Predators control pests and increase yield across crop types and climates: a meta-analysis.

Pesticides have well-documented negative consequences to control crop pests, and natural predators are alternatives and can provide an ecosystem service as biological control agents. However, there remains considerable uncertainty regarding whether such biological control can be a widely applicable solution, especially given ongoing climatic variation and climate change. Here, we performed a meta-analysis focused on field studies with natural predators to explore broadly whether and how predators might control pests and in turn increase yield. We also contrasted across studies pest suppression by a single and multiple predators and how climate influence biological control. Predators reduced pest populations by 73% on average, and increased crop yield by 25% on average. Surprisingly, the impact of predators did not depend on whether there were many or a single predator species. Precipitation seasonality was a key climatic influence on biological control: as seasonality increased, the impact of predators on pest populations increased. Taken together, the positive contribution of predators in controlling pests and increasing yield, and the consistency of such responses in the face of precipitation variability, suggest that biocontrol has the potential to be an important part of pest management and increasing food supplies as the planet precipitation patterns become increasingly variable.

The World Spider Trait database: a centralized global open repository for curated data on spider traits.

Spiders are a highly diversified group of arthropods and play an important role in terrestrial ecosystems as ubiquitous predators, which makes them a suitable group to test a variety of eco-evolutionary hypotheses. For this purpose, knowledge of a diverse range of species traits is required. Until now, data on spider traits have been scattered across thousands of publications produced for over two centuries and written in diverse languages. To facilitate access to such data, we developed an online database for archiving and accessing spider traits at a global scale. The database has been designed to accommodate a great variety of traits (e.g. ecological, behavioural and morphological) measured at individual, species or higher taxonomic levels. Records are accompanied by extensive metadata (e.g. location and method). The database is curated by an expert team, regularly updated and open to any user. A future goal of the growing database is to include all published and unpublished data on spider traits provided by experts worldwide and to facilitate broad cross-taxon assays in functional ecology and comparative biology. Database URL:https://spidertraits.sci.muni.cz/.

Intensive site preparation for reforestation wastes multi-trophic biodiversity potential in commercial oak woodlands.

Lowland oak forests host high biodiversity throughout Europe, and to maintain their high biodiversity potential we need to understand how current silviculture practices influence these ecosystems. The most common harvest approach in oak woodlands remains clear-cutting followed by site preparation for reforestation. The site preparation can vary in intensity and possibly affect forest biodiversity in various ways. We studied the impact of site preparation intensity on ground-dwelling arthropod predators and detritivores in commercial oak woodlands. The intensive management included mechanical site preparation with soil milling, stump removal, and herbicide application. The extensive management consisted of natural succession after reforestation with only such slight disturbances as regular hand-provided mowing of vegetation. We recorded 120 spider species, 92 species of ground beetles, 10 species of centipedes, 17 species of millipedes, and 7 species of woodlice, including a relatively large number of threatened species. We found that intensive post-logging management strongly homogenized the habitat structure, and this led to low multi-trophic taxonomic and functional diversity in comparison to that under extensive management. At sites with extensive management, there was not only high functional diversity but also high functional redundancy. Species of conservation concern almost vanished from clearings under intensive management. The high multi-trophic diversity and functional redundancy indicate that extensive site preparation may enhance ecosystem multi-functionality, including primary productivity and ecosystem resilience. Extensive post-harvest management is therefore strongly preferable, and it is also economically more feasible. Intensive post-harvest management should be prohibited in lowland forests within transition zones to localities under nature protection or in natural oak forests.

Disturbance by invasive pathogenic fungus alters arthropod predator-prey food-webs in ash plantations.

According to the disturbance-succession theory, natural disturbances support biodiversity and are expected to increase the complexity of food-webs in forest ecosystems by opening canopies and creating a heterogeneous environment. However, a limited number of studies have investigated the impact of disturbance by invasive pathogenic species and succession on arthropod predator-prey food-webs in forest ecosystems. Hymenoscyphus fraxineus is a pathogenic fungus of ash trees that is invasive in Europe and causes massive dieback, mainly of the common ash Fraxinus excelsior across its native range. Here we investigated how this pathogenic fungus affects food-webs of web-building spiders and their prey in understorey vegetation of ash plantations. In 23 young and middle-aged ash plantations that were distributed along a gradient of infestation by H. fraxineus (29%-86% infestation), we measured the vegetation structure (canopy openness, shrub coverage, herb/grass coverage), the trait composition of local spider communities (web type, body size), the prey availability and the prey intercepted by spider webs. We then evaluated the multivariate prey composition (prey type, body size) and network properties. Hymenoscyphus fraxineus opened the ash tree canopy, which resulted in denser shrub coverage. The dense shrub vegetation changed the composition of web types in local spider communities and increasing fungus infestation resulted in reduced mean body size of spiders. Infestation by H. fraxineus reduced the availability of predaceous Coleoptera and increased the availability of herbivorous Coleoptera as potential prey. The mean body size of captured prey and the per capita capture rates of most prey groups decreased with increasing fungus infestation. Hymenoscyphus fraxineus infestation indirectly reduced the complexity in bipartite networks and the trophic functional complementarity in local web-building spider communities. The plantation age affected the vegetation structure but did not affect the studied food-webs. Forest disturbance by the invasive pathogen affected four trophic levels (plant-herbivore-coleopteran intermediate predator-top predator web-building spiders) and, contrary to the disturbance-succession theory, disturbance by the fungus simplified the web-building spider-prey food-webs. The results support the view that H. fraxineus represents a threat to the biodiversity and ecosystem functioning in the simplified ecosystems of ash plantations.

The pest-specific effects of glyphosate on functional response of a wolf spider.

Although glyphosate is widely used for weed pest control, it might have negative side effects on natural enemies. Wolf spiders are one of the most representative predators found on soybean crops in Uruguay, preying on a wide variety of potential pests. However, the sublethal effects that pesticides might have on this group have been poorly explored for South American species. Herein, we explored the sublethal effects of glyphosate on the functional response of the wolf spider Hogna cf. bivittata against three potential pest insects, namely ant (Acromyrmex sp.), caterpillar (Anticarsia gemmatalis), and cricket (Miogryllus sp.). We contaminated residually adult females of the species Hogna cf. bivittata with glyphosate (Roundup®) and compared their functional response against non-contaminated spiders. We did not observe any mortality during the study. We found that overall Hogna cf. bivittata showed a functional response type II against crickets and caterpillars but no functional response to ants. Contaminated spiders killed less ants and caterpillars in comparison to the control group, probably as a consequence of the irritating effects of glyphosate. We did not observe differences in functional response to crickets at the evaluated densities, probably as a consequence of the low capture rate against this prey. Although glyphosate does not specifically target spiders, it might have negative sublethal effects on native predators such as Hogna cf. bivittata. Further studies should explore effect of glyphosate on other native predators from South American crops.

Transformational Mimicry in a Myrmecomorphic Spider.

Species that are Batesian mimics during postembryonic development shift between mimetic models as they grow in size. However, it has not yet been tested whether these successive mimetic phenotypes are similarly protected from predators. Early-instar phenotypes could represent an inaccurate phenotype or an accurate phenotype because of selection from different predators. Here, we tested the hypotheses of transformational Batesian mimicry in the ant-mimicking jumping spider Leptochestes berolinensis. We quantified the mimetic accuracy of different ontogenetic stages to potential ant models by using a multitrait approach. We measured movement, body profile, body size, and coloration. Analysis revealed adults to be more accurate mimics than juveniles. Adults were similar to smaller morphs of Camponotus or Lasius ants, whereas juveniles were more similar to Lasius and Colobopsis ants. We tested whether predators, mantises, and Pisaura spiders were deceived by mimics after having experience with ant models. These predators never captured any ant or a mimic but always captured the nonmyrmecomorphic spider. We conclude that L. berolinensis is a Batesian mimic of ants undergoing transformational mimicry, with all stages being accurate mimics.

Synergistic effects of glyphosate formulation herbicide and tank-mixing adjuvants on Pardosa spiders.

Glyphosate-based herbicides are the world's most consumed agrochemicals, and they are commonly used in various agroecosystems, including forests, as well as in urban zones and gardens. These herbicides are sold as formulations containing adjuvants. Other tank-mixing adjuvants (most often surfactants) are commonly added to these formulations prior to application. According to the manufacturers of agrochemicals, such tank mixes (as these are known in agronomic and horticultural practice) have modified properties and perform better than do the herbicides as used alone. The effects of these tank mixes on the environment and on beneficial arthropods are almost unknown. Therefore, we studied whether a herbicide formulation mixed with adjuvant has modified effects on one of the most common genera of ground-dwelling wolf spiders vis-à-vis the herbicide formulation and adjuvants themselves. Specifically, we studied the synergistic effect in the laboratory on the predatory activity (represented by the number of killed flies) of wolf spiders in the genus Pardosa after direct treatment using the glyphosate-based herbicide formulation Roundup klasik Pro®, Roundup klasik Pro® in a mixture with the surfactant Wetcit®, Roundup klasik Pro® in a mixture with the surfactant Agrovital®, and the surfactants alone. We found that pure surfactants as well as herbicide-and-surfactants tank mixes significantly decrease the predatory activity of Pardosa spiders in the short term even as Roundup klasik Pro® did not itself have any such effect. Our results support the hypothesis that plant protection tank mixes may have modified effect on beneficial arthropods as compared to herbicide formulations alone. Therefore, testing of pesticide tank mixes is highly important, because it is these tank mixes that are actually applied to the environment.

An updated perspective on spiders as generalist predators in biological control.

The role of generalist predators in biological control remains controversial as they may not only reduce pest populations but also disrupt biocontrol exerted by other natural enemies. Here, we focus on spiders as a model group of generalist predators. They are among the most abundant and most diverse natural enemies in agroecosystems. We review their functional traits that influence food-web dynamics and pest suppression at organisational levels ranging from individuals to communities. At the individual and population levels, we focus on hunting strategy, body size, life stage, nutritional target, and personality (i.e., consistent inter-individual differences in behaviour). These functional traits determine the spider trophic niches. We also focus on the functional and numerical response to pest densities and on non-consumptive effects of spiders on pests. At the community level, we review multiple-predator effects and effect of alternative prey on pest suppression. Evidence for a key role of spiders in pest suppression is accumulating. Importantly, recent research has highlighted widespread non-consumptive effects and complex intraguild interactions of spiders. A better understanding of these effects is needed to optimize biocontrol services by spiders in agroecosystems.

Behavioral predictability in a lynx spider is interactively influenced by mean behavior, prey density, and an insecticide.

Behavioral predictability, i.e., short-term intra-individual variability under relatively constant environmental conditions, has only recently begun to gain attention. It is unknown, however, whether predictability of individuals with distinct mean behavior changes differently as a response to ecological factors such as resource availability. Moreover, the response might be affected by anthropogenic contaminants that are ubiquitous in the environment and that can affect animals' variability in behavior. Here, we investigated the relationship between mean predatory activity and predictability in predatory activity along a prey density gradient in the lynx spider Oxyopes lineatipes. We further examined how this relationship is influenced by insecticides, azadirachtin, and a plant extract from Embelia ribes. We found that all studied variables affected the predictability. In the control and Embelia treatments, that did not differ significantly, the predictability decreased with increasing prey density in a mean behavior-specific way. Individuals with low mean predatory activity were relatively less predictable than were those with high activity from low to moderate prey densities but more predictable at high prey densities. Azadirachtin altered this pattern and the individuals with low predatory activity were less predictable than were those with high predatory activity along the whole gradient of prey density. Our results show that predictability can change along an environmental gradient depending on a mean behavior. The relative predictability of the individuals with distinct mean behavior can depend on the value of the environmental gradient. In addition, this relationship can be affected by anthropogenic contaminants such as pesticides.

Specific parasites indirectly influence niche occupation of non-hosts community members.

The coexistence of phylogenetically related species is an attractive topic because of the potentially intense interspecific competition. The most often investigated mechanisms mediating coexistence of these species are environmental filtering and niche partitioning. However, the role of other factors, such as species-specific parasites, is still poorly understood. Along the riparian understory of a tropical forest, we explored niche occupation and coexistence between Chrysso intervales and Helvibis longicauda, two related syntopic web-building spiders. We also investigated the effect of H. longicauda mortality induced by a specific fungus parasite, Gibellula pulchra, on the dynamic of C. intervales spatial distribution. Coexistence was mediated mainly by a fine-scale horizontal spatial segregation. H. longicauda built webs almost exclusively close to the river, while C. intervales occupied adjacent areas (10-20 m away from margins). We also found differentiation in other niche dimensions that might allow coexistence, such as in plants occupied, height of web placement, width of leaves used for thread attachment and phenology. H. longicauda mortality caused by fungi was higher during winter than in summer. Consequently, the abundance of C. intervales increased at distances close to the river, indicating competitive release through a density-mediated indirect effect. This demonstrates how non-competitive specific-antagonists can indirectly affect other non-hosts competing community members and influence their spatial distribution in fine-scale ranges. We suggest that environmental filtering restricts H. longicauda to cooler regions; niche partitioning leads populations to occupy different vertical strata and competitive exclusion precludes C. intervales to reach river margins, generating an unusual horizontal zonation pattern.

Insecticides alter prey choice of potential biocontrol agent Philodromus cespitum (Araneae, Philodromidae).

Even though pesticides can have various sublethal effects on behaviour of biocontrol agents, no study to date has investigated the effects of pesticides on the prey choice of generalist predators. Prey choice of generalist predators is among key factors determining the predation pressure they exert on pests, because it influences fitness of predators and consequently their densities and per capita capture rate. Here, we investigated the effect of Integro (a.i. methoxyfenozide) and SpinTor 480 Sc (a.i. spinosad) on prey choice and predatory activity of the spider Philodromus cespitum, which is known significantly to reduce hemipteran and dipteran pests in fruit orchards. We compared the prey preferences of philodromids between the psyllid pest Cacopsylla pyri (Hemiptera, Psyllidae) and beneficial Theridion sp. spiders in laboratory experiments. We found that both pesticides altered the prey preferences of philodromids. While the philodromids in a control treatment preferred theridiid spiders over the psyllid pest, philodromids in the pesticide treatments exhibited no significant prey preferences. The changes in prey preferences were caused by increased predation on the psyllids, while the predation on the theridiid remained similar. We suggest that the changes in prey preferences might theoretically be due to 1) impaired sensory systems, 2) altered taste, and/or 3) altered mobility. In combination with other studies finding reduced predation on fruit flies in P. cespitum after exposure to SpinTor, our results indicate that the sublethal effects of pesticides on predatory behaviour of generalist predators can depend on prey type and/or prey community composition.

Influence of no-tillage versus tillage system on common vole (Microtus arvalis) population density.

BACKGROUND: While the 'no-tillage' management system generally improves soil properties and helps to control arthropod pests, it may also intensify crop infestation by the common vole (Microtus arvalis Pallas). In this study, we evaluated the impact of soil management (no-tillage or tillage), crop and previous crop (winter wheat or winter rape), and season (spring or autumn) on common vole density using data from the Common Vole Monitoring Programme undertaken by the Plant Protection Service of the Czech Republic between 2000 and 2009. RESULTS: Models predicted low mean values of vole infestation across management types, crops, and seasons. The untilled fields hosted significantly more voles than the tilled fields in spring but not in autumn. More common voles were found in winter rape than in winter wheat during both seasons. CONCLUSION: Recent studies suggest that no-tillage management is more profitable than tillage management as a result of its positive impact on soil properties and pest control. During periods of high vole infestation, however, tillage may constitute an alternative strategy for reducing yield losses. © 2017 Society of Chemical Industry.

The Behavioral Type of a Top Predator Drives the Short-Term Dynamic of Intraguild Predation.

Variation in behavior among individual top predators (i.e., the behavioral type) can strongly shape pest suppression in intraguild predation (IGP). However, the effect of a top predator's behavioral type-namely, foraging aggressiveness (number of killed divided by prey time) and prey choosiness (preference degree for certain prey type)-on the dynamic of IGP may interact with the relative abundances of top predator, mesopredator, and pest. We investigated the influence of the top predator's behavioral type on the dynamic of IGP in a three-species system with a top predator spider, a mesopredator spider, and a psyllid pest using a simulation model. The model parameters were estimated from laboratory experiments and field observations. The top predator's behavioral type altered the food-web dynamics in a context-dependent manner. The system with an aggressive/nonchoosy top predator, without prey preferences between pest and mesopredator, suppressed the pest more when the top predator to mesopredator abundance ratio was high. In contrast, the system with a timid/choosy top predator that preferred the pest to the mesopredator was more effective when the ratio was low. Our results show that the behavioral types and abundances of interacting species need to be considered together when studying food-web dynamics, because they evidently interact. To improve biocontrol efficiency of predators, research on the alteration of their behavioral types is needed.

Different hunting strategies of generalist predators result in functional differences.

The morphological, physiological, and behavioural traits of organisms are often used as surrogates for actual ecological functions. However, differences in these traits do not necessarily lead to functional differences and/or can be context-dependent. Therefore, it is necessary to explicitly test whether the surrogates have general ecological relevance. To investigate the relationship between the hunting strategies of predators (i.e., how, where, and when they hunt) and their function, we used euryphagous spiders as a model group. We used published data on the diet composition of 76 spider species based on natural prey and laboratory prey acceptance experiments. We computed differences in the position and width of trophic niches among pairs of sympatrically occurring species. Pairs were made at different classification levels, ranked according to the dissimilarity in their hunting strategies: congeners, confamiliars (as phylogenetic proxies for similarity in hunting strategy), species from the same main class of hunting strategy, from the same supra-class, and from different supra-classes. As for niche position computed from the natural prey analyses, species from the same class differed less than species from different classes. A similar pattern was obtained from the laboratory studies, but the congeners differed less than the species from the same classes. Niche widths were most similar among congeners and dissimilar among species from different supra-classes. Functional differences among euryphagous spiders increased continuously with increasing difference in their hunting strategy. The relative frequency of hunting strategies within spider assemblages can, therefore, influence the food webs through hunting strategy-specific predator-prey interactions.

Sublethal effect of agronomical surfactants on the spider Pardosa agrestis.

In addition to their active ingredients, pesticides contain also additives - surfactants. Use of surfactants has been increasing over the past decade, but their effects on non-target organisms, especially natural enemies of pests, have been studied only very rarely. The effect of three common agrochemical surfactants on the foraging behavior of the wolf spider Pardosa agrestis was studied in the laboratory. Differences in short-term, long-term, and overall cumulative predatory activities were investigated. We found that surfactant treatment significantly affected short-term predatory activity but had no effect on long-term predatory activity. The surfactants also significantly influenced the cumulative number of killed prey. We also found the sex-specific increase in cumulative kills after surfactants treatment. This is the first study showing that pesticide additives have a sublethal effect that can weaken the predatory activity of a potential biological control agent. More studies on the effects of surfactants are needed to understand how they affect beneficial organisms in agroecosystems.

Impact of Canopy Openness on Spider Communities: Implications for Conservation Management of Formerly Coppiced Oak Forests.

Traditional woodland management created a mosaic of differently aged patches providing favorable conditions for a variety of arthropods. After abandonment of historical ownership patterns and traditional management and the deliberate transformation to high forest after World War II, large forest areas became darker and more homogeneous. This had significant negative consequences for biodiversity. An important question is whether even small-scale habitat structures maintained by different levels of canopy openness in abandoned coppiced forest may constitute conditions suitable for forest as well as open habitat specialists. We investigated the effect of canopy openness in former traditionally coppiced woodlands on the species richness, functional diversity, activity density, conservation value, and degree of rareness of epigeic spiders. In each of the eight studied locations, 60-m-long transect was established consisting of five pitfall traps placed at regular 15 m intervals along the gradient. Spiders were collected from May to July 2012. We recorded 90 spider species, including high proportions of xeric specialists (40%) and red-listed threatened species (26%). The peaks of conservation indicators, as well as spider community abundance, were shifted toward more open canopies. On the other hand, functional diversity peaked at more closed canopies followed by a rapid decrease with increasing canopy openness. Species richness was highest in the middle of the canopy openness gradient, suggesting an ecotone effect. Ordinations revealed that species of conservation concern tended to be associated with sparse and partly opened canopy. The results show that the various components of biodiversity peaked at different levels of canopy openness. Therefore, the restoration and suitable forest management of such conditions will retain important diversification of habitats in formerly coppiced oak forest stands. We indicate that permanent presence of small-scale improvements could be suitable conservation tools to prevent the general decline of woodland biodiversity in the intensified landscape of Central Europe.

Phenotypic integration in a series of trophic traits: tracing the evolution of myrmecophagy in spiders (Araneae).

Several hypotheses have been put forward to explain the evolution of prey specificity (stenophagy). Yet little light has so far been shed on the process of evolution of stenophagy in carnivorous predators. We performed a detailed analysis of a variety of trophic adaptations in one species. Our aim was to determine whether a specific form of stenophagy, myrmecophagy, has evolved from euryphagy via parallel changes in several traits from pre-existing characters. For that purpose, we studied the trophic niche and morphological, behavioural, venomic and physiological adaptations in a euryphagous spider, Selamia reticulata. It is a species that is branching off earlier in phylogeny than stenophagous ant-eating spiders of the genus Zodarion (both Zodariidae). The natural diet was wide and included ants. Laboratory feeding trials revealed versatile prey capture strategies that are effective on ants and other prey types. The performance of spiders on two different diets - ants only and mixed insects - failed to reveal differences in most fitness components (survival and developmental rate). However, the weight increase was significantly higher in spiders on the mixed diet. As a result, females on a mixed diet had higher fecundity and oviposited earlier. No differences were found in incubation period, hatching success or spiderling size. S. reticulata possesses a more diverse venom composition than Zodarion. Its venom is more effective for the immobilisation of beetle larvae than of ants. Comparative analysis of morphological traits related to myrmecophagy in the family Zodariidae revealed that their apomorphic states appeared gradually along the phylogeny to derived prey-specialised genera. Our results suggest that myrmecophagy has evolved gradually from the ancestral euryphagous strategy by integrating a series of trophic traits.