Changes in herbivore control in arable fields by detrital subsidies depend on predator species and vary in space

Prey from the decomposer subsystem may help sustain predator populations in arable fields. Adding organic residues to agricultural systems may therefore enhance pest control. We investigated whether resource addition (maize mulch) strengthens aboveground trophic cascades in winter wheat fields. Evaluating the flux of the maize-borne carbon into the food web after 9 months via stable isotope analysis allowed differentiating between prey in predator diets originating from the above- and belowground subsystems. Furthermore, we recorded aphid populations in predator-reduced and control plots of no-mulch and mulch addition treatments. All analyzed soil dwelling species incorporated maize-borne carbon. In contrast, only 2 out of 13 aboveground predator species incorporated maize carbon, suggesting that these 2 predators forage on prey from the above- and belowground systems. Supporting this conclusion, densities of these two predator species were increased in the mulch addition fields. Nitrogen isotope signatures suggested that these generalist predators in part fed on Collembola thereby benefiting indirectly from detrital resources. Increased density of these two predator species was associated by increased aphid control but the identity of predators responsible for aphid control varied in space. One of the three wheat fields studied even lacked aphid control despite of mulch-mediated increased density of generalist predators. The results suggest that detrital subsidies quickly enter belowground food webs but only a few aboveground predator species include prey out of the decomposer system into their diet. Variation in the identity of predator species benefiting from detrital resources between sites suggest that, depending on locality, different predator species are subsidised by prey out of the decomposer system and that these predators contribute to aphid control. Therefore, by engineering the decomposer subsystem via detrital subsidies, biological control by generalist predators may be strengthened.     

Generalist predators in organically and conventionally managed grass‐clover fields: implications for conservation biological control

Organically managed agroecosystems rely in part on biological control to prevent pest outbreaks. Generalist predators (Araneae, Carabidae and Staphylinidae) are a major component of the natural enemy community in agroecosystems. We assessed the seasonal dynamics of major generalist predator groups in conventionally and organically managed grass–clover fields that primarily differed by fertilisation strategy. We further established an experiment, manipulating the abundant wolf spider genus Pardosa, to identify the importance of these predators for herbivore suppression in the same system and growth period. Organic management significantly enhanced ground‐active spider numbers early and late in the growing season, with potentially positive effects of plant cover and non‐pest decomposer prey. However, enhancing spider numbers in the field experiment did not improve biological control in organically managed grass–clover fields. Similar to the survey results, reduced densities of Pardosa had no short‐term effect on any prey taxa; however, spider guild structure changed in response to Pardosa manipulation. In the presence of fewer Pardosa, other ground‐running spiders were more abundant; therefore, their impact on herbivore numbers may have been elevated, possibly cancelling increases in herbivore numbers because of reduced predation by Pardosa. Our results indicate positive effects of organic farming on spider activity density; however, our survey data and the predator manipulation experiment failed to find evidence that ground‐running spiders reduced herbivore numbers. We therefore suggest that a positive impact of organic fertilisers on wolf spiders in grass–clover agroecosystems may not necessarily improve biological control when compared with conventional farming.     

Intraguild interference and biocontrol effects of generalist predators in a winter wheat field

Arable land typically harbours communities of polyphagous invertebrate natural enemies, among them numerous soil-surface dwelling predators such as ground beetles (Carabidae) and spiders (Lycosidae, Linyphiidae). Numbers of these predators were experimentally manipulated in a winter wheat field in order to study the predation impact of a generalist predator assemblage on herbivorous insects, the possible interferences among the predators concerned, and subsequent effects on wheat plant parameters. Removing ground beetles doubled numbers of Lycosidae indicative of intraguild interference between these two predator groups. Aphid densities were highest in carabid removal plots implying a substantial predation impact of ground beetles on the pest population. The predation impact of ground beetles was strongest earlier and disappeared later in the season. In mid-season, at intermediate aphid densities, the combined impact of carabid beetles and spiders appeared to be responsible for the reduction in aphid abundance. This result was probably due to a biomass effect rather than to a synergistic effect of the predator community. Thysanoptera decreased when spiders were removed (perhaps because spiders were preying on a predator of thrips), while Cicadellidae and Delphacidae showed no effect at all. The rise of aphid numbers in carabid removal plots corresponded to an increase in protein content of the wheat grains, while other plant parameters such as plant numbers and grain mass were not affected. In conclusion, this study provided field evidence for intraguild interference among generalist ground predators in arable land. Despite this interference the polyphagous predator community was able to depress numbers of aphids in winter wheat, a result cascading down to plant quality parameters.     

Arable weeds in organically managed wheat fields foster carabid beetles by resource- and structure-mediated effects

Arable weeds in organically managed fields may foster arthropod generalist predators by the provision of shelter and favorable microclimate (structure-mediated effects) and the provision of additional animal and floral food resources (resource-mediated effects). In three organically managed winter wheat fields in Central Germany, we investigated the impact of weed removal and introduction of artificial weed-like structure on the activity density and species richness of carabid beetles with respect to trophic groups, microclimatic conditions, and densities of potential prey. Removal of weeds reduced both carabid activity density and species richness but did not affect trophic group composition. The decline in carabid activity density was dampened by the addition of artificial structure. Mean daily surface temperature and light intensity were significantly lower under weeds and artificial plants than under wheat plants alone. Weed removal reduced the abundance of leafhoppers and true bugs, but the response was inconsistent across fields. We conclude that the presence of arable weeds in organically managed wheat fields fosters carabid activity density and species richness via resource-mediated effects, such as a higher availability of weed-borne resources (e.g. seeds and pollen) and herbivorous prey. Structure-mediated effects (altering the microclimate) add to this positive effect. The presence of weeds in organically managed wheat fields enhances carabid activity density and diversity and needs to be integrated into future management strategies for natural enemy conservation.     

Aphid cards – Useful model for assessing predation rates or bias prone nonsense?

Predation on pest organisms is an essential ecosystem function supporting yields in modern agriculture. However, assessing predation rates is intricate, and they can rarely be linked directly to predator densities or functions. We tested whether sentinel prey aphid cards are useful tools to assess predation rates in the field. Therefore, we looked at aphid cards of different sizes on the ground level as well as within the vegetation. Additionally, by trapping ground-dwelling predators, we examined whether obtained predation rates could be linked to predator densities and traits. Predation rates recorded with aphid cards were independent of aphid card size. However, predation rates on the ground level were three times higher than within the vegetation. We found both predatory carabid activity densities as well as community weighted mean body size to be good predictors for predation rates. Predation rates obtained from aphid cards are stable over card type and related to predator assemblages. Aphid cards, therefore, are a useful, efficient method for rapidly assessing the ecosystem function predation. Their use might especially be recommended for assessments on the ground level and when time and resource limitations rule out more elaborate sentinel prey methods using exclosures with living prey animals.     

Subsidy from the detrital food web, but not microhabitat complexity, affects the role of generalist predators in an aboveground herbivore food web

The activity and density of generalist predators, such as carabid beetles, rove beetles and spiders, may increase in response to: (1) increased availability of prey from the belowground subsystem and/or (2) enhanced complexity of aboveground vegetation. Organic farming practices support decomposer populations and enhance habitat complexity due to an increased weed density. A response by generalist predators to such below‐ or aboveground changes could affect predation rates on herbivores in the aboveground food web. We tested this hypothesis in a replicated field experiment conducted in a winter wheat field, where increased predator activity could lead to improved control of herbivorous pests. In a crossed design, we increased and lowered densities of decomposer prey, and manipulated vegetation complexity using artificial plants in order to examine the effect of structural complexity in isolation from effects of plant‐attracted additional prey. Isotomid Collembola exhibited lowest activity‐densities (AD) in plots treated with soil insecticide and had gradually increasing AD in untreated plots and plots receiving detrital subsidies. Carabid beetles and cursorial spiders did not respond to increased availability of isotomid prey, and they unexpectedly displayed higher AD in the structurally less‐complex plots. Aphid density mirrored the positive response of isotomids to detrital subsidies, suggesting that aphids benefited from reduced predation due to predators switching to abundant prey in the decomposer subsystem. The absence of a numerical response by surface‐active predators apparently strengthened this indirect effect of isotomids on aphids. Our results suggest that indirect predator‐mediated prey‐prey interactions can reduce beneficial effects of detrital subsidies on pest suppression. We further demonstrated that generalist predators may not per se benefit from structural complexity. Both results document the challenges associated with management practices that support generalist predators, as these measures may not necessarily improve herbivore suppression.     

Cascading effects of belowground predators on plant communities are density‐dependent

Soil food webs comprise a multitude of trophic interactions that can affect the composition and productivity of plant communities. Belowground predators feeding on microbial grazers like Collembola could decelerate nutrient mineralization by reducing microbial turnover in the soil, which in turn could negatively influence plant growth. However, empirical evidences for the ecological significance of belowground predators on nutrient cycling and plant communities are scarce. Here, we manipulated predator density (Hypoaspis aculeifer: predatory mite) with equal densities of three Collembola species as a prey in four functionally dissimilar plant communities in experimental microcosms: grass monoculture (Poa pratensis), herb monoculture (Rumex acetosa), legume monoculture (Trifolium pratense), and all three species as a mixed plant community. Density manipulation of predators allowed us to test for density‐mediated effects of belowground predators on Collembola and lower trophic groups. We hypothesized that predator density will reduce Collembola population causing a decrease in nutrient mineralization and hence detrimentally affect plant growth. First, we found a density‐dependent population change in predators, that is, an increase in low‐density treatments, but a decrease in high‐density treatments. Second, prey suppression was lower at high predator density, which caused a shift in the soil microbial community by increasing the fungal: bacterial biomass ratio, and an increase of nitrification rates, particularly in legume monocultures. Despite the increase in nutrient mineralization, legume monocultures performed worse at high predator density. Further, individual grass shoot biomass decreased in monocultures, while it increased in mixed plant communities with increasing predator density, which coincided with elevated soil N uptake by grasses. As a consequence, high predator density significantly increased plant complementarity effects indicating a decrease in interspecific plant competition. These results highlight that belowground predators can relax interspecific plant competition by increasing nutrient mineralization through their density‐dependent cascading effects on detritivore and soil microbial communities.     

Control of aphids on wheat by generalist predators: effects of predator density and the presence of alternative prey

There is evidence for both positive and negative effects of generalist predators on pest populations and the various reasons for these contrasting observations are under debate. We studied the influence of a generalist predator, Pardosa lugubris (Walckenaer) (Araneae: Lycosidae), on an aphid pest species, Rhopalosiphum padi (L.) (Hemiptera: Aphididae; low food quality for the spider), and its host plant wheat, Triticum spec. (Poaceae). We focused on the role of spider density and the availability of alternative prey, Drosophila melanogaster Meigen (Diptera: Drosophilidae; high food quality). The presence of spiders significantly affected plant performance and aphid biomass. Alternative prey and spider density strongly interacted in affecting aphids and plants. High spider density significantly improved plant performance but also at low spider density plants benefited from spiders especially in the presence of alternative prey. The results suggest that generalist arthropod predators may successfully reduce plant damage by herbivores. However, their ability to control prey populations varies with predator nutrition, the control of low-quality prey being enhanced if alternative higher-quality prey is available.     

植被边界带对相邻麦田地表步甲和蜘蛛分布及蚜虫发生的影响

通过管理半自然生境提高农田中天敌多样性及农田害虫的生物控制效率是当前生态农业研究的基本问题之一。为探讨不同类型半自然植被边界带对相邻麦田地表天敌的发生以及对小麦蚜虫生物控制潜力的影响,在北京顺义区赵全营镇采用陷阱法调查了不同类型植被边界带（人工林地、人工种植草带、自然演替草带）相邻的小麦田中,距离边界0,15 m和30 m处步甲和蜘蛛两类重要地表天敌的多度分布及小麦田蚜虫发生情况,同时分析了边界带植被群落结构对步甲和蜘蛛群落分布的影响。结果表明：人工林地相邻的小麦田中维持了显著较高的步甲、蜘蛛和蚜虫发生密度;不同类型植被边界带相邻小麦田中步甲和蜘蛛群落组成显著不同,植被边界带宽度以及乔木层盖度是影响相邻麦田步甲和蜘蛛群落分布的主要因子;距离植被边界带不同取样界面处,不同迁移方向的步甲和蜘蛛的活动密度无显著性差异;与林地相邻的麦田中,蜘蛛活动密度从农田边界向农田内部处依次降低,与人工种植草带以及自然演替草带相邻的麦田中,农田边界处蜘蛛的活动密度显著高于农田内部。研究显示,半自然植被边界带与麦田交界处维持了较高的蜘蛛活动密度,并具有向农田内部扩散的趋势,然而,并非高天敌密度就意味着低害虫发生率,深入研究天敌群落与害虫发生关系及其影响因素是未来天敌保护和害虫生物防治的重要方面。同时,因为不同类型植被边界带维持的天敌群落具有高度异质性,未来的研究需要充分考虑不同类型植被边界带对天敌维持的作用,以及不同类型生境界面地表天敌的活动规律。     

A field experiment on the effectiveness of spiders and carabid beetles as biocontrol agents in soybean

1 Spiders and carabid beetles are abundant generalist predators that prey upon insect pests of soybean. A field experiment was conducted to determine the impact of spiders and carabids on soybean yield. Prior to planting, three 7 × 7 m plots were fenced in order to reduce spider and carabid immigration. Carabids that emerged within the plots were not removed, but spiders that ballooned into these predator‐reduction plots or that entered by climbing the fence were removed by pitfall trapping and searching the vegetation. Three unmanipulated, unfenced plots served as the control treatment. 2 Densities of spiders on soybean vegetation, and activity‐densities of spiders and carabids determined by pitfall trapping, were c. 75% lower in the spider‐carabid reduction treatment than in control plots. Despite clear differences between treatments in numbers and activity of these major generalist predators, the weight of soybeans harvested did not differ between control and spider‐carabid reduction plots. 3 Paralleling the absence of an effect of predator reduction on soybean yield was the absence of any significant difference between treatments in densities of whiteflies (Aleyrodidae), leafhoppers (Cicadellidae), thrips (Thysanoptera), Lepidoptera larvae and herbivorous Coleoptera. 4 Our experiment provides no evidence that spiders and carabid beetles at ambient densities affect soybean yield. Low populations of pest species or low predation pressure on soybean pests by spiders and carabids at the ambient densities of this experiment could be responsible for this result.     

Collembola as alternative prey sustaining spiders in arable ecosystems: prey detection within predators using molecular markers

Collembola comprise a major source of alternative prey to linyphiid spiders in arable fields, helping to sustain and retain these predators as aphid control agents within the crop. Polymerase chain reaction primers were developed for the amplification, from spider gut samples, of DNA from three of the most abundant species of Collembola in wheat crops in Europe, namely Isotoma anglicana, Lepidocyrtus cyaneus and Entomobrya multifasciata. The primers amplified fragments of the mitochondrial cytochrome oxidase subunit I (COI) gene and were designed following alignment of comparable sequences for a range of predator and prey species. Each of the primer pairs proved to be species-specific to a Collembola species, amplifying DNA fragments from 211 to 276 base pairs in length. Following consumption of a single collembolan, prey DNA was detectable in 100% of spiders after 24 h of digestion. We report the first use of DNA-based techniques to detect predation by arthropods on natural populations of prey in the field. All three species of Collembola were consumed by the spiders. By comparing the ratios of the Collembola species in the field with the numbers of spiders that gave positive results for each of those species, it was possible to demonstrate that the spiders were exercising prey choice. Overall, a single target species of Collembola was eaten by 48% of spiders while a further 16% of spiders contained DNA from two different species of Collembola. Preference was particularly evident for I. anglicana, the species most frequently found in spider guts yet the least numerous of the three target species in the field.     

Organic farming affects the potential of a granivorous carabid beetle to control arable weeds at local and landscape scales

相似文献   

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Aggregation of polyphagous predators in response to multiple prey: ladybirds (Coleoptera: Coccinellidae) foraging in alfalfa

The spatial distribution of polyphagous predators may often reflect the integration of aggregative responses to local densities of multiple species of prey, and as such may have consequences for the indirect linkages among the prey sharing these predators. In a factorial field experiment in which we manipulated local prey densities within a field of alfalfa in Utah (USA), we tested whether aphidophagous ladybirds would aggregate not only in response to their primary aphid prey, but also in response to an abundant alternative prey, the alfalfa weevil (Hypera postica [Gyllenhal]). Native North American ladybirds (primarily Hippodamia convergens Guerin and H. quinquesignata quinquesignata [Kirby]) responded only to spatial variation in aphid density. In contrast, the introduced ladybird, Coccinella septempunctata L., aggregated also at local concentrations of the weevil late in the experiment when weevil density was high and aphid density was relatively low throughout all experimental plots. The results support the hypothesis that C. septempunctata is more responsive than are native ladybirds to the availability of alternative prey in alfalfa, which may account in part for the displacement of native ladybirds from alfalfa by the introduced species as aphid numbers have declined. The differing responses of the native and introduced ladybirds to spatial patterns of the alternative prey underscore the importance of extending the study of predator aggregation to understand better how polyphagous predators distribute themselves in response to spatial patterns of multiple species of potential prey.     

A pragmatic approach to the analysis of diets of generalist predators: the use of next‐generation sequencing with no blocking probes

Predicting whether a predator is capable of affecting the dynamics of a prey species in the field implies the analysis of the complete diet of the predator, not simply rates of predation on a target taxon. Here, we employed the Ion Torrent next‐generation sequencing technology to investigate the diet of a generalist arthropod predator. A complete dietary analysis requires the use of general primers, but these will also amplify the predator unless suppressed using a blocking probe. However, blocking probes can potentially block other species, particularly if they are phylogenetically close. Here, we aimed to demonstrate that enough prey sequence could be obtained without blocking probes. In communities with many predators, this approach obviates the need to design and test numerous blocking primers, thus making analysis of complex community food webs a viable proposition. We applied this approach to the analysis of predation by the linyphiid spider Oedothorax fuscus in an arable field. We obtained over two million raw reads. After discarding the low‐quality and predator reads, the libraries still contained over 61 000 prey reads (3% of the raw reads; 6% of reads passing quality control). The libraries were rich in Collembola, Lepidoptera, Diptera and Nematoda. They also contained sequences derived from several spider species and from horticultural pests (aphids). Oedothorax fuscus is common in UK cereal fields, and the results showed that it is exploiting a wide range of prey. Next‐generation sequencing using general primers but without blocking probes provided ample sequences for analysis of the prey range of this spider and proved to be a simple and inexpensive approach.     

Monoclonal antibodies reveal changes in predator efficiency with prey spatial pattern

Spatially explicit predator–prey interactions can alter the predatory potential of natural enemies augmented through conservation biological control. To test hypotheses regarding such interactions and predatory efficiency, we used a combination of molecular techniques and mark–release–recapture to study the foraging behaviour of a generalist carabid predator, Poecilus cupreus , in response to spatial patterns of its cereal aphid prey ( Metapolophium dirhodum and Sitobion avenae ). Beetle and aphid numbers were measured across two grids of sampling locations, within which aphid spatial pattern had been manipulated to generate patchy and more homogenous distributions. Aphid consumption was measured by enzyme-linked immunosorbent assays (ELISA) of beetle gut contents, using an aphid-specific monoclonal antibody. Movement and distribution patterns suggest that P. cupreus does not aggregate at, nor instigate prey-taxis within, aphid patches. However, more than two-thirds of the 2169 P. cupreus tested by ELISA had consumed aphids and the proportion of beetles containing aphid proteins was positively related to aphid density. Against expectation, the proportion of predators feeding on aphids was greatest where prey were homogenously distributed, and this was attributed to the loss of partial refuges for prey in aphid patches. The functional value of this type of uniform foraging strategy is ideally suited to early colonization of the crop habitat, when aphid numbers are low, before populations build up and form strong spatial patterns.     

Detection of secondary predation by PCR analyses of the gut contents of invertebrate generalist predators

Predation by generalist predators is difficult to study in the field because of the complex effects of positive and negative interactions within and between predator species and guilds. Predation can be monitored by molecular means, through identification of prey DNA within predators. However, polymerase chain reaction (PCR) amplification of prey DNA from predators cannot discriminate between primary and secondary predation (hyperpredation), in which one predator feeds on another that has recently eaten the target prey. Here we quantify, for the first time, the potential error caused by detection of prey DNA following secondary predation, using an aphid-spider-carabid model. First, the aphid Sitobion avenae was fed to the spider Tenuiphantes tenuis and the carabid Pterostichus melanarius, and the postconsumption detection periods, for prey DNA within predators, were calculated. Aphids were then fed to spiders and the spiders to carabids. Aphid DNA was detected in the predators using primers that amplified 245- and 110-bp fragments of the mitochondrial cytochrome oxidase I gene. Fragment size and predator sex had no significant effect on detection periods. Secondary predation could be detected for up to 8 h, when carabids fed on spiders immediately after the latter had consumed aphids. Beetles tested positive up to 4 h after eating spiders that had digested their aphid prey for 4 h. Clearly, the extreme sensitivity of PCR makes detection of secondary predation more likely, and the only reliable answer in future may be to use PCR to identify, in parallel, instances of intraguild predation.     

Evaluation of lycosid,micryphantid and linyphiid spiders as predators ofRhopalosiphum padi (Hom.: Aphididae) and their functional response to prey density-laboratory experiments

Laboratory experiments were performed to determine the potential of dominant spider species in winter wheat in Germany,Erigone atra (Blackwall),Lepthyphantes tenuis (Blackwall) andPardosa agrestis (Westring) adults and youngs, in suppressing the population ofRhopalosiphum padi (L.) on wheat plants and their functional response to different aphid densities. The presence of spiders significantly caused between 34 and 58% reduction in aphid population development on wheat plants compared to the aphid population in the absence of spiders. The functional response curves for these spiders as predators ofR. padi seem to descrive a typical type II functional response with the prey consumed increasing to a plateau as aphid densities increased. Prey killed without eating was linear on prey density.