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.     

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.     

Effect of spiders (Arachnida: Araneae) as predators of insect pest in alfalfa crops (Medicago sativa) (Fabaceae) in Argentina

Spiders are predators that reduce insect pest populations in agroecosystems. Trials were conducted to measure the selectivity against different insect preys, the daily consumption, effect of predators alone and together with a known number of preys, and the indirect effect of predators on vegetation. For this, experimental units (1 x 1m) were used covered with a fine plastic mesh. Misumenops pallidus, Oxyopes salticus and Araneus sp. were used as generalist predators, and aphids, weevils, locusts, chrysomelids and Lepidoptera larvae as their potential preys. Among the preys offered, the spiders preferred Lepidoptera larvae compared to the other two pests groups (weevils and aphids). The maximum consumption rate was of 93.33% for Lepidoptera larvae, 25.33% for aphids and 11.67% for weevils. The Q Index values for the three species of spiders showed a positive selectivity only for defoliating larvae. O. salticus showed the highest values of consumption rates while Rachiplusia nu was the most consumed. The maximum value of consumption in 24 hours was showed by O. salticus on R. nu (C) = 2.8. The association of several species of predatory spiders increased the total number of insects captured, and also showed that the addition of spiders caused a decrease in the number of leaves damaged by the effect of lepidopterous larvae.     

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.     

Selective predation by larval Agabus (Coleoptera: Dytiscidae) on mosquitoes: support for conservation-based mosquito suppression in constructed wetlands

1. Wetland insect predators can structure aquatic prey communities via selective predation, but receive considerably less attention than vertebrate predators. We conducted laboratory experiments to test selective predation by two species of larval dytiscid beetles ( Agabus ; Coleoptera: Dytiscidae) and the potential contribution of these beetles to suppression of mosquito populations in constructed wetlands.
2.  Agabus consumed copepods, ostracods and mosquito larvae in no-choice tests. When offered a choice, 76% of all prey consumed were mosquito larvae, indicating selective predation. Subsequent experiments revealed this preference was due to ease of capture of mosquito larvae over alternative prey.
3. Cannibalism and intraguild predation were common within and between species of Agabus , which may reduce the overall impact of the observed selective predation.
4.  Agabus larvae selectively preyed on mosquito larvae over alternative prey, which is not characteristic of some fish used as biological control agents for mosquitoes. Predator exclusion or similar experiments in the field could document how these results translate into a natural setting.
5. The findings of this study suggest developing mosquito suppression strategies focused on conservation of native wetland predators. These strategies are preferable to introducing non-native generalist predators, or applying pesticides.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Predation by generalist predators on the codling moth versus a closely‐related emerging pest the oriental fruit moth: a molecular analysis

• 1 Biological control by conservation of native natural enemies can, at its best, reduce the need for pesticides and prevent detrimental effects upon the environment. The present study investigated the role of ground‐active generalist predators as natural enemies of two tortricid pests in apple orchards.
• 2 Predation rates were compared on the well established codling moth Cydia pomonella and the emerging oriental fruit moth Grapholita molesta, which has recently switched hosts to apples.
• 3 The present study hypothesized that the ground‐active predators consumed the two tortricid pests in significant numbers without preference, and attacked the pests at different developmental stages.
• 4 Using diagnostic polymerase chain reaction on the gut contents of field‐caught ground‐active predators, no difference in predation rates was found on these two pests. Spiders were the most efficient predators of emergent adult moths in spring, whereas the carabid beetles, feeding on diapausing larvae, were important in the autumn.
• 5 The temporal complementarity between spiders and carabid beetles, attacking different stages of the pests at different times of year, highlights the need for diverse predator assemblages to optimize biological control.

     

Impact on soil-dwelling arthropods in citrus orchards of spraying horticultural mineral oil, carbaryl or methidathion

Abstract  Impacts of an n C24 horticultural mineral oil (HMO) and two synthetic insecticides (carbaryl and methidathion) on the abundance and species diversity of soil-dwelling arthropods were evaluated in two citrus orchards in the coastal region of New South Wales, Australia. In the first orchard, mature Valencia orange trees were sprayed in summer with one of HMO, carbaryl or methidathion delivered at low (2000 L/ha) or high (10 000 L/ha) volumes; the sprays were applied either once (February) or twice (December and February). HMO had no significant impact but the synthetic insecticides, irrespective of spray frequency or spray volume, significantly reduced the abundance and species diversity of the arthropods including springtails and generalist predators, such as spiders, staphylinid beetles and mesostigmatid mites. In the second orchard, blocks of Washington navel and Valencia orange trees were sprayed with either HMO or methidathion; sprays were applied twice, 2 weeks apart, in late summer (February), first at a rate of 6500 L/ha and second at 5500 L/ha. Results were similar to those in the first orchard, except that the effect of methidathion on carabid beetles and ants was not significant.     

Effects of kaolin on the composition of generalist predator assemblages and parasitism of Choristoneura rosaceana (Lep., Tortricidae) in apple orchards

Abstract:  In three apple orchards, we tested how the hydrophilic kaolin clay particle film Surround WP affected the diversity of generalist arthropod predator assemblages in orchard foliage and the parasitism of the pest species Choristoneura rosaceana (Harris) (obliquebanded leafroller) (Lep., Tortricidae). In two orchards, kaolin was applied to orchard foliage once a week for 4 weeks, between mid-June and mid-July in 2004 and 2005. In the third orchard kaolin was applied to foliage twice over 2 weeks in June 2004. We quantified the proportion of larvae C. rosaceana parasitized, larval populations, and the relative abundance and assemblage composition of generalist predators (spiders and insects) in the orchards. Kaolin altered the species composition of the generalist predator assemblages and reduced the relative abundances of certain generalist predators, most notably jumping and crab spiders (Salticidae and Philodromidae), assassin bugs (Reduviidae), ants (Formicidae) and coccinellids (Coccinellidae). In contrast, the relative abundances of web-spinning spiders (Araneidae, Dictynidae, Theridiidae) were not affected. Kaolin did not affect the proportion of parasitized C. rosaceana larvae, which ranged from 24% to 47% in control and kaolin treatments, or the relative proportions of parasitoid taxa. The kaolin formulation did not affect the abundance of C. rosaceana larvae, but in one orchard, kaolin did reduce the abundance of the combined numbers of C. rosaceana and another tortricid pest, Argyrotaenia velutiana . Although kaolin does not affect parasitism of C. rosaceana , it significantly changes the composition of generalist predator assemblages in orchard foliage.     

Laboratory and field tests of predation on the cereal leaf beetle,Oulema melanopus (Coleoptera: Chrysomelidae)

The cereal leaf beetle (CLB), Oulema melanopus (L.) (Coleoptera: Chrysomelidae), is an invasive pest in North America recently reported in the Canadian Prairies. We performed a series of laboratory assays to identify potential predators and a field study to quantify predation of CLB eggs. In no-choice Petri dish assays, ground beetles (Carabidae), rove beetles (Staphylinidae), and several common lady beetle species (Coccinellidae) were the most consistent predators of eggs and larvae. Nabis spp. (Hemiptera: Nabidae) and wolf spiders (Araneae: Lycosidae) consumed many larvae, but did not consume eggs. Hippodamia spp., Coccinella septempunctata (L.) (Coleoptera: Coccinellidae), and Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) also fed on CLB eggs on potted plants when an alternative food source was available, Sitobion avenae (Fabricius) (Hemiptera: Aphididae). In our field study, we found an average of 24.5% of sentinel eggs disappeared over a 24?h period, likely due to predation. Our results suggest that generalist predators can play an important role in the biological control of CLB, and warrant further study.     

Predator Interference and the Establishment of Generalist Predator Populations for Biocontrol

We conducted a field experiment to determine the extent to which interference among generalist predators limits their effectiveness as biocontrol agents. We manipulated immigration of a guild of actively hunting generalist ground predators, carabid beetles and lycosid spiders, by intercepting them as they attempted to enter fenced 50-m² vegetable gardens. Immigration was blocked, allowed at the mean rate measured at our field site, or doubled. Altered immigration rates were maintained through a spring garden of cabbage, bean, eggplant, and cucumber, followed by a summer garden of squash. We monitored densities of carabids and lycosids to discover if altering their immigration rate changed their densities in the plots. We also measured densities of other predators on the ground and in plant foliage, pest numbers, and vegetable yields. Doubling the immigration rate of carabids and lycosids approximately doubled the densities of carabids inside the plots, but did not increase lycosid densities. Increasing the rate of immigration of carabids and lycosids depressed densities of nonlycosid ground spiders. In the spring gardens, manipulation of carabid and lycosid immigration did not influence numbers of predators or herbivores in the foliage and did not affect vegetable productivity. In contrast, in the summer gardens, foliage-dwelling predators were lower, pest densities were marginally lower, and squash productivity was higher in the carabid and lycosid immigration plots compared to the no-immigration treatment. Doubling carabid and lycosid immigration rate never increased the magnitude of their effects on other predators, pests, or plant productivity. Predator interference limited lycosid establishment, reduced densities of other predator taxa, and apparently prevented a doubling of carabid densities from having an increased impact on pest numbers. Nevertheless, despite widespread effects of predator interference, allowing immigration of lycosids and carabids increased squash productivity.     

多种捕食性天敌控制麦蚜当量系统研究

通过饲养,明确了麦蚜主要天敌食蚜蝇类、瓢虫娄、草蛉类、蜘蛛类之中8种幼虫和6种成虫(蛛)的取食量。经校正当量系数模式等统计．建立了上述天敌控制麦蚜的当量系统。实例表明,该系统能准确测定复合天敌的控蚜效果,为促进麦蚜综合治理进展提供了一条依据。     

Manipulating carabid beetle abundance alters prey removal rates in corn fields

Generalist natural enemies such as carabid beetles have the potential to maintain a variety of pests below outbreak levels in annual crops. To assess the relationship between carabid beetle abundance and field rates of prey removal, we created plots surrounded by different boundaries that selectively affected dispersal of edaphic arthropods, primarily carabids. Three treatments were established: (1) naturally occurring communities, (2) augmented communities using ingress boundaries, and (3) reduced communities using egress boundaries. Selective boundaries altered carabid communities with minimal habitat alteration and without use of insecticides. Three times during the growing season, a fixed number of onion fly pupae were placed in plots to evaluate the impact of carabid abundance on predation rates. A combination of vertebrate and invertebrate exclosures allowed us to evaluate prey removal by invertebrates alone. In comparison to the no boundary treatment, carabids increased 54.2% and decreased 83.1% in plots surrounded by ingress and egress boundaries respectively. Predation rates were positively correlated with carabid abundance (r² = 0.70, p < 0.0001). Significantly more pupae were removed from exclosures allowing access to invertebrates alone than from total exclosures, suggesting that invertebrates represented an important group of predators. Laboratory trials tested the feeding potential of the four most abundant carabid species and showed that they readily consumed onion fly pupae, supporting our hypothesis that carabids were the main predators in field tests. This study corroborates and extends previous observations of the importance of carabid beetles as generalist predators of insect pests in agricultural fields.     

Predation by Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on immature Rhagoletis mendax Curran (Diptera: Tephritidae) in semi-field and field conditions

Practices that enhance abundance and diversity of generalist predators are often employed with the objective of improving biological control of insect pests. Ground beetles and other predators can prey on blueberry maggot, an important pest of blueberries, when mature larvae pupate in the ground. We conducted mesocosm and field experiments to determine if Pterostichus melanarius, a common predatory ground beetle, lowers maggot numbers in compost mulch or when predator and alternative prey abundances are manipulated. At background (field) densities of alternative prey, increasing densities of P. melanarius did not significantly reduce pest numbers in mesocosms containing compost or soil. When alternative prey were removed from compost, beetles reduced pest numbers by up to 35%. In field experiments, maggot numbers were higher when beetles and other predators were excluded from soil plots, but beetle exclusion had no effect in compost plots where both predator and alternative prey numbers were high. Our results indicate that there can be some reduction of blueberry maggot by P. melanarius and other potential predators when there are few alternative prey. However, despite attracting large numbers of predators compost mulch did not lead to a significant reduction in blueberry maggot; in fact, the high abundance of alternative food associated with compost appeared to interfere with beetle predation on blueberry maggot.     

A test of the hypothesis that a pathway of intraguild predation limits densities of a wolf spider

Abstract.  1. The high number of potential predatory interactions among the many arthropod generalist predators in terrestrial food webs makes exhaustive testing of interaction strengths by field experiments unfeasible. Thus, correlative patterns and laboratory observations of behaviour often form the basis of inferences about the strength of interaction pathways involving generalist predators (intraguild predation).
2. Previous research has revealed a negative correlation between survival of juvenile wolf spiders of the genus Schizocosa (Lycosidae) and densities of another abundant spider family, the Gnaphosidae.
3. Feeding trials in laboratory microcosms with a leaf-litter substrate revealed that gnaphosids prey on juvenile Schizocosa in a structurally complex habitat.
4. Gnaphosid densities were manipulated in two different field experiments, each conducted in a different year, in order to test directly the hypothesis that intraguild predation by gnaphosids limits densities of juvenile Schizocosa .
5. Reducing numbers of gnaphosids, and doubling their numbers to two times the mean natural density, had no impact on the survival of juvenile Schizocosa in either field experiment. This finding suggests that correlative patterns in nature and feeding trials in the laboratory may at times provide deceptively simple and potentially misleading generalisations about the strengths of interaction pathways in complex networks of generalist predators.     

Complementary effects of resident natural enemies on the suppression of the introduced moth Epiphyas postvittana

Generalist predators may disrupt or complement biological control by parasitoids. Past studies have examined how predators and parasitoids interact to affect aphid suppression, but more information is needed from other host taxa. Here, we explore the interactive effects of a spider (Cheiracanthium mildei) and a generalist parasitoid (Meteorus ictericus) on the light brown apple moth (Epiphyas postvittana), a recent introduction to North America. The spider negatively affected adult parasitoids in a field experiment, and reduced numbers of parasitized larvae in the laboratory. Nonetheless, the combined effects of parasitoids and spiders on larval mortality of the moth were additive. Percent parasitism was not affected by the presence of the spider in field or laboratory experiments, and results were similar when single or multiple larval instars of the moth were included. The spider’s lack of prey preference for unparasitized or parasitized larvae likely precluded any disruptive effects on parasitism. Results suggest that resident generalist parasitoids and predators can work in conjunction to hinder the invasion success of a novel herbivore prey species.     

Management intensity at field and landscape levels affects the structure of generalist predator communities

Agricultural intensification is recognised as a major driver of biodiversity loss in human-modified landscapes. Several agro-environmental measures at different spatial scales have been suggested to mitigate the negative impact of intensification on biodiversity and ecosystem services. The effect of these measures on the functional structure of service-providing communities remains, however, largely unexplored. Using two distinct landscape designs, we examined how the management options of organic farming at the field scale and crop diversification at the landscape level affect the taxonomic and functional structure of generalist predator communities and how these effects vary along a landscape complexity gradient. Organic farming as well as landscapes with longer and more diversified crop rotations enhanced the activity-density of spiders and rove beetles, but not the species richness or evenness. Our results indicate that the two management options affected the functional composition of communities, as they primarily enhanced the activity-density of functionally similar species. The two management options increased the functional similarity between spider species in regards to hunting mode and habitat preference. Organic farming enhanced the functional similarity of rove beetles. Management options at field and landscape levels were generally more important predictors of community structure when compared to landscape complexity. Our study highlights the importance of considering the functional composition of generalist predators in order to understand how agro-environmental measures at various scales shape community assemblages and ecosystem functioning in agricultural landscapes.     

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.     

Enhancing spider families and spider webs in Indian rice fields for conservation biological control,considering local and landscape management

Spiders are omnipresent, occurring in almost all terrestrial habitats. They are generalist predators and important for conservation biocontrol in agricultural fields, helping to reduce pesticide applications. In this study, we examined how spider families and spider web types in rice fields are related to local and landscape management. Samples were taken in differently managed rice fields adjoining either homegarden polycultures or banana monocultures. Furthermore, landscape structure, prey abundance, herb richness and cover were taken into account. The results showed that prey availability explained most of the variation in spiders and their web’s abundance, indicating that spiders in the rice fields can potentially contribute to pest control. Adjacent habitat had no effect on the spider populations, but maintaining fallow fields in the surrounding landscape seems to be a useful measure to promote Erigoninae in rice fields. There was no evidence that local management practices such as fertiliser and pesticide use had an impact on spider families, which appeared to be due to the low level of these inputs. Spider web sampling can complement spider sampling as it detects spiders hidden at the base of the rice tillers, which are likely to be missed by sweep netting. Additionally, tetragnathid webs are easy to observe and thus can be used as an indication for farmers not to spray pesticides as spiders are potentially controlling the pest species. Interviews with farmers made clear that many farmers in the study area showed their interest in management methods that promote biological pest control.     

Stable isotope analyses document intraguild predation in wolf spiders (Araneae: Lycosidae) and underline beneficial effects of alternative prey and microhabitat structure on intraguild prey survival

Intraguild predation (IGP) is common among generalist predators and an important issue in food web theory, because IGP may destabilise communities by increasing extinction of species. Also, IGP may interfere with the effectiveness of generalist predators as biological control agents. In general, occurrence of IGP in laboratory or field studies is inferred from abundance data or direct observations only. We investigated if tracing stable isotopes allows distinction between different types of predation and confirmation of IGP. Wolf spiders were chosen as model organisms for generalist predators; IGP of third instar A. cuneata on second instar P. palustris was investigated in a laboratory experiment. The availability of alternative prey and the complexity of the microhabitat were manipulated, since both factors are thought to facilitate coexistence of predators.
Stable isotope analysis documented predation of A. cuneata on P. palustris and predation on alternative prey by both juveniles. Both the presence of alternative prey and the availability of shelter reduced mortality of juvenile P. palustris during the first week. During the second week mortality increased in complex structure without alternative prey presumably due to enhanced activity and cannibalism among starving P. palustris . Thus, microhabitat complexity and prey abundance may foster coexistence of wolf spiders in the field.
In conclusion, stable isotope analysis was proven a powerful tool to investigate animal behaviour without direct observation. The method allowed disentangling predator feeding behaviour when more than one type of prey was present.