Imperfect diet choice reduces the performance of a predatory mite.

Two mutually unexclusive hypotheses prevail in the theory of nutritional ecology: the balanced diet hypothesis states that consumers feed on different food items because they have complementary nutrient and energy compositions. The toxin-dilution hypothesis poses that consumers feed on different food items to dilute the toxins present in each. Both predict that consumers should not feed on low-quality food when ample high-quality food forming a complete diet is present. We investigated the diet choice of Phytoseiulus persimilis, a predatory mite of web-producing spider mites. It can develop and reproduce on single prey species, for example the spider mite Tetranychus urticae. A closely related prey, T. evansi, is of notorious bad quality for P. persimilis and other predator species. We show that juvenile predators feeding on this prey have low survival and do not develop into adults. Adults stop reproducing and have increased mortality when feeding on it. Feeding on a mixed diet of the two prey decreases predator performance, but short-term effects of feeding on the low-quality prey can be partially reversed by subsequently feeding on the high-quality prey. Yet, predators consume low-quality prey in the presence of high-quality prey, which is in disagreement with both hypotheses. We suggest that it is perhaps not the instantaneous reproduction on single prey or mixtures of prey that matters for the fitness of predators, but that it is the overall reproduction by a female and her offspring on an ephemeral prey patch, which may be increased by including inferior prey in their diet.

Prey exploitation and dispersal strategies vary among natural populations of a predatory mite.

When predators commonly overexploit local prey populations, dispersal drives the dynamics in local patches, which together form a metapopulation. Two extremes in a continuum of dispersal strategies are distinguished: the "Killer" strategy, where predators only start dispersing when all prey are eliminated, and the "Milker" strategy, in which predator dispersal occurs irrespective of prey availability. Theory shows that the Milker strategy is not evolutionarily stable if local populations are well connected by dispersal. Using strains of the predatory mite Phytoseiulus persimilis, collected from 11 native populations from coastal areas in Turkey and Sicily, we investigated whether these two strategies occur in nature. In small wind tunnels, we measured dispersal rates and population dynamics of all populations in a system consisting of detached rose leaves, spider mites (Tetranychus urticae) as prey, and P. persimilis. We found significant variation in the exploitation and dispersal strategies among predator populations, but none of the collected strains showed the extreme Killer or Milker strategy. The results suggest that there is genetic variation for prey exploitation and dispersal strategies. Thus, different dispersal strategies in the Milker-Killer continuum may be selected for under natural conditions. This may affect the predator-prey dynamics in local populations and is likely to determine persistence of predator-prey systems at the metapopulation level.

Spatial patterns generated by simultaneous cooperation and exploitation favour the evolution of altruism.

In kin selection theory in the evolution of social behaviours, the relatedness between interacting individuals is influenced by the spatial structure of the population. It is generally considered that in the 'viscous' population competition among individuals tends to suppress the evolution of altruism. We consider that more complex interactions produce specific spatial patterns in the presence of competitive interaction, which could alter the process of kin selection in a given space. Here, we theoretically studied the joint evolution of altruism and resource exploitation in a spatially structured population. The simulations indicated that joint evolution can result in self-organisation of regularly arranged cluster structures, which creates a new 'level of selection', and significantly promotes the evolution of altruism through a promotion of kin selection. The analysis also suggested synergetic effects of the joint evolution of two traits, including an evolutionary suicide of the population.

Predatory interactions between prey affect patch selection by predators.

ABSTRACT: When predators can use several prey species as food sources, they are known to select prey according to foraging efficiency and food quality. However, interactions between the prey species may also affect prey choice, and this has received limited attention. The effect of one such interaction, intraguild predation between prey, on patch selection by predators was studied here. The predatory mite Neoseiulus californicus preys on young larvae of the western flower thrips Frankliniella occidentalis and on all stages of the two-spotted spider mite Tetranychus urticae. The two prey species co-occur on several plant species, on which they compete for resources, and western flower thrips feed on eggs of the spider mites. A further complicating factor is that the thrips can also feed on the eggs of the predator. We found that performance of the predatory mite was highest on patches with spider mites, intermediate on patches with spider mites plus thrips larvae and lowest on patches with thrips larvae alone. Patch selection and oviposition preference of predators matched performance: predators preferred patches with spider mites over patches with spider mites plus thrips. Patches with thrips only were not significantly more attractive than empty patches. We also investigated the cues involved in patch selection and found that the attractiveness of patches with spider mites was significantly reduced by the presence of cues associated with killed spider mite eggs. This explains the reduced attractiveness of patches with both prey. Our results point at the importance of predatory interactions among prey species for patch selection by predators. SIGNIFICANCE STATEMENT: Patch selection by predators is known to be affected by factors such as prey quality, the presence of competitors and predators, but little is known on the effects of interactions among prey species present on patch selection. In this paper, we show that patch selection by a predator is affected by such interactions, specifically by the feeding of one prey species on eggs of the other.

The role of web sharing, species recognition and host-plant defence in interspecific competition between two herbivorous mite species.

When competing with indigenous species, invasive species face a problem, because they typically start with a few colonizers. Evidently, some species succeeded, begging an answer to the question how they invade. Here, we investigate how the invasive spider mite Tetranychus evansi interacts with the indigenous species T. urticae when sharing the solanaceous host plant tomato: do they choose to live together or to avoid each other's colonies? Both species spin protective, silken webs on the leaf surfaces, under which they live in groups of con- and possibly heterospecifics. In Spain, T. evansi invaded the non-crop field where native Tetranychus species including T. urticae dominated. Moreover, T. evansi outcompetes T. urticae when released together on a tomato plant. However, molecular plant studies suggest that T. urticae benefits from the local down-regulation of tomato plant defences by T. evansi, whereas T. evansi suffers from the induction of these defences by T. urticae. Therefore, we hypothesize that T. evansi avoids leaves infested with T. urticae whereas T. urticae prefers leaves infested by T. evansi. Using wild-type tomato and a mutant lacking jasmonate-mediated anti-herbivore defences, we tested the hypothesis and found that T. evansi avoided sharing webs with T. urticae in favour of a web with conspecifics, whereas T. urticae more frequently chose to share webs with T. evansi than with conspecifics. Also, T. evansi shows higher aggregation on a tomato plant than T. urticae, irrespective of whether the mites occur on the plant together or not.

Why do males choose heterospecific females in the red spider mite?

In some species, males readily show courtship behaviour towards heterospecific females and even prefer them to females of their own species. This behaviour is generally explained by indiscriminate mating to acquire more mates, but may partly be explained by male mate preference mechanisms that have developed to choose among conspecific females, as male preference for larger females causes mating with larger heterospecific females. Recently, we found that males of the red spider mite, Tetranychus evansi collected from Spain (invasive population), prefer to mate with females of the two-spotted spider mite, T. urticae rather than with conspecific females. In spider mites, mate preference for non-kin individuals has been observed. Here, we investigated if T. evansi males collected from the area of its origin (Brazil) also show preference for heterospecific females. Secondly, we investigated if mate preference of T. evansi males for heterospecific females is affected by their relatedness to conspecific females which are offered together with heterospecific females. We found that mate preference for heterospecific females exists in Brazilian T. evansi, suggesting that the preference for heterospecific females is not a lack of evolved premating isolation with an allopatric species. We found that T. evansi males showed lower propensity to mate with heterospecific females when alternative females were non-kin in the two iso-female lines collected from Brazil. However, the effect of relatedness on male mate preference was not significant. We discuss alternative hypotheses explaining why T. evansi males prefer to mate with T. urticae females.

No adaptation of a herbivore to a novel host but loss of adaptation to its native host.

Most herbivorous arthropods are host specialists and the question is which mechanisms drive the evolution of such specialization. The theory of antagonistic pleiotropy suggests that there is a trade-off between adaptation of herbivores to a novel host and their native host. The mutation accumulation hypothesis proposes that herbivores on a novel host lose their adaptation to the native host through the accumulation of mutations with negligible effects on performance on the novel host. Experimental evidence for either of the two hypotheses is scarce. We compared the fitness of two sympatric moth strains from an introduced host and a native host. The strain from the novel host did not perform better on this host than the strain from the native host. The strain from the novel host performed less well on the native host than did the strain from the native host. Hence, selection on the novel host did not result in noticeable gain in performance, but adaptation to the native host was lost. These results are more readily explained by the mutation-accumulation hypothesis than by the trade-off hypothesis.

Cry-wolf signals emerging from coevolutionary feedbacks in a tritrophic system.

For a communication system to be stable, senders should convey honest information. Providing dishonest information, however, can be advantageous to senders, which imposes a constraint on the evolution of communication systems. Beyond single populations and bitrophic systems, one may ask whether stable communication systems can evolve in multitrophic systems. Consider cross-species signalling where herbivore-induced plant volatiles (HIPVs) attract predators to reduce the damage from arthropod herbivores. Such plant signals may be honest and help predators to identify profitable prey/plant types via HIPV composition and to assess prey density via the amount of HIPVs. There could be selection for dishonest signals that attract predators for protection from possible future herbivory. Recently, we described a case in which plants release a fixed, high amount of HIPVs independent of herbivore load, adopting what we labelled a 'cry-wolf' strategy. To understand when such signals evolve, we model coevolutionary interactions between plants, herbivores and predators, and show that both 'honest' and 'cry-wolf' types can emerge, depending on the assumed plant-herbivore encounter rates and herbivore population density. It is suggested that the 'cry-wolf' strategy may have evolved to reduce the risk of heavy damage in the future. Our model suggests that eco-evolutionary feedback loops involving a third species may have important consequences for the stability of this outcome.

Distribution and oviposition site selection by predatory mites in the presence of intraguild predators.

When intraguild prey and intraguild predators feed and reproduce in the same habitat and relatively immobile juveniles are the vulnerable stage, predation risk depends on oviposition site selection by the adult females. We studied how the availability of oviposition sites affected the distribution of two predatory mite species, Neoseiulus cucumeris (Oudemans) and Iphiseius degenerans (Berlese), over two patches that both contained food. The two plant-inhabiting species feed on pollen and thrips, prey on each other's juveniles, and prefer to oviposit on hairy parts of the leaf. When an artificial oviposition site was provided on one of two connected patches, both predator species strongly preferred this patch. Whereas the distributions of adults and eggs of N. cucumeris over the two patches were not affected by the presence of heterospecifics, the proportions of adults and eggs of I. degenerans on the patch with an oviposition site were reduced by the presence of N. cucumeris. A similar change in distribution was induced by cues of N. cucumeris on the oviposition site, without these mites being present. Hence, intraguild prey can weaken the strength of intraguild predation through patch selection, which in turn may promote coexistence of the two predator species.

Plant Resources as a Factor Altering Emergent Multi-Predator Effects.

Multiple predator effects (MPEs) can modify the strength of pest regulation, causing positive or negative deviations from those that are predicted from independent effects of isolated predators. Despite increasing evidence that omnivory can shape predator-prey interactions, few studies have examined the impact of alternative plant food on interactions between multiple predators. In the present study, we examined the effects and interactions of two omnivorous mirids, Μacrolophus pygmaeus and Nesidiocoris tenuis, on different densities of their aphid prey, Myzus persicae. Prey were offered to the to single or pairs of mirid predator individuals, either conspecific or heterospecific on a leaf, while simultaneously adding or excluding a flower as an alternative food resource. Data were compared with calculated expected values using the multiplicative risk model and the substitutive model. We showed that predation of aphids was reduced in the presence of the alternative flower resource in treatments with single M. pygmaeus individuals, but not with single N. tenuis individuals. When the predators had access only to prey, the effects of multiple predation, either conspecific or heterospecific, were additive. The addition of an alternative plant resource differently affected MPEs depending on the nature of predator pairings. Predation risk was increased in conspecific M. pygmaeus treatments at intermediate prey densities, whereas it was reduced in conspecific N. tenuis treatments at high prey densities. Observations of foraging behaviour concerning the location of conspecific pairings revealed that M. pygmaeus individuals showed a clear tendency to reside mainly in the flower, whereas N. tenuis individuals were found to reside at different posts in the dish. We suggest that the competition between omnivorous predators may be mediated through the diversity of their plant feeding preferences, which directly affects the strength of MPEs. Consequently, the preferences of the interacting predators for different plant resources should be considered in studies evaluating the outcomes of MPEs.

Beyond Predation: The Zoophytophagous Predator Macrolophus pygmaeus Induces Tomato Resistance against Spider Mites.

Many predatory insects that prey on herbivores also feed on the plant, but it is unknown whether plants affect the performance of herbivores by responding to this phytophagy with defence induction. We investigate whether the prior presence of the omnivorous predator Macrolophus pygmaeus (Rambur) on tomato plants affects plant resistance against two different herbivore species. Besides plant-mediated effects of M. pygmaeus on herbivore performance, we examined whether a plant defence trait that is known to be inducible by herbivory, proteinase inhibitors (PI), may also be activated in response to the interactions of this predator with the tomato plant. We show that exposing tomato plants to the omnivorous predator M. pygmaeus reduced performance of a subsequently infesting herbivore, the two-spotted spider mite Tetranychus urticae Koch, but not of the greenhouse whitefly Trialeurodes vaporariorum (Westwood). The spider-mite infested tomato plants experience a lower herbivore load, i.e., number of eggs deposited and individuals present, when previously exposed to the zoophytophagous predator. This effect is not restricted to the exposed leaf and persists on exposed plants for at least two weeks after the removal of the predators. The decreased performance of spider mites as a result of prior exposure of the plant to M. pygmaeus is accompanied by a locally and systemically increased accumulation of transcripts and activity of proteinase inhibitors that are known to be involved in plant defence. Our results demonstrate that zoophytophagous predators can induce plant defence responses and reduce herbivore performance. Hence, the suppression of populations of certain herbivores via consumption may be strengthened by the induction of plant defences by zoophytophagous predators.

Incomplete premating and postmating reproductive barriers between two parapatric populations of a social spider mite.

Closely related species with overlapping distributions often show premating reproductive barriers to avoid hybridization. Stigmaeopsis miscanthi (Saito) is a social spider mite infesting Chinese silver grass, and the species consists of two parapatric groups with frequent contacts within the contact zone. They differ in male-male aggressiveness, male morphology, female diapause traits, and life history parameters. There is incomplete but strong post-mating reproductive isolation between the two groups, and their DNA sequences are slightly different, suggesting that they diverged recently. In this study, we investigated premating reproductive barriers. We found that females from different groups frequently shared nest webs, indicating no barriers in the phase of nest establishment. However, inside nests, males from either group showed less courtship behaviour to females of the other group and they copulated less frequently with them when compared to females of the same group. However, the premating reproductive barrier was incomplete and asymmetric. Females of one group frequently resisted courtship by males from the other group, but females of the other group did not. We conclude that some gene flow may occur in the contact zone between the two groups.

Spider mites suppress tomato defenses downstream of jasmonate and salicylate independently of hormonal crosstalk.

Plants respond to herbivory by mounting a defense. Some plant-eating spider mites (Tetranychus spp.) have adapted to plant defenses to maintain a high reproductive performance. From natural populations we selected three spider mite strains from two species, Tetranychus urticae and Tetranychus evansi, that can suppress plant defenses, using a fourth defense-inducing strain as a benchmark, to assess to which extent these strains suppress defenses differently. We characterized timing and magnitude of phytohormone accumulation and defense-gene expression, and determined if mites that cannot suppress defenses benefit from sharing a leaf with suppressors. The nonsuppressor strain induced a mixture of jasmonate- (JA) and salicylate (SA)-dependent defenses. Induced defense genes separated into three groups: 'early' (expression peak at 1 d postinfestation (dpi)); 'intermediate' (4 dpi); and 'late', whose expression increased until the leaf died. The T. evansi strains suppressed genes from all three groups, but the T. urticae strain only suppressed the late ones. Suppression occurred downstream of JA and SA accumulation, independently of the JA-SA antagonism, and was powerful enough to boost the reproductive performance of nonsuppressors up to 45%. Our results show that suppressing defenses not only brings benefits but, within herbivore communities, can also generate a considerable ecological cost when promoting the population growth of a competitor.

Defense suppression benefits herbivores that have a monopoly on their feeding site but can backfire within natural communities.

BACKGROUND: Plants have inducible defenses to combat attacking organisms. Hence, some herbivores have adapted to suppress these defenses. Suppression of plant defenses has been shown to benefit herbivores by boosting their growth and reproductive performance. RESULTS: We observed in field-grown tomatoes that spider mites (Tetranychus urticae) establish larger colonies on plants already infested with the tomato russet mite (Aculops lycopersici). Using laboratory assays, we observed that spider mites have a much higher reproductive performance on russet mite-infested plants, similar to their performance on the jasmonic acid (JA)-biosynthesis mutant def-1. Hence, we tested if russet mites suppress JA-responses thereby facilitating spider mites. We found that russet mites manipulate defenses: they induce those mediated by salicylic acid (SA) but suppress those mediated by JA which would otherwise hinder growth. This suppression of JA-defenses occurs downstream of JA-accumulation and is independent from its natural antagonist SA. In contrast, spider mites induced both JA- and SA-responses while plants infested with the two mite species together display strongly reduced JA-responses, yet a doubled SA-response. The spider mite-induced JA-response in the presence of russet mites was restored on transgenic tomatoes unable to accumulate SA (nahG), but russet mites alone still did not induce JA-responses on nahG plants. Thus, indirect facilitation of spider mites by russet mites depends on the antagonistic action of SA on JA while suppression of JA-defenses by russet mites does not. Furthermore, russet mite-induced SA-responses inhibited secondary infection by Pseudomonas syringae (Pst) while not affecting the mite itself. Finally, while facilitating spider mites, russet mites experience reduced population growth. CONCLUSIONS: Our results show that the benefits of suppressing plant defenses may diminish within communities with natural competitors. We show that suppression of defenses via the JA-SA antagonism can be a consequence, rather than the cause, of a primary suppression event and that its overall effect is determined by the presence of competing herbivores and the distinct palette of defenses these induce. Thus, whether or not host-defense manipulation improves an herbivore's fitness depends on interactions with other herbivores via induced-host defenses, implicating bidirectional causation of community structure of herbivores sharing a plant.

The endosymbionts Wolbachia and Cardinium and their effects in three populations of the predatory mite Neoseiulus paspalivorus.

Whereas endosymbiont-induced incompatibility is known to occur in various arthropod taxa, such as spider mites, insects and isopods, it has been rarely reported in plant-inhabiting predatory mites (Acari: Phytoseiidae). Recent cross-breeding studies with the phytoseiid mite Neoseiulus paspalivorus De Leon revealed a complete post-mating reproductive isolation between specimens collected from three geographic origins-Northeast Brazil (South America), Benin and Ghana (West Africa)-even though they are morphologically similar. We carried out a study to assess to what extent these populations exhibit genetic differences and whether endosymbionts are involved in the incompatibility. First, we used the mitochondrial cytochrome oxidase I (COI) gene to assess genetic diversity among the three populations. Second, we used a PCR-based method to check for the presence of Wolbachia and/or Cardinium in these populations, and we determined their phylogenetic relationships using specific primers for Wolbachia and Cardinium 16S rDNA genes. Third, we also conducted a test using an antibiotic (tetracycline) in an attempt to eliminate the symbionts and evaluate their effects on the reproductive compatibility of their host. Based on the DNA sequences of their COI genes, specimens of the three populations appear to be genetically similar. However, the 16S rDNA gene sequences of their associated endosymbionts differed among the three populations: the Benin and Brazil populations harbour different strains of Wolbachia symbionts, whereas the Ghana population harbours Cardinium symbionts. In response to antibiotic treatment females of each of the three populations became incompatible with untreated males of their own population, similar to that observed in crossings between females from one geographic population and males from another. Compatibility was restored in crosses involving uninfected Brazil females and uninfected Benin males, whereas the reciprocal crosses remained incompatible. Cardinium symbionts seem to be essential for oviposition in the Ghana population. It is concluded that their associated bacterial symbionts are the cause of the post-mating reproductive isolation previously observed among the three geographic populations. This insight is relevant to biological control of coconut mites for which N. paspalivorus is an effective predator, because introducing one geographic strain into the population of another (e.g. in field releases or mass cultures) may cause population growth depression.

Effects of kinship or familiarity? Small thrips larvae experience lower predation risk only in groups of mixed-size siblings.

In many species of insects, larvae are distributed in an aggregated fashion. As they may differ in size and size matters to predation risk, small larvae may be less likely to fall prey to predators when near large and therefore better-defended larvae. We hypothesize that the small larvae may profit even more when these large larvae are siblings. We tested this hypothesis on kinship-dependent survival in groups of larvae of the Western flower thrips (Frankliniella occidentalis) exposed to a predatory mite (Iphiseius degenerans). Our experiments showed that small larvae in sibling groups survive significantly better than in non-sibling groups, but only when such groups consisted of a mixture of small and large larvae. To test whether the survival effect we found is due to familiarity of thrips larvae growing up together (i.e. on one leaf), we also measured survival in sibling groups of larvae grown up on different leaves and in non-sibling groups of larvae grown up on the same leaf. These experiments showed an increased survival of small thrips larvae only in groups of sibling larvae from the same leaf. Non-sibling larvae did not show an increased survival when they come from the same leaf. Our results indicated that the increased survival in sibling groups was only partly due to the familiarity effect we tested. Growing up together did not return the same survival effect for non-siblings as it did for siblings. We conclude that growing up together is a necessary but not sufficient condition for discrimination in thrips larvae.

Neoseiulus paspalivorus, a predator from coconut, as a candidate for controlling dry bulb mites infesting stored tulip bulbs.

The dry bulb mite, Aceria tulipae, is the most important pest of stored tulip bulbs in The Netherlands. This tiny, eriophyoid mite hides in the narrow space between scales in the interior of the bulb. To achieve biological control of this hidden pest, candidate predators small enough to move in between the bulb scales are required. Earlier experiments have shown this potential for the phytoseiid mite, Neoseiulus cucumeris, but only after the bulbs were exposed to ethylene, a plant hormone that causes a slight increase in the distance between tulip bulb scales, just sufficient to allow this predator to reach the interior part of the bulb. Applying ethylene, however, is not an option in practice because it causes malformation of tulip flowers. In fact, to prevent this cosmetic damage, bulb growers ventilate rooms where tulip bulbs are stored, thereby removing ethylene produced by the bulbs (e.g. in response to mite or fungus infestation). Recently, studies on the role of predatory mites in controlling another eriophyoid mite on coconuts led to the discovery of an exceptionally small phytoseiid mite, Neoseiulus paspalivorus. This predator is able to move under the perianth of coconuts where coconut mites feed on meristematic tissue of the fruit. This discovery prompted us to test N. paspalivorus for its ability to control A. tulipae on tulip bulbs under storage conditions (ventilated rooms with bulbs in open boxes; 23 °C; storage period June-October). Using destructive sampling we monitored predator and prey populations in two series of replicated experiments, one at a high initial level of dry bulb mite infestation, late in the storage period, and another at a low initial dry bulb mite infestation, halfway the storage period. The first and the second series involved treatment with N. paspalivorus and a control experiment, but the second series had an additional treatment in which the predator N. cucumeris was released. Taking the two series of experiments together we found that N. paspalivorus controlled the populations of dry bulb mites both on the outer scale of the bulbs as well as in the interior part of the bulbs, whereas N. cucumeris significantly reduced the population of dry bulb mites on the outer scale, but not in the interior part of the bulb. Moreover, N. paspalivorus was found predominantly inside the bulb, whereas N. cucumeris was only found on the outer scale, thereby confirming our hypothesis that the small size of N. paspalivorus facilitates access to the interior of the bulbs. We argue that N. paspalivorus is a promising candidate for the biological control of dry bulb mites on tulip bulbs under storage conditions in the Netherlands.

Single versus multiple enemies and the impact on biological control of spider mites in cassava fields in West-Africa.

To determine whether to use single or multiple predator species for biological pest control requires manipulative field experiments. We performed such tests in Benin (West Africa) in cassava fields infested by the cassava green mite Mononychellus tanajoa, and the cotton red mite Oligonychus gossypii. These fields also harboured the cassava apex-inhabiting predator Typhlodromalus aripo and either the leaf-inhabiting predator Amblydromalus manihoti or Euseius fustis. We manipulated predator species composition on individual plants to determine their effect on prey and predator densities. In fields with T. aripo plus A. manihoti, M. tanajoa densities were reduced by T. aripo alone or together with A. manihoti, but neither of these predators, alone or together, reduced O. gossypii densities. In fields with T. aripo plus E. fustis, T. aripo alone or together with E. fustis exerted significant control over O. gossypii, but weak control over M. tanajoa. Densities of any of the predator species were not affected by co-occurring predator species, suggesting a minor role for intraguild predation in the field, contrary to earlier experiments on small plants in the laboratory. We conclude that (1) T. aripo is the most effective predator species in suppressing M. tanajoa, (2) two predator species, T. aripo and E. fustis, are needed to suppress O. gossypii, and (3) predator species together on the same plant do not negatively affect each other nor the extent to which they control their prey. We argue that intraguild predation is reduced due to partial niche separation among predator species.