Interactions among predators and phytophagous mites on apple; possible impact on biocontrol of Panonychus ulmi by Typhlodromus pyri in orchards

Laboratory experiments were conducted to determine the potential impact of the phytoseiid Euseius finlandicus, the mirid Blepharidopterus angulatus and the anthocorid Orius majusculus on the Typhlodromus pyri/Panonychus ulmi predator/prey relationship on apple. Euseius finlandicus consumed more immature spider mites than did T. pyri. When both phytoseiids were present and spider mite prey was abundant, there was no evidence of a negative interaction between the predators. In experiments where each predatory mite was confined with large numbers of the other predator, interspecific predation was exhibited by adults of each species on immatures of the other, but more so by E. finlandicus. In the predatory insect/phytoseiid experiments, when confined with spider mites and large numbers of T. pyri, both B. angulatus and O. majusculus consumed some T. pyri, but spider mites were the preferred prey. In experiments with B. angulatus, O. majusculus and T. pyri feeding on P. ulmi, there was no evidence of negative interactions between the predatory insects and T. pyri.     

Is the predatory mite <Emphasis Type="Italic">Kampimodromus aberrans</Emphasis> a candidate for the control of phytophagous mites in European apple orchards?

The most important biocontrol agents of phytophagous mites (mainly Tetranychidae) in European apple orchards are the predatory mites Amblyseius andersoni, Typhlodromus pyri and Euseius finlandicus (Phytoseiidae). A similar situation is found in Trentino (north-eastern Italy), an important apple production area in Europe. Another phytoseiid mite, Kampimodromus aberrans, can be dominant in neglected fruit orchards but is rare in commercial orchards because of its susceptibility to pesticides. However, pesticide resistant strains of K. aberrans have recently been found in vineyards. In the late 1990s, one of these resistant strains was successfully released on an experimental farm in Trentino. Kampimodromus aberrans spread to an apple orchard, despite the fact that it was colonized by T. pyri, A. andersoni and E. finlandicus, and became the dominant species. Since K. aberrans’ colonization appeared to be affected by apple cultivars, experiments were conducted on potted plants of three selected apple cultivars (Golden Delicious, Red Chief, Reinette du Canada). The results stressed the effect of cultivars on apple colonization by K. aberrans and suggested the role of leaf morphology in influencing this phenomenon. Field releases of K. aberrans were successfully performed in four commercial apple orchards. The incidence of K. aberrans in the total phytoseiid population increased over time and the predator became dominant in the season following its release in two orchards. Kampimodromus aberrans persists in these orchards as the dominant species. The adaptation of K. aberrans to varying environmental conditions, its tolerance to pesticides, and its competitiveness towards other phytoseiid species suggest a potential role of this species in the biological control of phytophagous mites in European apple orchards. Handling editor: Eric Lucas.     

The predatory mite Typhlodromalus aripo prefers green-mite induced plant odours from pubescent cassava varieties

It is well known that plant-inhabiting predators use herbivore-induced plant volatiles to locate herbivores being their prey. Much less known, however, is the phenomenon that genotypes of the same host plant species vary in the attractiveness of these induced chemical signals, whereas they also differ in characteristics that affect the predator’s foraging success, such as leaf pubescence. In a series of two-choice experiments (using a Y-tube olfactometer) we determined the preference of Typhlodromalus aripo for pubescent versus glabrous cassava cultivars infested with the cassava green mite Mononychellus tanajoa and also the preference for cultivars within each of the two groups. We found that when offered a choice between pubescent and glabrous cassava cultivars (either apex or leaves), T. aripo was significantly more attracted to pubescent cultivars. For each cultivar, M. tanajoa infested leaves and apices were equally attractive to T. aripo. There was however some variation in the response of T. aripo to M. tanajoa-infested plant parts within the group of pubescent cultivars, as well as within the group of glabrous cultivars. Our study confirms not only that T. aripo uses herbivore-induced plant volatiles to search for prey in cassava fields, but it also shows that it can discriminate between glabrous and pubescent cultivars and prefers the latter. This knowledge can be useful in selecting cultivars that are attractive and suitable to T. aripo, which, in turn, may promote biological control of the cassava green mite.     

Risk assessment of non-target effects caused by releasing two exotic phytoseiid mites in Japan: can an indigenous phytoseiid mite become IG prey?

Two exotic phytoseiid mites, Neoseiulus cucumeris and Amblyseius swirskii, are commercially available in Japan for the control of thrips and other pest insects. As part of a risk assessment of the non-target effects of releasing these two species, we investigated intraguild predation (IGP) between these exotic phytoseiid mites and an indigenous phytoseiid mite Gynaeseius liturivorus, which is promising as an indigenous natural enemy for the control of thrips in Japan. To understand IGP relations between the exotic and indigenous phytoseiid mites after use of the exotic mites for biological control, we investigated IGP between them in the absence of their shared prey. When an IG prey was offered to an IG predator, both exotic and indigenous females consumed the IG prey at all immature stages (egg, larva, protonymph, deutonymph), especially at its larval stages. The propensity for IGP in a no-choice test was measured by the survival time of IG prey corrected using the survival time of thrips offered to the IG predator. There was no significant difference in the propensity for IGP between N. cucumeris and G. liturivorus, but the propensity was significantly higher in A. swirskii than G. liturivorus. The propensity for IGP in a choice test was measured by the prey choice of the IG predator when a conspecific and a heterospecific larva were offered simultaneously as IG prey. Both exotic females consumed the heterospecific larva only. The indigenous female preferentially consumed the heterospecific larva when the heterospecific larva was N. cucumeris, but consumed the conspecific larva when the heterospecific larva was A. swirskii. We concluded that further investigation would be necessary for the exotic mites’ risk assessment, since the propensity for IGP of the two exotic females was similar to or higher than that of the indigenous female in both the no-choice and choice tests.     

Effects of predator-mediated apparent competition on the population dynamics of Tetranychus urticae on apples

The two spotted spider mite, Tetranychus urticae Koch, is an important herbivore pest of apple trees in Northwest China. This spider mite and another less damaging spider mite, Eotetranychus pruni Oudemans, are attacked by a common and often effective phytoseiid predator, Euseius finlandicus (Oudemans). Functional relationships were studied in the field to evaluate the impact of E. pruni and E. finlandicus on T. urticae. The results from this study showed that the predator-mediated apparent competition strongly affected the population dynamics of T. urticae. The addition of the apparent competitor E. pruni alone had little impact on T. urticae densities. Although the release of the predator E. finlandicus alone could result in reduction in T. urticae densities, the greatest reduction in T. urticae densities occurred in plots where both the predator E. finlandicus and apparent competitor E. pruni were released. In apple orchards, the early introduction of both the apparent competitor E. pruni and predator E. finlandicus would evidently increase the population size of the predator E. finlandicus and consequently significantly enhance the control of T. urticae populations. It is concluded from the study that the predator and apparent competitor release might be an appropriate control for the target species.     

Effects of foliar apple trichomes on Galendromus occidentalis (Nesbitt) (Acari: Phytoseiidae): preferences,fecundity and prey consumption

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Diet‐dependent intraguild predation between the predatory mites Neoseiulus californicus and Neoseiulus cucumeris

Based on the hypothesis that matching diets of intraguild (IG) predator and prey indicate strong food competition and thus intensify intraguild predation (IGP) as compared to non‐matching diets, we scrutinized diet‐dependent mutual IGP between the predatory mites Neoseiulus cucumeris and N. californicus. Both are natural enemies of herbivorous mites and insects and used in biological control of spider mites and thrips in various agricultural crops. Both are generalist predators that may also feed on plant‐derived substances such as pollen. Irrespective of diet (pollen or spider mites), N. cucumeris females had higher predation and oviposition rates and shorter attack latencies on IG prey than N. californicus. Predation rates on larvae were unaffected by diet but larvae from pollen‐fed mothers were a more profitable prey than those from spider‐mite fed mothers resulting in higher oviposition rates of IG predator females. Pollen‐fed protonymphs were earlier attacked by IG predator females than spider‐mite fed protonymphs. Spider mite‐fed N. californicus females attacked protonymphs earlier than did pollen‐fed N. californicus females. Overall, our study suggests that predator and prey diet may exert subtle influences on mutual IGP between bio‐control agents. Matching diets did not intensify IGP between N. californicus and N. cucumeris but predator and prey diets proximately influenced IGP through changes in behaviour and/or stoichiometry.     

Testing intraguild predation theory in a field system: does numerical dominance shift along a gradient of productivity?

Although ecological theory exists to predict dynamics in communities with intraguild predation (IGP), few empirical tests have examined this theory. IGP theory, in particular, predicts that when two competitors interact via IGP, with increasing resource productivity: (1) the IG predator will increase in abundance as the IG prey declines, and (2) increasing dominance of the IG predator will cause resource density to increase. Here, we provide a first test of these predictions in a field community consisting of a scale insect and its two specialist parasitoids, Aphytis melinus (the IG predator) and Encarsia perniciosi (the IG prey). The shared resource, California red scale, is a pest of citrus, and its productivity varies across a threefold range among citrus cultivars. We examined both absolute and relative densities of parasitoids along this natural gradient of scale productivity in three citrus cultivars (orange, grapefruit and lemon). Although both parasitoid species were found in all three cultivars, their abundances reflected those predicted by IGP theory: the IG prey species dominated at low productivity and the IG predator dominated at high productivity. This relationship was caused by an increase in Aphytis density with productivity. In addition, the density of scale increased with the dominance of the IG predator. These results from a field system demonstrate the important dynamic outcomes for food webs with IGP.     

Impact of intraguild predation on parasitoid foraging behaviour

1. Trophic interactions between predators and parasitoids can be described as intraguild predation (IGP) and are often asymmetric. Parasitoids (typically the IG prey) may respond to the threat of IGP by mitigating the predation risk for their offspring. 2. We used a system with a facultative predator Macrolophus caliginosus, the parasitoid Aphidius colemani, and their shared prey, the aphid Myzus persicae. We examined the functional responses of the parasitoid in the presence/absence of the predator on two host plants (aubergine and sweet pepper) with differing IGP risk. 3. Estimated model parameters such as parasitoid handling time increased on both plants where the predator was present, but impact of the predator varied with plant species. The predator, which could feed herbivorously on aubergine, had a reduced impact on parasitoid foraging on that plant. IG predator presence could reduce the searching effort of the IG prey depending on the plant, and on likely predation risk. 4. The results are discussed with regard to individual parasitoid's foraging behaviour and population stability; it is suggested that the presence of the predator can contribute to the stabilisation of host–parasitoid dynamics     

Intraguild predation among the predatory mites Amblyseius eharai,Amblyseius cucumeris and Amblyseius barkeri

Predator species with the same prey interact not only by competition for food and space but also by intraguild predation (IGP). The impact of IGP on introduced phytoseiid mites and native species in the context of biological control is a matter of considerable debate. Amblyseius eharai is the dominant native citrus species in central China, while Amblyseius cucumeris and Amblyseius barkeri are candidates for importation. All three species can feed on the spider mite Panonychus citri, which is the main pest in citrus. This study investigated, in the laboratory, possible IGP among these species in the absence and presence of P. citri, respectively. IGP in different densities of shared prey and intraguild (IG) prey was also studied. All three species consumed heterospecific larvae and eggs but not adults, and the IGP rate of larvae was significantly higher than that of eggs in the absence of shared prey. Additionally, the IGP rate of each group was reduced dramatically in the presence of both shared and IG prey when compared to the absence of shared prey. This occurs most likely because the three species prefer to feed on their natural prey P. citri, rather than on IG prey. Our results showed that A. eharai seems to be a more voracious IG predator than A. cucumeris. A. eharai was much more prone to IGP than A. barkeri.     

Intraguild predation of immature stages of British and Japanese coccinellids by the invasive ladybird Harmonia axyridis

Declines in native aphidophages in North America have been linked to intraguild predation (IGP) by the invasive coccinellid Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). It is feared that many British species will face a similar fate following the recent establishment of H. axyridis in the UK. Meanwhile, H. axyridis exists in apparent ecological equilibrium with other members of its guild in Japan. The impact of H. axyridis on British coccinellids is uncertain but intraguild predatory interactions do occur, particularly amongst immature stages. This study investigates IGP between immature stages of H. axyridis and various British and Japanese coccinellids. The only asymmetric IG predator of H. axyridis at first instar was Anatis ocellata (Linnaeus). Harmonia axyridis engaged in symmetric IGP with Coccinella septempunctata Linnaeus, Calvia quatuordecimguttata (Linnaeus), Harmonia quadripunctata (Pontoppidan) and Eocaria muiri Timberlake, but was the asymmetric IG predator of all other species studied. The level of IGP was high between fourth instar larvae, and frequently biased towards H. axyridis, except in the case of A. ocellata, which again was the only IG predator of H. axyridis. In interactions between fourth instar larvae and pre-pupae, IGP was unidirectional towards H. axyridis for all species except A. ocellata, which acted as both IG predator and IG prey. Pupae were better protected against IGP than pre-pupae but most species were still susceptible to attack by H. axyridis, although IGP was symmetric with A. ocellata, and H. quadripunctata pupae were never attacked. The differences in susceptibility of the various species and developmental stages to IGP by H. axyridis are discussed in relation to physical defence structures. We find no evidence that Japanese species have superior defences to British ones and suggest that behavioural strategies may enable co-existence in the native range. We discuss the relevance of IGP by H. axyridis to the species it is likely to encounter in Britain.     

Invasion success in communities with reciprocal intraguild predation depends on the stage structure of the resident population

The probability of individuals being targeted as prey often decreases as they grow in size. Such size‐dependent predation risk is very common in systems with intraguild predation (IGP), i.e. when predatory species interact through predation and competition. Theory on IGP predicts that community composition depends on productivity. When recently testing this prediction using a terrestrial experimental system consisting of two phytoseiid mite species, Iphiseius degenerans as the IG‐predator and Neoseiulus cucumeris as the IG‐prey, and pollen (Typha latifolia) as the shared resource, we could not find the predicted community shift. Instead, we observed that IG‐prey excluded IG‐predators when the initial IG‐prey/IG‐predator ratio was high, whereas the opposite held when the initial ratio was low, which is also not predicted by theory. We therefore hypothesized that the existence of vulnerable and invulnerable stages in the two populations could be an important driver of the community composition. To test this, we first demonstrate that IG‐prey adults indeed attacked IG‐predator juveniles in the presence of the shared resource. Second, we show that the invasion capacity of IG‐predators at high productivity levels indeed depended on the structure of resident IG‐prey populations. Third, we further confirmed our hypothesis by mimicking successive invasion events of IG‐predators into an established population of IG‐prey at high productivity levels, which consistently failed. Our results show that the interplay between stage structure of populations and reciprocal intraguild predation is decisive at determining the species composition of communities with intraguild predation.     

Phenotypic plasticity in anti-intraguild predator strategies: mite larvae adjust their behaviours according to vulnerability and predation risk

Interspecific threat-sensitivity allows prey to maximize the net benefit of antipredator strategies by adjusting the type and intensity of their response to the level of predation risk. This is well documented for classical prey-predator interactions but less so for intraguild predation (IGP). We examined threat-sensitivity in antipredator behaviour of larvae in a predatory mite guild sharing spider mites as prey. The guild consisted of the highly vulnerable intraguild (IG) prey and weak IG predator Phytoseiulus persimilis, the moderately vulnerable IG prey and moderate IG predator Neoseiulus californicus and the little vulnerable IG prey and strong IG predator Amblyseius andersoni. We videotaped the behaviour of the IG prey larvae of the three species in presence of either a low- or a high-risk IG predator female or predator absence and analysed time, distance, path shape and interaction parameters of predators and prey. The least vulnerable IG prey A. andersoni was insensitive to differing IGP risks but the moderately vulnerable IG prey N. californicus and the highly vulnerable IG prey P. persimilis responded in a threat-sensitive manner. Predator presence triggered threat-sensitive behavioural changes in one out of ten measured traits in N. californicus larvae but in four traits in P. persimilis larvae. Low-risk IG predator presence induced a typical escape response in P. persimilis larvae, whereas they reduced their activity in the high-risk IG predator presence. We argue that interspecific threat-sensitivity may promote co-existence of IG predators and IG prey and should be common in predator guilds with long co-evolutionary history.     

Functional response of Euseius finlandicus and Amblyseius andersoni to Panonychus ulmi on apple and peach leaves in the laboratory

The functional response of adult females of the predatory mites Euseius (Amblyseius) finlandicus and Amblyseius andersoni to larvae and adult females of the fruit tree red spider mite Panonychus ulmi was determined on apple and peach leaf disks in the laboratory at 25°C and 16:8 (L:D). For adult females of P. ulmi the predation efficiency of E. finlandicus was higher on peach than on apple, whereas that of A. andersoni was higher on apple than on peach. Efficiency of predation on larvae of P. ulmi by either predator did not differ significantly between apple and peach. On both plants, A. andersoni had a higher predation rate than E. finlandicus on larvae of P. ulmi. It is concluded that in the laboratory the host plant has a substantial effect on predation efficiency of A. andersoni and E. finlandicus when they preyed on adults but not when they preyed on larvae of P. ulmi.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.     

Trichomes and spider-mite webbing protect predatory mite eggs from intraguild predation

Predaceous arthropods are frequently more abundant on plants with leaves that are pubescent or bear domatia than on plants with glabrous leaves. We explored the hypothesis that for some predatory mites this is because pubescence affords protection from intraguild predation. In laboratory experiments, we tested whether apple leaf pubescence protected Typhlodromus pyri eggs from predation by western flower thrips, Frankliniella occidentalis. To investigate the effect of pubescence further, we added cotton fibers to trichome-free leaves. We also determined whether webbing produced by Tetranychus urticae protected Phytoseiulus persimilis eggs from predation by F. occidentalis. Predation by thrips on T. pyri eggs oviposited on field-collected pubescent "Erwin Bauer" apple leaves was significantly less than on glabrous "Crittenden" apple leaves. Phytoseiid eggs oviposited in the cotton fibers were preyed upon significantly less than those on the trichome-free bean disk. Increasing the cotton fiber density from 5 to 20 fibers only slightly further reduced predation by thrips on T. pyri eggs. Thrips fed upon significantly fewer P. persimilis eggs oviposited in Te. urticae webbing than eggs oviposited on a surface that differed only in the absence of Te. urticae web. We conclude that a complex leaf topography reduces intensity of intraguild predation in this system.     

Morphological and physiological correlates of niche breadth in two species of Espeletia (Compositae) in the Venezuelan Andes

Summary We compared adaptive strategies in two plants of Venezuelan páramos (alpine areas): the widely distributed, caulescent, and pubescent Espeletia schultzii Wedd. with the acaulescent, nearly glabrous E. atropurpurea A.C. Smith which is restricted to mesic sites just above treeline. Both species occur together at 3,450 m, near treeline.Physiologically, E. schultzii was more drought resistant than E. atropurpurea, and was better adapted for carbon dioxide fixation under low temperatures. The densely pubescent leaves of E. schultzii are highly reflective; this increases the intensity of light needed for photosynthetic saturation and influences leaf temperature. Leaf pubescence may reduce the level of insect predation.Measurements of leaf productivity indicate higher values for E. atropurpurea during the rainy season and higher values for E. schultzii during the dry season. However, annual values of leaf productivity are similar for both species.Benefits of specialization in E. atropurpurea include reduced costs for stem and leaf hair production, higher growth rates during the rainy season and the ability to grow beneath canopies of some larger arborescent species. Costs of specialization include lower growth rates during the dry season, great susceptibility to insect predation and restriction to low elevation, mesic sites.     

Dynamics of a intraguild predation model with generalist or specialist predator

Intraguild predation (IGP) is a combination of competition and predation which is the most basic system in food webs that contains three species where two species that are involved in a predator/prey relationship are also competing for a shared resource or prey. We formulate two intraguild predation (IGP: resource, IG prey and IG predator) models: one has generalist predator while the other one has specialist predator. Both models have Holling-Type I functional response between resource-IG prey and resource-IG predator; Holling-Type III functional response between IG prey and IG predator. We provide sufficient conditions of the persistence and extinction of all possible scenarios for these two models, which give us a complete picture on their global dynamics. In addition, we show that both IGP models can have multiple interior equilibria under certain parameters range. These analytical results indicate that IGP model with generalist predator has “top down” regulation by comparing to IGP model with specialist predator. Our analysis and numerical simulations suggest that: (1) Both IGP models can have multiple attractors with complicated dynamical patterns; (2) Only IGP model with specialist predator can have both boundary attractor and interior attractor, i.e., whether the system has the extinction of one species or the coexistence of three species depending on initial conditions; (3) IGP model with generalist predator is prone to have coexistence of three species.     

The geographic distribution of pubescence in the sea daisy, Borrichia aborescens, on Bahamian Islands

Borrichia aborescens (L.) DC. is a salt‐tolerant perennial herb that is common on small islands in the central Bahamas. Two morphotypes are present: one with densely pubescent leaves and one with glabrous leaves. I conducted surveys in three archipelagos to document the geographical distribution of pubescence in this species and to infer the underlying mechanisms. B. aborescens was also grown from seed in a greenhouse. The pubescent form of B. aborescens was relatively more abundant on small islands than on nearby large ‘mainland’ islands. In two of the three archipelagoes, pubescence increased with distance on small islands. The pubescent form was relatively more abundant on small islands exposed to the open ocean compared to small islands that were protected by mainland islands or reefs on all sides. On a large mainland island, the pubescent form decreased in relative abundance inland from the coast. B. aborescens cultivated in a greenhouse revealed the effect of a genetic factor on the expression of pubescence. The observed patterns of variation are consistent with a physiological explanation for the adaptive benefit of pubescence. Trichomes may prevent physical blockage of the stomata by accumulation of salt in areas near breaking waves and salt spray.     

Effect of pollen supplement on intraguild predatory interactions between two omnivores: The importance of spatial dynamics

Arthropods often engage in complex trophic interactions such as intraguild predation (IGP), true omnivory (i.e., feeding on plants and prey), and apparent competition. Theoretical treatments of the effects of such interactions on herbivore populations have been concerned almost entirely with equilibrium conditions. Yet these interactions are common in non-equilibrium settings such as agroecosystems, where they are likely to have a strong influence on pest populations. We therefore tested short-term effects of IGP and food supplementation on interactions between two predators (the phytoseiid mite Neoseiulus cucumeris and the anthocorid bug Orius laevigatus) and their shared prey, Frankliniella occidentalis, on strawberry plants. All three consumers feed on strawberry pollen, both mites and bugs prey on thrips, and the bug also feeds on the mites (IGP). Strong IGP on mites (IG prey) by the bugs (IG predator) was recorded in structurally-simple arenas. In a more complex setting (whole-plants), however, the intensity of IGP differed among plant structures. Likewise, pollen supplementation reduced both IGP and predation on thrips in a structurally simple setting. In the whole-plant experiment, IGP was more intense on pollen-bearing than pollen-free flowers. The study illustrated how spatial dynamics, generated when consumers track food sources differently in the habitat and possibly when herbivorous and IG prey alter their distribution to escape predation, led to site-specific configuration of interacting populations. The intensity of resulting trophic interactions was weakened by food supplementation and by increased complexity of the habitat.