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Scientists have largely neglected the effects of grazing on soil microbial communities despite their importance as drivers of ecosystem functions and services. We hypothesized that changes in soil properties resulting from grazing regulate the diversity of soil microbes by releasing/suppressing subordinate microbial taxa via competition. To test this, we examined how intensity of vertebrate herbivores influences the diversity and composition of soil bacteria and fungi at 216 soil samples from 54 sites across four microsites. Increasing grazing intensity reduced soil carbon, suppressing the dominant bacterial phylum Actinobacteria (indirectly promoting bacterial diversity) and increasing the dominant fungal phylum Ascomycetes (indirectly reducing fungal diversity). Our data provide novel evidence that grazing modulates the diversity and composition of soil microbes via increases or reductions in competition by dominant taxa. Our results suggest that grazing can potentially alter soil function by altering microbial community composition, providing a clear link between grazing management, carbon availability and ecosystem functions.

1. Grazing is one of the most widespread forms of intensive management on Earth and is linked to reductions in soil health. However, little is known about the relative influence of herbivore type, herbivore intensity and site productivity on soil health. This lack of knowledge reduces our capacity to manage landscapes where grazing is a major land use. 2. We used structural equation modelling to assess the effects of recent (cattle, sheep, goats, kangaroos and rabbit dung) and historic (cattle, sheep/goat livestock tracks) herbivore activity on soil health at 451 sites across 0.5 M km² of eastern Australia. We assessed the direct and indirect effects of increasing herbivore intensity, using dung and livestock tracks, on 15 morphological, physical and chemical attributes that are indicative of soil health, and we used these attributes to derive three indices representing the capacity of the soil to maintain its structural integrity (stability), cycle nutrients (nutrients) and maintain water flow (infiltration). 3. Grazing had negative effects on the three soil health indices, but these effects varied with productivity. Grazing intensity was associated with strong reductions in the stability and nutrient indices under low productivity, but these effects diminished with increasing productivity. Herbivore effects on individual attributes varied in relation to productivity level and were strongly herbivore specific, with most due to cattle grazing, and to a lesser extent, sheep, goats and rabbits. Few effects due to kangaroos or historic grazing by livestock were observed. 4. Synthesis and applications. Our study shows that livestock and rabbits degrade soil health through grazing, and its effects are strongest under low or moderate productivity; however, kangaroo effects are benign. Our findings support calls for resource management agencies to consider site productivity, as well as herbivore type and intensity, when developing strategies to manage grazing by livestock, and feral and native herbivores.

1. Conservation biological control aims to control pests by promoting wild populations of natural enemies. One challenge is to attract and retain efficient natural enemies in crop fields, which often are a suboptimal environment. Towards this goal, the attract-and-reward strategy relies on combining attractive synthetically produced herbivore-induced plant volatiles (HIPVs) with companion plants (non-crop plants which provide alternative resources to the targeted natural enemies). Although severely overlooked, the spatial arrangement of HIPV dispensers and rewards inside crop fields may strongly influence the foraging behaviour and persistence of natural enemies and thus the success of this pest management strategy.2. We tested the impact of two contrasting spatial arrangements of HIPV dispensers and rewards, alternatively inside and around a block of target apple trees, on the efficacy of the biological control of Aphis citricola populations by the common predatory ladybird Propylea japonica in apple orchards in northern China. We used synthetic methyl salicylate (MeSA) as an attractant and the companion plant Calendula officinalis as a reward. To better understand how the spatial arrangement of MeSA dispensers and companion plants affected the attraction and foraging behaviour of adult ladybirds, we conducted indoor experiments in a flight mill, an olfactometer and a wind-tunnel.3. Blocks of target trees treated with MeSA dispensers inside and companion plants around provided the most efficient pest control in orchards, compared with the opposite spatial arrangement.4. The synthetic MeSA dispenser and the companion plant synergistically attracted ladybirds in the olfactometer and enhanced their flight activity in the flight mill. In the wind-tunnel, MeSA served as a spatial cue for ladybirds to find nearby prey, while companion plants were sought in the absence of prey.5. Synthesis and applications. The present study will help further improvements of aphid control in apple orchards through a careful spatial arrangement of herbivore-induced plant volatiles dispensers (HIPVs) and rewards (companion plants) in optimized attract-and-reward strategies. Without such assessment, these strategies may be hazardous even with well-identified targeted natural enemies. Associated lab experiments highlight that HIPVs and companion plants interactively influence ladybird foraging pattern, and that their spatial arrangement can modulate the ability of such key predators to find their prey. KEYWORDS attract-and-reward, foraging behaviour, integrated pest management, methyl salicylate, natural enemies, semiochemicals,synomone How to cite this article: Jaworski CC, Xiao D, Xu Q, et al. Varying the spatial arrangement of synthetic herbivore-induced plant volatiles and companion plants to improve conservation biological control. J Appl Ecol. 2019;00:1-13.

1. Mammalian herbivores can strongly influence nitrogen (N) cycling and herbivore urine could be a central component of the N cycle in grazed ecosystems. Despite its potential role for ecosystem productivity and functioning, the fate of N derived from urine has rarely been investigated in grazed ecosystems. 2. This study explored the fate of ¹⁵N-enriched urea in tundra sites that have been either lightly or intensively grazed by reindeer for more than 50 years. We followed the fate of the ¹⁵N applied to the plant canopy, at 2 weeks and 1 year after tracer addition, in the different ecosystem N pools. 3. ¹⁵N-urea was rapidly incorporated in cryptogams and in above-ground parts of vascular plants, while the soil microbial pool and plant roots sequestered only a marginal proportion. Furthermore, the litter layer constituted a large sink for the ¹⁵N-urea, at least in the short term, indicating a high biological activity in the litter in the first phases of organic matter 15 layer and high immobilization decomposition. 4. Mosses and lichens still constituted the largest sink for the ¹⁵N-urea 1 year after tracer addition at both levels of grazing intensity demonstrating their large ability to capture and retain N from urine. Despite large fundamental differences in their traits, deciduous and evergreen shrubs were just as efficient as graminoids in taking up the ¹⁵N-urea. The total recovery of ¹⁵N-urea was lower in the intensively grazed sites, suggesting that reindeer reduce ecosystem N retention. 5. Synthesis. The rapid incorporation of the applied ¹⁵N-urea indicates that arctic plants can take advantage of a pulse of incoming N from urine. In addition, δ¹⁵N values of all taxa in the heavily grazed sites converged towards the δ¹⁵N values for urine, bringing further evidence that urine is an important N source for plants in grazed tundra ecosystems.

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission.

Insect pests cause significant global agricultural damage and lead to major financial and environmental costs. Crops contain intrinsic defenses to protect themselves from such pests, including a wide array of specialized secondary metabolite-based defense chemicals. These chemicals can be induced upon attack (phytoalexins) or are constitutive (phytoanticipins), and can have a direct impact on the pests or be used indirectly to attract their natural enemies. They form part of a global arms race between the crops and their insect pests, with the insects developing methods of suppression, avoidance, detoxification, or even capture of their hosts defensive chemicals. Harnessing and optimizing the chemical defense capabilities of crops has the potential to aid in the continuing struggle to enhance or improve agricultural pest management. Such strategies include breeding for the restoration of defense chemicals from ancestral varieties, or cross-species transfer of defense metabolite production.

1. Wind is an important abiotic factor that influences an array of biological processes, but it is rarely considered in studies on plant-herbivore interactions. 2. Here, we tested whether wind exposure could directly or indirectly affect the performance of two insect herbivores, Plutella xylostella and pieris brassicae, feeding on Brassica nigra plants. 3. In a greenhouse study using a factorial design, B. nigra plants were exposed to different wind regimes generated by fans before and after caterpillars were introduced on plants in an attempt to separate the effects of direct and indirect wind exposure on herbivores. 4. Wind exposure delayed flowering, decreased plant height and increased leaf concentrations of amino acids and glucosinolates. 5. Plant-mediated effects of wind on herbivores, that is effects of exposure of plants to wind prior to herbivore feeding, were generally small. However, development time of both herbivores was extended and adult body mass of P. xylostella was reduced when they were directly exposed to wind. By contrast, wind-exposed adult P. brassicae butterflies were significantly larger, revealing a trade-off between development time and adult size. 6. Based on these results, we conducted a behavioural experiment to study preference by an avian predator, the great tit (Parus major) for last instar P. brassicae caterpillars on plants that were exposed to either control (no wind) or wind (fan-exposed) treatments. Tits captured significantly more caterpillars on still than on wind-exposed plants. 7. Our results suggest that P. brassicae caterpillars are able to perceive the abiotic environment and to trade off the costs of extended development time against the benefits of increased size depending on the perceived risk of predation mediated by wind exposure. Such adaptive phenotypic plasticity in insects has not yet been described in response to wind exposure.

1. Anthropogenic disturbance and climate change might negatively affect the ecosystem services provided by mutualistic networks. However, the effects of such forces remain poorly characterized. They may be especially important in dry forests, which (1) experience chronic anthropogenic disturbances (CADs) as human populations exploit forest resources, and (2) are predicted to face a 22% decline in rainfall under climate change. 2. In this study, we investigated the separate and combined effects of CADs and rainfall levels on the specialization of mutualistic networks in the Caatinga, a seasonally dry tropical forest typical of north-eastern Brazil. More specifically, we examined interactions between plants bearing extrafloral nectaries (EFNs) and ants. We analysed whether differences in network specialization could arise from environmentally mediated variation in the species composition, namely via the replacement of specialist by generalist species. 3. We characterized these ant-plant networks in 15 plots (20 × 20 m) that varied in CAD intensity and mean annual rainfall. We quantified CAD intensity by calculating three indices related to the main sources of disturbance in the Caatinga: livestock grazing (LG), wood extraction (WE) and miscellaneous resource use (MU). We determined the degree of ant-plant network specialization using four metrics: generality, vulnerability, interaction evenness and H₂'. 4. Our results indicate that CADs differentially influenced network specialization: we observed positive, negative, and neutral responses along LG, MU and WE gradients, respectively. The pattern was most pronounced with LG. Rainfall also shaped network specialization, markedly increasing it. While LG and rainfall were associated with changes in network species composition, this trend was not related to the degree of species specialization. This result suggests that shifts in network specialization might be related to changes in species behaviour, not species composition. 5. Our study highlights the vulnerability of such dry forest ant-plant networks to climate change. Moreover, dry forests experience highly heterogeneous anthropogenic disturbances, creating a geographic mosaic of selective forces that may shape the co-evolution of interactions between ants and EFN-bearing plants.

A \"landscape of fear\" (LOF) is a map that describes continuous spatial variation in an animal's perception of predation risk. The relief on this map reflects, for example, places that an animal avoids to minimize risk. Although the LOF concept is a potentially unifying theme in ecology that is often invoked to explain the ecological and conservation significance of fear, little is known about the daily dynamics of an LOF. Despite theory and data to the contrary, investigators often assume, implicitly or explicitly, that an LOF is a static consequence of a predator's mere presence within an ecosystem. We tested the prediction that an LOF in a large-scale, free-living system is a highly dynamic map with \"peaks\" and \"valleys\" that alternate across the diel (24-h) cycle in response to daily lulls in predator activity. We did so with extensive data from the case study of Yellowstone elk (Cervus elaphus) and wolves (Canis lupus) that was the original basis for the LOF concept. We quantified the elk LOF, defined here as spatial allocation of time away from risky places and times, across nearly 1,000-km² of northern Yellowstone National Park and found that it fluctuated with the crepuscular activity pattern of wolves, enabling elk to use risky places during wolf downtimes. This may help explain evidence that wolf predation risk has no effect on elk stress levels, body condition, pregnancy, or herbivory. The ability of free-living animals to adaptively allocate habitat use across periods of high and low predator activity within the diel cycle is an underappreciated aspect of animal behavior that helps explain why strong antipredator responses may trigger weak ecological effects, and why an LOF may have less conceptual and practical importance than direct killing.

Summary The ecological function of secondary metabolites in plant defence against herbivores is well established, but their role in plant–pollinator interactions is less obvious. Nectar is the major reward for pollinators, so the occurrence of defence chemicals in the nectar of many species is unexpected. However, increasing evidence supports a variety of potential benefits for both plant and pollinator from these compounds. Beneficial effects may include: (i) mediating specialization in plant–pollinator interactions, (ii) protecting nectar from robbery or larceny and (iii) microbial activity including preservation of nutrients in nectar from degradation and reduction in disease levels in pollinators. Secondary metabolites in nectar can be toxic or repellent to flower visitors, but equally they can go undetected or even make nectar more apparent or attractive. These biological effects are concentration dependent, so must be considered at a range of ecologically relevant doses. For example, caffeine occurs in nectar and improves honeybee memory for odours associated with food rewards, which enhances pollen transfer at naturally occurring concentrations but is repellent to honeybees at higher concentrations. This review synthesizes evidence from recent literature that supports selection for secondary metabolites in floral nectar as an adaptation that drives the co‐evolution between plants and their pollinators. However, their presence in nectar could still simply be a consequence of their defensive role elsewhere in the plant (pleiotropy). We highlight the need for more studies demonstrating measurable benefits to the plant, the importance of exposure levels and effects on target species beyond the current emphasis on alkaloids and bees. A Lay Summary is available for this article. Lay Summary