-bordered fritillary, Boloria euphrosyne, over larger scales, habitat is likely to influence population dynamics. Although declines in local-scale synchrony conformed to typical movement in this species, synchrony showed no significant trend with distance when studied at larger (between-site) scales. By focusing on specific site comparisons, we draw the conclusion that heterogeneity in habitat successional stage drives asynchrony between sites at larger distances and is, therefore, likely to be a more important driver of population dynamics over large distances than dispersal. Within-site assessments of synchrony highlight differences in dispersal based on habitat type, with movement shown to be most inhibited between transect sections with contrasting habitat permeability. While synchrony has implications for metapopulation stability and extinction risk, no significant difference was found in average site synchrony between sites that had gone extinct during the study period and those remaining occupied. We demonstrate that population synchrony may be used to assess local-scale movement between sedentary populations, as well as to understand barriers to dispersal and guide conservation management.

The effectiveness of flower strips and hedgerows on pest control, pollination services and crop yield: a quantitative synthesis.

Floral plantings are promoted to foster ecological intensification of agriculture through provisioning of ecosystem services. However, a comprehensive assessment of the effectiveness of different floral plantings, their characteristics and consequences for crop yield is lacking. Here we quantified the impacts of flower strips and hedgerows on pest control (18 studies) and pollination services (17 studies) in adjacent crops in North America, Europe and New Zealand. Flower strips, but not hedgerows, enhanced pest control services in adjacent fields by 16% on average. However, effects on crop pollination and yield were more variable. Our synthesis identifies several important drivers of variability in effectiveness of plantings: pollination services declined exponentially with distance from plantings, and perennial and older flower strips with higher flowering plant diversity enhanced pollination more effectively. These findings provide promising pathways to optimise floral plantings to more effectively contribute to ecosystem service delivery and ecological intensification of agriculture in the future.

Resistance and tolerance to the brown planthopper, Nilaparvata lugens (Stål), in rice infested at different growth stages across a gradient of nitrogen applications.

High resource availability can reduce anti-herbivore resistance (a plant's ability to defend against herbivores and reduce damage) in rice, Oryza sativa L, but may also increase tolerance (a plant's ability to withstand damage by, for example, compensatory growth). Through a series of greenhouse, screenhouse and field experiments, this study examines fitness (survival and development × reproduction) of the brown planthopper, Nilaparvata lugens (Stål), on resistant (IR62) and susceptible (IR22) rice varieties and age-related rice tolerance to planthopper damage under varying resource (nitrogenous fertilizer) availability. Planthoppers reared on IR62 in the greenhouse had lower fitness than planthoppers on IR22. IR62 became increasingly resistant as plants aged. IR22 was generally more tolerant of planthopper damage, and tolerance increased in IR22, but declined in IR62, as the plants aged. Rice plants infested at pre-tillering stages (3-4 leaf stage) in the screenhouse had greater losses to root, shoot and grain yield per unit weight of planthopper than plants infested at tillering stages, particularly in IR22. These trends were mainly due to the impact of planthoppers during pre-tillering stages and the length of exposure to the planthoppers. High nitrogen compromised IR62 resistance, particularly in tillering plants in the greenhouse study; however, high nitrogen did not increase planthopper biomass-density on IR62 in greenhouse or field cages. Tolerance to damage in IR62 at mid-tillering stages declined under increasing levels of nitrogen, but nitrogen increased tolerance during late-tillering stages. Planthopper damage to IR22 in field cages was severe and hopperburn (plant death) occurred in 83% of IR22 plants under high nitrogen (60-150 kg N ha-1). In contrast, despite planthopper infestations, damage to IR62 was low in field-grown plants and productivity (tillers, roots, shoots and grain) increased in IR62 under increasing nitrogen. Our results indicate that, whereas nitrogenous fertilizer increases planthopper fitness on susceptible and resistant varieties, the net effects of high nitrogen on IR62 include decreased planthopper biomass-density (apparent in all experiments) and higher tolerance to damage during later growth stages (observed in the greenhouse, and during one of two seasons in field cages).

Legacy effects of grassland management on soil carbon to depth.

The importance of managing land to optimize carbon sequestration for climate change mitigation is widely recognized, with grasslands being identified as having the potential to sequester additional carbon. However, most soil carbon inventories only consider surface soils, and most large-scale surveys group ecosystems into broad habitats without considering management intensity. Consequently, little is known about the quantity of deep soil carbon and its sensitivity to management. From a nationwide survey of grassland soils to 1 m depth, we show that carbon in grassland soils is vulnerable to management and that these management effects can be detected to considerable depth down the soil profile, albeit at decreasing significance with depth. Carbon concentrations in soil decreased as management intensity increased, but greatest soil carbon stocks (accounting for bulk density differences), were at intermediate levels of management. Our study also highlights the considerable amounts of carbon in subsurface soil below 30 cm, which is missed by standard carbon inventories. We estimate grassland soil carbon in Great Britain to be 2097 Tg C to a depth of 1 m, with ~60% of this carbon being below 30 cm. Total stocks of soil carbon (t ha(-1) ) to 1 m depth were 10.7% greater at intermediate relative to intensive management, which equates to 10.1 t ha(-1) in surface soils (0-30 cm), and 13.7 t ha(-1) in soils from 30 to 100 cm depth. Our findings highlight the existence of substantial carbon stocks at depth in grassland soils that are sensitive to management. This is of high relevance globally, given the extent of land cover and large stocks of carbon held in temperate managed grasslands. Our findings have implications for the future management of grasslands for carbon storage and climate mitigation, and for global carbon models which do not currently account for changes in soil carbon to depth with management.

Effects of species evenness and dominant species identity on multiple ecosystem functions in model grassland communities.

Ecosystems provide multiple services upon which humans depend. Understanding the drivers of the ecosystem functions that support these services is therefore important. Much research has investigated how species richness influences functioning, but we lack knowledge of how other community attributes affect ecosystem functioning. Species evenness, species spatial arrangement, and the identity of dominant species are three attributes that could affect ecosystem functioning, by altering the relative abundance of functional traits and the probability of synergistic species interactions such as facilitation and complementary resource use. We tested the effect of these three community attributes and their interactions on ecosystem functions over a growing season, using model grassland communities consisting of three plant species from three functional groups: a grass (Anthoxanthum odoratum), a forb (Plantago lanceolata), and a N-fixing forb (Lotus corniculatus). We measured multiple ecosystem functions that support ecosystem services, including ecosystem gas exchange, water retention, C and N loss in leachates, and plant biomass production. Species evenness and dominant species identity strongly influenced the ecosystem functions measured, but spatial arrangement had few effects. By the end of the growing season, evenness consistently enhanced ecosystem functioning and this effect occurred regardless of dominant species identity. The identity of the dominant species under which the highest level of functioning was attained varied across the growing season. Spatial arrangement had the weakest effect on functioning, but interacted with dominant species identity to affect some functions. Our results highlight the importance of understanding the role of multiple community attributes in driving ecosystem functioning.

Herbivore species richness, composition and community structure mediate predator richness effects and top-down control of herbivore biomass.

Changes in predator species richness can have important consequences for ecosystem functioning at multiple trophic levels, but these effects are variable and depend on the ecological context in addition to the properties of predators themselves. Here, we report an experimental study to test how species identity, community attributes, and community structure at the herbivore level moderate the effects of predator richness on ecosystem functioning. Using mesocosms containing predatory insects and aphid prey, we independently manipulated species richness at both predator and herbivore trophic levels. Community structure was also manipulated by changing the distribution of herbivore species across two plant species. Predator species richness and herbivore species richness were found to negatively interact to influence predator biomass accumulation, an effect which is hypothesised to be due to the breakdown of functional complementarity among predators in species-rich herbivore assemblages. The strength of predator suppression of herbivore biomass decreased as herbivore species richness and distribution across host plants increased, and positive predator richness effects on herbivore biomass suppression were only observed in herbivore assemblages of relatively low productivity. In summary, the study shows that the species richness, productivity and host plant distribution of prey communities can all moderate the general influence of predators and the emergence of predator species richness effects on ecosystem functioning.

New understanding of the direct effects of spectral balance on behaviour in Myzus persicae.

The study of insect responses to colour has mainly focused on flying species and morphs, however colour cues are likely to be important for insect positioning within the canopy. We examine the role of illumination colour in canopy positioning of apterous Myzus persicae (Sulzer) using both a field experiment, utilising various UV-manipulating optical filters, and a laboratory experiment using video tracking of individuals illuminated by a variable intensity UVA-Blue-Green LED-array. In the field experiment, approximately twice as many aphids were located on exposed leaf surfaces under UV-deficient environments compared to UV-rich environments. The lab experiment showed all three M. persicae photoreceptors were involved in a visually-mediated feeding/avoidance behaviour. Highly UV-rich, green-deficient environments were up to 3 times as likely to trigger an avoidance behaviour compared to UV-absent, green-rich environments such as those found below the leaf surface. We show that apterous M. persicae use this, in addition to other cues, in order to locate feeding positions that minimise exposure to direct sunlight. This has relevance to both the fundamental understanding of photoprotective behaviour in Hemiptera as well as to applied research of crop production environments that disrupt pest behaviour.

Equivocal Evidence for Colony Level Stress Effects on Bumble Bee Pollination Services.

Climate change poses a threat to global food security with extreme heat events causing drought and direct damage to crop plants. However, by altering behavioural or physiological responses of insects, extreme heat events may also affect pollination services on which many crops are dependent. Such effects may potentially be exacerbated by other environmental stresses, such as exposure to widely used agro-chemicals. To determine whether environmental stressors interact to affect pollination services, we carried out field cage experiments on the buff-tailed bumble bee (Bombus terrestris). Using a Bayesian approach, we assessed whether heat stress (colonies maintained at an ambient temperature of 25 °C or 31 °C) and insecticide exposure (5 ng g-1 of the neonicotinoid insecticide clothianidin) could induce behavioural changes that affected pollination of faba bean (Vicia faba). Only the bumble bee colonies and not the plants were exposed to the environmental stress treatments. Bean plants exposed to heat-stressed bumble bee colonies (31 °C) had a lower proportional pod set compared to colonies maintained at 25 °C. There was also weak evidence that heat stressed colonies caused lower total bean weight. Bee exposure to clothianidin was found to have no clear effect on plant yields, either individually or as part of an interaction. We identified no effect of either colony stressor on bumble bee foraging behaviours. Our results suggest that extreme heat stress at the colony level may impact on pollination services. However, as the effect for other key yield parameters was weaker (e.g. bean yields), our results are not conclusive. Overall, our study highlights the need for further research on how environmental stress affects behavioural interactions in plant-pollinator systems that could impact on crop yields.

Stability lies in flowers: Plant diversification mediating shifts in arthropod food webs.

Arthropod community composition in agricultural landscapes is dependent on habitat characteristics, such as plant composition, landscape homogeneity and the presence of key resources, which are usually absent in monocultures. Manipulating agroecosystems through the insertion of in-field floral resources is a useful technique to reduce the deleterious effects of habitat simplification. Food web analysis can clarify how the community reacts to the presence of floral resources which favour ecosystem services such as biological control of pest species. Here, we reported quantitative and qualitative alterations in arthropod food web complexity due to the presence of floral resources from the Mexican marigold (Tagetes erecta L.) in a field scale lettuce community network. The presence of marigold flowers in the field successfully increased richness, body size, and the numerical and biomass abundance of natural enemies in the lettuce arthropod community, which affected the number of links, vulnerability, generality, omnivory rate and food chain length in the community, which are key factors for the stability of relationships between species. Our results reinforce the notion that diversification through insertion of floral resources may assist in preventing pest outbreaks in agroecosystems. This community approach to arthropod interactions in agricultural landscapes can be used in the future to predict the effect of different management practices in the food web to contribute with a more sustainable management of arthropod pest species.

Getting More Power from Your Flowers: Multi-Functional Flower Strips Enhance Pollinators and Pest Control Agents in Apple Orchards.

Flower strips are commonly recommended to boost biodiversity and multiple ecosystem services (e.g., pollination and pest control) on farmland. However, significant knowledge gaps remain regards the extent to which they deliver on these aims. Here, we tested the efficacy of flower strips that targeted different subsets of beneficial arthropods (pollinators and natural enemies) and their ecosystem services in cider apple orchards. Treatments included mixes that specifically targeted: (1) pollinators ('concealed-nectar plants'); (2) natural enemies ('open-nectar plants'); or (3) both groups concurrently (i.e., 'multi-functional' mix). Flower strips were established in alleyways of four orchards and compared to control alleyways (no flowers). Pollinator (e.g., bees) and natural enemy (e.g., parasitoid wasps, predatory flies and beetles) visitation to flower strips, alongside measures of pest control (aphid colony densities, sentinel prey predation), and fruit production, were monitored in orchards over two consecutive growing seasons. Targeted flower strips attracted either pollinators or natural enemies, whereas mixed flower strips attracted both groups in similar abundance to targeted mixes. Natural enemy densities on apple trees were higher in plots containing open-nectar plants compared to other treatments, but effects were stronger for non-aphidophagous taxa. Predation of sentinel prey was enhanced in all flowering plots compared to controls but pest aphid densities and fruit yield were unaffected by flower strips. We conclude that 'multi-functional' flower strips that contain flowering plant species with opposing floral traits can provide nectar and pollen for both pollinators and natural enemies, but further work is required to understand their potential for improving pest control services and yield in cider apple orchards.

Understanding biodiversity effects on prey in multi-enemy systems.

Biodiversity-ecosystem functioning theory would predict that increasing natural enemy richness should enhance prey consumption rate due to functional complementarity of enemy species. However, several studies show that ecological interactions among natural enemies may result in complex effects of enemy diversity on prey consumption. Therefore, the challenge in understanding natural enemy diversity effects is to predict consumption rates of multiple enemies taking into account effects arising from patterns of prey use together with species interactions. Here, we show how complementary and redundant prey use patterns result in additive and saturating effects, respectively, and how ecological interactions such as phenotypic niche shifts, synergy and intraguild predation enlarge the range of outcomes to include null, synergistic and antagonistic effects. This study provides a simple theoretical framework that can be applied to experimental studies to infer the biological mechanisms underlying natural enemy diversity effects on prey.

Harvester ant response to spatial and temporal heterogeneity in seed availability: pattern in the process of granivory.

The influence of temporal and spatial heterogeneity in seed availability on the foraging behaviour of the harvester ant Messor arenarius was studied in an arid shrubland in the Negev Desert, Israel. The study investigated the implications of behavioural responses to heterogeneity in seed availability for the seed predation process and the potential for feedback effects on vegetation. Vegetation and seed rain were monitored across two landscape patch types (shrub patches and inter-shrub patches) in 1997. Shrub patches were shown to have higher plant and seed-rain density than inter-shrub patches. Patch use and seed selection by M. arenarius foragers were monitored through the spring, summer and autumn of 1997. After a pulse of seed production in the spring, the ants exhibited very narrow diet breadth, specialising on a single annual grass species, Stipa capensis. At this time, ants were foraging and collecting seeds mainly from inter-shrub patches. In the summer, diet breadth broadened and use of shrub patches increased, although the rate of seed collection per unit area was approximately equal in the two patch types. The increase in the use of shrub patches was due to colony-level selection of foraging areas with relatively high shrub cover and an increase in the use of shrub patches by individual foragers. In the autumn, a pulse of seed production by the shrub species Atractylis serratuloides and Noaea mucronata led to a reduction in diet breadth as foragers specialised on these species. During this period, foragers exhibited a large increase in the proportion of time spent in shrub patches and in the proportion of food items collected from shrub patches. The seasonal patterns in foraging behaviour showed linked changes in seed selection and patch use resulting in important differences in the seed predation process between the two landscape patch types. For much of the study period, there was higher seed predation pressure on the inter-shrub patches, which were of relatively low productivity compared with the shrub patches. This suggests that the seed predation process may help maintain the spatial heterogeneity in the density of ephemeral plants in the landscape.

Plant species composition alters the sign and strength of an emergent multi-predator effect by modifying predator foraging behaviour.

The prediction of pest-control functioning by multi-predator communities is hindered by the non-additive nature of species functioning. Such non-additivity, commonly termed an emergent multi-predator effect, is known to be affected by elements of the ecological context, such as the structure and composition of vegetation, in addition to the traits of the predators themselves. Here we report mesocosm experiments designed to test the influence of plant density and species composition (wheat monoculture or wheat and faba bean polyculture) on the emergence of multi-predator effects between Adalia bipunctata and Chrysoperla carnea, in their suppression of populations of the aphid Metopolophium dirhodum. The mesocosm experiments were followed by a series of behavioural observations designed to identify how interactions among predators are modified by plant species composition and whether these effects are consistent with the observed influence of plant species composition on aphid population suppression. Although plant density was shown to have no influence on the multi-predator effect on aphid population growth, plant composition had a marked effect. In wheat monoculture, Adalia and Chrysoperla mixed treatments caused greater suppression of M. dirhodum populations than expected. However this positive emergent effect was reversed to a negative multi-predator effect in wheat and faba bean polyculture. The behavioural observations revealed that although dominant individuals did not respond to the presence of faba bean plants, the behaviour of sub-dominants was affected markedly, consistent with their foraging for extra-floral nectar produced by the faba bean. This interaction between plant composition and predator community composition on the foraging behaviour of sub-dominants is thought to underlie the observed effect of plant composition on the multi-predator effect. Thus, the emergence of multi-predator effects is shown to be strongly influenced by plant species composition, mediated, in this case, by the provision of extra-floral nectar by one of the plant species.

DENSITY-DEPENDENT EFFECTS OF PREDATOR SPECIES-RICHNESS IN DIVERSITY-FUNCTION STUDIES.

Ecological systems often exhibit a positive but saturating diversity-function curve. Variation in the mechanisms generating this relationship can alter the slope and variance of the curve, with implications for the optimal management of biodiversity for ecosystem services. In biological control, prevalence of selection effects supports augmentation of the most effective natural enemy, but complementarity effects support augmentation of natural enemy diversity. Optimization of biological control strategies from the results of diversity-function studies is limited because few consider changes in function with relative or absolute changes in abundance, and many confound the relative importance of richness and density through experimental designs (additive and substitutive). By manipulating species richness across an abundance gradient we show that effects of species richness are density dependent and indicate how this may be incorporated into experimental designs or models predicting resource consumption in diverse communities. Furthermore, the underlying mechanisms causing an observed diversity-function response, and its associated variation, changed across the richness-abundance gradient. Finally, species-rich assemblages provided higher levels of minimum function than species-poor assemblages, without any compromise on the maximum function possible.