Assembly rules for functional groups of North American shrews: effects of geographic range and habitat partitioning

Trophic cascades exist in numerous terrestrial systems, including many systems with ants as the top predator. Many studies show how behavioral modifications of herbivores are especially important in mediating species interactions and trophic cascades. Although most studies of trophic cascades focus on predator-herbivore-plant links, the trophic cascades concept could be applied to almost any level of trophic interactions. Especially considering the importance of parasites we consider here the interactions between the parasitic phorid fly, Pseudacteon sp. (Diptera: Phoridae), its ant host, Azteca instabilis (F. Smith) (Hymenoptera: Formicidae), and the herbivore, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) in the coffee agroecosystem. We investigated the effects of phorid flies on ant behavior by monitoring ant recruitment to tuna baits over a 30-min period in the presence or absence of phorid flies. To study the indirect effects of phorids on larvae, we placed baits on coffee plants to elevate ant foraging levels to levels near to ant nests, placed larvae near baits, and recorded the effects of ants on larvae either in the presence or absence of phorid flies. We found that phorid fly presence significantly reduced ant ability to recruit to baits through behavioral modifications and also significantly lessened ant ability to attack, carry away, or force herbivores off plants. We conclude there is a behaviorally-modified species-level trophic cascade in the coffee agroecosystem, with potentially important effects in ant and herbivore communities as well as for coffee production.     

Do Herbivores Eavesdrop on Ant Chemical Communication to Avoid Predation?

Strong effects of predator chemical cues on prey are common in aquatic and marine ecosystems, but are thought to be rare in terrestrial systems and specifically for arthropods. For ants, herbivores are hypothesized to eavesdrop on ant chemical communication and thereby avoid predation or confrontation. Here I tested the effect of ant chemical cues on herbivore choice and herbivory. Using Margaridisa sp. flea beetles and leaves from the host tree (Conostegia xalapensis), I performed paired-leaf choice feeding experiments. Coating leaves with crushed ant liquids (Azteca instabilis), exposing leaves to ant patrolling prior to choice tests (A. instabilis and Camponotus textor) and comparing leaves from trees with and without A. instabilis nests resulted in more herbivores and herbivory on control (no ant-treatment) relative to ant-treatment leaves. In contrast to A. instabilis and C. textor, leaves previously patrolled by Solenopsis geminata had no difference in beetle number and damage compared to control leaves. Altering the time A. instabilis patrolled treatment leaves prior to choice tests (0-, 5-, 30-, 90-, 180-min.) revealed treatment effects were only statistically significant after 90- and 180-min. of prior leaf exposure. This study suggests, for two ecologically important and taxonomically diverse genera (Azteca and Camponotus), ant chemical cues have important effects on herbivores and that these effects may be widespread across the ant family. It suggests that the effect of chemical cues on herbivores may only appear after substantial previous ant activity has occurred on plant tissues. Furthermore, it supports the hypothesis that herbivores use ant chemical communication to avoid predation or confrontation with ants.     

Cheating on a mutualism: indirect benefits of ant attendance to a coccidophagous coccinellid

Coccinellids (Coleoptera: Coccinellidae) are generally unable to prey on ant-tended prey. However, particular coccinellid species have morphological, behavioral, or chemical characteristics that render them immune to ant attacks, and some species are even restricted to ant-tending areas. The benefit gained from living in close association with ants can be twofold: (1) gaining access to high-density prey areas and (2) gaining enemy-free space. Here, the myrmecophily of Azya orbigera Mulsant (Coleoptera: Coccinellidae), an important predator of the green coffee scale, Coccus viridis (Green) (Hemiptera: Coccidae), is reported. In this paper, three main questions were studied. (1) Are the waxy filaments of A. orbigera larvae effective as defense against attacks of the mutualistic ant partner of C. viridis, Azteca instabilis F. Smith (Hymenoptera: Formicidae)? (2) Does A. instabilis reduce the rate at which A. orbigera larvae prey on scales? (3) Do A. orbigera larvae gain enemy-free space by living in close association with A. instabilis? Laboratory and field experiments were conducted to answer these questions. We found that, because of the sticky waxy filaments of A. orbigera larvae, A. instabilis is incapable of effectively attacking them and, therefore, the predation rate of A. orbigera on C. viridis does not decrease in the presence of ants. Furthermore, A. instabilis showed aggressive behavior toward A. orbigera's parasitoids, and the presence of ants reduced the parasitism suffered by A. orbigera. This is the first time that this kind of indirect positive effect is reported for an ant and a coccidophagous coccinellid. Furthermore, this indirect positive effect may be key to the persistence of A. orbigera's populations.     

Cascading indirect effects in a coffee agroecosystem: effects of parasitic phorid flies on ants and the coffee berry borer in a high-shade and low-shade habitat

Nonconsumptive effects (NCE) of parasites on hosts vary with habitat complexity thereby modifying trait-mediated effects on lower trophic levels. In coffee agroecosystems, Pseudacteon sp. phorid fly parasites negatively affect Azteca instabilis F. Smith ants via NCE thereby indirectly benefiting prey. It is unknown how differences in habitat complexity influence Azteca-phorid interactions or how phorids affect the coffee berry borer (Hypothenemus hampei Ferrari), an important pest of coffee (Coffea arabica L). We tested the following hypotheses in field and lab experiments to find the impact of NCE of phorids on A. instabilis and trait-mediated indirect effects of phorids on the coffee berry borer: (1) Phorid effects on A. instabilis differ between complex and simple shade habitats and (2) Phorids, by modifying A. instabilis behavior, indirectly affect coffee berry borer abilities to invade coffee berries. Phorids had greater impacts on A. instabilis activity in low-shade farms, but differences in phorid impacts were not mediated by phorid density or light availability. In the lab, phorids had strong cascading effects on abilities of A. instabilis to deter coffee berry borers. Without phorids, A. instabilis limited coffee berry borer attacks, whereas when the coffee berry borer was alone or with A. instabilis and phorids, more coffee fruits were attacked by coffee berry borer. These results indicate that A. instabilis has stronger biological control potential in high-shade farms, but the exact mechanism deserves further attention.     

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

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

Effects of predatory ants on lower trophic levels across a gradient of coffee management complexity

1. Ants are important predators in agricultural systems, and have complex and often strong effects on lower trophic levels. Agricultural intensification reduces habitat complexity, food web diversity and structure, and affects predator communities. Theory predicts that strong top-down cascades are less likely to occur as habitat and food web complexity decrease. 2. To examine relationships between habitat complexity and predator effects, we excluded ants from coffee plants in coffee agroecosystems varying in vegetation complexity. Specifically, we studied the effects of eliminating ants on arthropod assemblages, herbivory, damage by the coffee berry borer and coffee yields in four sites differing in management intensification. We also sampled ant assemblages in each management type to see whether changes in ant assemblages relate to any observed changes in top-down effects. 3. Removing ants did not change total arthropod densities, herbivory, coffee berry borer damage or coffee yields. Ants did affect densities of some arthropod orders, but did not affect densities of different feeding groups. The effects of ants on lower trophic levels did not change with coffee management intensity. 4. Diversity and activity of ants on experimental plants did not change with coffee intensification, but the ant species composition differed. 5. Although variation in habitat complexity may affect trophic cascades, manipulating predatory ants across a range of coffee agroecosystems varying in management intensity did not result in differing effects on arthropod assemblages, herbivory, coffee berry borer attack or coffee yields. Thus, there is no clear pattern that top-down effects of ants in coffee agroecosystems intensify or dampen with decreased habitat complexity.     

Stage‐dependent responses to emergent habitat heterogeneity: consequences for a predatory insect population in a coffee agroecosystem

Interactions among members of biological communities can create spatial patterns that effectively generate habitat heterogeneity for other members in the community, and this heterogeneity might be crucial for their persistence. For example, stage‐dependent vulnerability of a predatory lady beetle to aggression of the ant, Azteca instabilis, creates two habitat types that are utilized differently by the immature and adult life stages of the beetle. Due to a mutualistic association between A. instabilis and the hemipteran Coccus viridis – which is A. orbigera main prey in the area – only plants around ant nests have high C. viridis populations. Here, we report on a series of surveys at three different scales aimed at detecting how the presence and clustered distribution of ant nests affect the distribution of the different life stages of this predatory lady beetle in a coffee farm in Chiapas, Mexico. Both beetle adults and larvae were more abundant in areas with ant nests, but adults were restricted to the peripheries of highest ant activity and outside the reach of coffee bushes containing the highest densities of lady beetle larvae. The abundance of adult beetles located around trees with ants increased with the size of the ant nest clusters but the relationship is not significant for larvae. Thus, we suggest that A. orbigera undergoes an ontogenetic niche shift, not through shifting prey species, but through stage‐specific vulnerability differences against a competitor that renders areas of abundant prey populations inaccessible for adults but not for larvae. Together with evidence presented elsewhere, this study shows how an important predator is not only dependent on the existence of two qualitatively distinct habitat types, but also on the spatial distribution of these habitats. We suggest that this dependency arises due to the different responses that the predator's life stages have to this emergent spatial pattern.     

Arboreal Ant Abundance and Leaf Miner Damage in Coffee Agroecosystems in Mexico

Shaded coffee agroecosystems traditionally have few pest problems potentially due to higher abundance and diversity of predators of herbivores. However, with coffee intensification (e.g., shade tree removal or pruning), some pest problems increase. For example, coffee leaf miner outbreaks have been linked to more intensive management and increased use of agrochemicals. Parasitic wasps control the coffee leaf miner, but few studies have examined the role of predators, such as ants, that are abundant and diverse in coffee plantations. Here, we examine linkages between arboreal ant communities and coffee leaf miner incidence in a coffee plantation in Mexico. We examined relationships between incidence and severity of leaf miner attack and: (1) variation in canopy cover, tree density, tree diversity, and relative abundance of Inga spp. shade trees; (2) presence of Azteca instabilis, an arboreal canopy dominant ant; and (3) the number of arboreal twig‐nesting ant species and nests in coffee plants. Differences in vegetation characteristics in study plots did not correlate with leaf miner damage perhaps because environmental factors act on pest populations at a larger spatial scale. Further, presence of A. instabilis did not influence presence or severity of leaf miner damage. The proportion of leaves with leaf miner damage was significantly lower where abundance of twig‐nesting ants was higher but not where twig‐nesting ant richness was higher. These results indicate that abundance of twig‐nesting ants in shaded coffee plantations may contribute to maintenance of low leaf miner populations and that ants provide important ecosystem services in coffee agroecosystems.     

Herbivore and predator diversity interactively affect ecosystem properties in an experimental marine community

Interacting changes in predator and prey diversity likely influence ecosystem properties but have rarely been experimentally tested. We manipulated the species richness of herbivores and predators in an experimental benthic marine community and measured their effects on predator, herbivore and primary producer performance. Predator composition and richness strongly affected several community and population responses, mostly via sampling effects. However, some predators survived better in polycultures than in monocultures, suggesting complementarity due to stronger intra- than interspecific interactions. Predator effects also differed between additive and substitutive designs, emphasizing that the relationship between diversity and abundance in an assemblage can strongly influence whether and how diversity effects are realized. Changing herbivore richness and predator richness interacted to influence both total herbivore abundance and predatory crab growth, but these interactive diversity effects were weak. Overall, the presence and richness of predators dominated biotic effects on community and ecosystem properties.     

Prey‐specific foraging tactics in a web‐building spider

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Agroforestry management affects coffee pests contingent on season and developmental stage

1 Management of vegetational diversity in agroecosystems is a potentially regulating factor of pest population dynamics and may affect developmental stages in different ways.
2 We investigated the population dynamics of red spider mites, coffee leaf miners, and coffee berry borers in three management types of coffee agroforests: increasing plant diversity from a few shade tree species (simple-shade agroforests), intermediate-shade tree species (complex-shade agroforests) to high-shade tree species (abandoned coffee agroforests) in Ecuador. Furthermore, we studied how changes in agroforestry management affect population stage structure of each coffee pest.
3 Our results show that agroforestry management affected seasonal patterns of coffee pests in that higher densities of red spider mites were observed from August to December, coffee leaf miners from December to February, and coffee berry borers from May to July. Moreover, specific developmental stages of red spider mites, coffee leaf miners, and coffee berry borers differed in their responses to agroforestry management. During all stages, red spider mite reached higher densities in simple-shade agroforests compared with complex-shade and abandoned agroforests. Meanwhile, coffee leaf miner densities decreased from simple-shade to complex-shade and abandoned agroforests, but only for larvae, not pupae. Similarly, only coffee berry borer adults (but not eggs, larvae and pupae) demonstrated a response to agroforestry management. Environmental variables characterizing each agroforestry type proved to be important drivers of pest population densities in the field.
4 We emphasize the importance of considering seasonal differences and population structure while investigating arthropod responses to different habitat types because responses change with time and developmental stages.     

A Keystone Ant Species Provides Robust Biological Control of the Coffee Berry Borer Under Varying Pest Densities

Species’ functional traits are an important part of the ecological complexity that determines the provisioning of ecosystem services. In biological pest control, predator response to pest density variation is a dynamic trait that impacts the provision of this service in agroecosystems. When pest populations fluctuate, farmers relying on biocontrol services need to know how natural enemies respond to these changes. Here we test the effect of variation in coffee berry borer (CBB) density on the biocontrol efficiency of a keystone ant species (Azteca sericeasur) in a coffee agroecosystem. We performed exclosure experiments to measure the infestation rate of CBB released on coffee branches in the presence and absence of ants at four different CBB density levels. We measured infestation rate as the number of CBB bored into fruits after 24 hours, quantified biocontrol efficiency (BCE) as the proportion of infesting CBB removed by ants, and estimated functional response from ant attack rates, measured as the difference in CBB infestation between branches. Infestation rates of CBB on branches with ants were significantly lower (71%-82%) than on those without ants across all density levels. Additionally, biocontrol efficiency was generally high and did not significantly vary across pest density treatments. Furthermore, ant attack rates increased linearly with increasing CBB density, suggesting a Type I functional response. These results demonstrate that ants can provide robust biological control of CBB, despite variation in pest density, and that the response of predators to pest density variation is an important factor in the provision of biocontrol services. Considering how natural enemies respond to changes in pest densities will allow for more accurate biocontrol predictions and better-informed management of this ecosystem service in agroecosystems.     

Weathering the storm: how lodgepole pine trees survive mountain pine beetle outbreaks

Climate change leads to phenology shifts of many species. However, not all species shift in parallel, which can desynchronize interspecific interactions. Within trophic cascades, herbivores can be top–down controlled by predators or bottom–up controlled by host plant quality and host symbionts, such as plant-associated micro-organisms. Synchronization of trophic levels is required to prevent insect herbivore (pest) outbreaks. In a common garden experiment, we simulated an earlier arrival time (~2 weeks) of the aphid Rhopalosiphum padi on its host grass Lolium perenne by enhancing the aphid abundance during the colonization period. L. perenne was either uninfected or infected with the endophytic fungus Epichloë festucae var. lolii. The plant symbiotic fungus produces insect deterring alkaloids within the host grass. Throughout the season, we tested the effects of enhanced aphid abundance in spring on aphid predators (top–down) and grass–endophyte (bottom–up) responses. Higher aphid population sizes earlier in the season lead to overall higher aphid abundances, as predator occurrence was independent of aphid abundances on the pots. Nonetheless, after predator occurrence, aphids were controlled within 2 weeks on all pots. Possible bottom–up control of aphids by increased endophyte concentrations occurred time delayed after high herbivore abundances. Endophyte-derived alkaloid concentrations were not significantly affected by enhanced aphid abundance but increased throughout the season. We conclude that phenology shifts in an herbivorous species can desynchronize predator–prey and plant–microorganism interactions and might enhance the probability of pest outbreaks with climate change.     

Ants defend coffee from berry borer colonization

Ants frequently prevent herbivores from damaging plants. In agroecosystems they may provide pest control services, although their contributions are not always appreciated. Here we compared the ability of eight ant species to prevent the coffee berry borer from colonizing coffee berries with a field exclusion experiment. We removed ants from one branch (exclusion) and left ants to forage on a second branch (control) before releasing 20 berry borers on each branch. After 24 h, six of eight species had significantly reduced the number of berries bored by the berry borer compared to exclusion treatment branches. While the number of berries per branch was a significant covariate explaining the number of berries bored, ant activity (that varied greatly among species) was not a significant factor in models. This study is the first field experiment to provide evidence that a diverse group of ant species limit the berry borer from colonizing coffee berries.     

Avian Foraging Behavior in Two Different Types of Coffee Agroecosystem in Chiapas, Mexico

This study describes the foraging ecology of birds during summer and winter in two different types of coffee agroecosystems in Chiapas, Mexico. Avian foraging behavior is documented in two agroecosystems of differing management intensity, structurally similar but with different levels of floristic diversity, during summer and winter seasons. The distribution of tree species used by birds was more even, and birds used a greater diversity of tree species, in the more diverse coffee shade system. Much of the variation in resource use derived from shifts in the use of flowers and fruit, highlighting the importance in resource phenology for birds. Insectivory was more frequent in winter than summer for the coffee layer, and in summer for the shade layer. Given the vegetative structural similarity of the two coffee agroecosystems included in this study, floristic differences probably accounted for much of the difference in the bird communities between the management systems, especially given the strong seasonal response to flowering and fruiting. This work suggests that plentiful and diverse food resources associated with the high diversity of plant species may facilitate coexistence of the high number of bird species found in shade-grown coffee agroecosystems.     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.     

Effect of plants on the searching efficiency of a generalist predator: the importance of predator-prey spatial association

In most studies of tritrophic interactions, the effect of plants on predators is confounded with changes in prey and predator behaviors after an encounter event. Here, we estimate how the effect of plants on prey distribution (in the absence of the predator) and on predator foraging behavior (in the absence of prey) may influence predation rate of Orius insidiosus (Say) (Heteroptera: Anthocoridae) in 11 plant by prey species combinations. The within-leaf distributions of O. insidiosus and its prey overlapped most on bean plants. The predator's foraging behavior (e.g., walking speed, turning rate) also differed among plant species. Simulations, using the prey distribution data and predator's foraging patterns on leaf surfaces of each plant species, show that, overall, the searching efficiency of O. insidiosus was higher on leaves of bean and corn than of tomato. However, the predator's searching efficiency was not consistent within plant species. Thus, the combined effect of plants directly on the predator and indirectly through the prey influenced the predator's searching efficiency.     

A Relationship Between Avian Foraging Behavior and Infestation by Trombiculid Larvae (Acari) in Chiapas, Mexico

Birds face varying risk from parasites as they select and utilize habitat. Unfortunately, behavioral and habitat correlates of parasitism in birds are not well documented. This study combines data from a foraging behavior study with results from a banding study to test whether behavior and habitat affect an ectoparasite infestation by trombiculid mite (chigger) larvae on the bird community found in two different coffee agroecosystems in Chiapas, Mexico. Individuals from bird species with regular prevalence ( i.e. , infestation) foraged more frequently in lower vegetative layers and had significantly lower foraging height than those from species with little or no prevalence, suggesting that foraging near the ground increases exposure risk to chigger larvae. Using linear regression, across species, parasite prevalence decreased with increasing average foraging height. Lower infestation rates were found in coffee agroecosystems with higher management intensity ( i.e. , less shade and drier conditions), suggesting that management activities influence infestation rates. Consequently, drier tropical habitats may pose less risk to birds from ectoparasites, though seasonal prevalence was highest during the winter dry season. Although no direct link was found between host condition and infestation by chigger larvae on the wintering grounds, birds were sampled during the middle of the over-wintering period, not the end when infestation could affect birds fattening for migration.     

Strong and weak trophic cascades along a productivity gradient

Variation in the strengths of predator effects, although commonly observed in natural communities, and predicted from theoretical models, remains poorly understood in the study of food web interactions and community structure. In this study, I first showed that prey species in benthic pond food webs were highly variable in their susceptibility to predators. Some were vulnerable throughout their lives, whereas others were vulnerable as juveniles, but able to grow to a size-refuge. Next, I showed that herbivore and predator abundance increased along a natural productivity gradient among 29 ponds, and herbivore species composition shifted from dominance by vulnerable to dominance by invulnerable herbivore species along this gradient. However, there was a considerable amount of variation both in herbivore biomass and composition at intermediate productivity; some were dominated by small species and others by larger species. Finally, in in situ exclosure experiments, I found that predator effects were strong and cascaded to plants in a low productivity pond and in an intermediate productivity pond dominated by small herbivore species. Alternatively, in a high productivity pond and in an intermediate productivity pond dominated by larger herbivores, I found that predator effects on prey biomass were weak, and did not cascade to plants.