Cory S. Straub

SPECIES IDENTITY DOMINATES THE RELATIONSHIP BETWEEN PREDATOR BIODIVERSITY AND HERBIVORE SUPPRESSION

Agricultural pest suppression is an important ecosystem service that may be threatened by the loss of predator diversity. This has stimulated interest in the relationship between predator biodiversity and biological control. Multiple-predator studies have shown that predators may complement or interfere with one another, but few experiments have determined if the resulting effects on prey are caused by changes in predator abundance, identity, species richness, or some combination of these factors. We experimentally isolated the effect of predator species richness on the biological control of an important agricultural pest, the green peach aphid. We found no evidence that increasing predator species richness affects aphid biological control; overall there was no strong complementarity or interference among predator species that altered the strength of aphid suppression. Instead, our experiments revealed strong effects of predator species identity, because predators varied dramatically in their per capita consumption rates. Our results are consistent with other multiple-predator studies finding strong species-identity effects and suggest that, for the biological control of aphids, conservation strategies that directly target key species will be more effective than those targeting predator biodiversity more broadly.

INCREASING ENEMY BIODIVERSITY STRENGTHENS HERBIVORE SUPPRESSION ON TWO PLANT SPECIES

Concern over biodiversity loss, especially at higher trophic levels, has led to a surge in studies investigating how changes in natural enemy diversity affect community and ecosystem functioning. These studies have found that increasing enemy diversity can strengthen, weaken, and not affect prey suppression, demonstrating that multi-enemy effects on prey are context-dependent. Here we ask how one factor, plant species identity, influences multi-enemy effects on prey. We focused on two plant species of agricultural importance, potato (Solanum tuberosum), and collards (Brassica oleracea L.). These species share a common herbivorous pest, the green peach aphid (Myzus persicae), but vary in structural and chemical traits that affect aphid reproductive rates and which may also influence inter-enemy interactions. In a large-scale field experiment, overall prey exploitation varied dramatically among the plant species, with enemies reducing aphid populations by approximately 94% on potatoes and approximately 62% on collards. Increasing enemy diversity similarly strengthened aphid suppression on both plants, however, and there was no evidence that plant species identity significantly altered the relationship between enemy diversity and prey suppression. Microcosm experiments suggested that, on both collards and potatoes, intraspecific competition among natural enemies exceeded interspecific competition. Enemy species showed consistent and significant differences in where they foraged on the plants, and enemies in the low-diversity treatment tended to spend less time foraging than enemies in the high-diversity treatment. These data suggest that increasing enemy diversity may strengthen aphid suppression because interspecific differences in where enemies forage on the plant allow for greater resource partitioning. Further, these functional benefits of diversity appear to be robust to changes in plant species identity.

Evidence for a Trade-Off between Host-Range Breadth and Host-Use Efficiency in Aphid Parasitoids

“The jack-of-all-trades is a master of none” describes the widely held belief that engaging in many tasks comes at the cost of being unable to do those tasks well. However, empirical evidence for generalist fitness costs remains scarce. We used published data from a long-term field survey of aphid parasitoids to determine whether relative specialists are more abundant than generalists on their shared hosts, a pattern that would be expected if generalists suffer a trade-off between host-range breadth and host-use efficiency. Relative specialists were more abundant than generalists on their shared hosts, but only when we used a measure of specialization that accounts for the taxonomic differences among parasitoids’ hosts. These results suggest that a generalist-specialist trade-off exists within this group of parasitoids and that the generalist fitness cost depends on the taxonomic breadth, rather than the number, of host species that are used.

Experimental evidence that the effectiveness of conservation biological control depends on landscape complexity

Summary 1. The expansion of intensive agricultural practices is a major threat to biodiversity and to the delivery of ecosystem services on which humans depend. Local-scale conservation management strategies, such as agri-environment schemes to preserve biodiversity, have been widely adopted to reduce the negative impacts of agricultural intensification. However, it is likely that the effectiveness of these local-scale management actions depend on the structure and composition of the surrounding landscape. 2. We experimentally tested the utility of floral resource strips to improve local-scale biological control of crop pests, when placed within a gradient of moderately simple through to highly complex landscapes. 3. We found that experimental provision of floral resources enhanced parasitism rates of two globally important crop pests in moderately simple landscapes but not in highly complex ones, and this translated into reduced pest abundances and increased crop yield. 4. Synthesis and applications. Our results lend experimental support for the ‘intermediate landscape complexity hypothesis’, which predicts that local conservation management will be most effective in moderately simple agricultural landscapes, and less effective in either very simple landscapes where there is no capacity for response, or in highly complex landscapes where response potential is already saturated. This knowledge will allow more targeted and cost-effective implementation of conservation biological control programs based on an improved understanding of landscape-dependent processes, which will reduce the negative impacts of agricultural intensification.

Experimental Approaches to Understanding the Relationship Between Predator Biodiversity and Biological Control

Conservation biological control (CBC) involves the manipulation of the environment to favour the natural enemies of pests. Alternative agricultural practices, such as organic farming, are more biodiversity-friendly than conventional agricultural practices and generally lead to greater predator species richness and abundance. This is desirable from a conservation perspective, but it is unclear how greater predator diversity affects biological control. Unfortunately, the predator ecology literature provides little guidance: increasing the number of predator species has been shown to enhance, diminish, and not affect prey suppression. In this chapter we explore how the experimental approach used in biodiversity and ecosystem functioning (BEF) studies, which focus on the ecological consequences of species loss, may be used to study how increasing predator diversity affects biological control. The notable features of this approach are: 1) realistic levels of species richness (i.e., < 2 predator species), 2) the use of substitutive, rather than additive, experimental designs, and 3) experimentally distinguishing the effect of species richness from the effects of species abundance, composition, and identity. This experimental approach can be used to identify which components of predator biodiversity—species richness, abundance, composition, and identity—should be targeted by CBC practitioners to maximize pest suppression. Further, it can be used to assess whether predator biodiversity conservation and biological control are truly compatible goals. Ultimately, we hope that this chapter will serve to motivate future research into this important problem.

Relationships between natural enemy diversity and biological control

Natural enemy diversity generally strengthens biological control, but individual studies have found everything from positive to negative effects. We discuss the factors that promote these different outcomes. We argue that a trait-based approach is helpful to improve our understanding of the relationship between enemy diversity and biological control, and suggest that enemy diversity is likely to be particularly important as an insurance against effects of climate change. Future research should increase the scale and ecological realism of enemy diversity studies, and consider both the strength and stability of biological control. Such research is likely to reveal even stronger evidence that conserving enemy biodiversity will improve biological pest control.

Multi–predator Effects Produced by Functionally Distinct Species Vary with Prey Density

Abstract Determining when multiple predator species provide better pest suppression than single species is a key step towards developing ecologically—informed biological control strategies. Theory and experiments predict that resource partitioning among functionally different predator species can strengthen prey suppression, because as a group they can access more prey types than functionally redundant predators. However, this prediction assumes that competition limits predation by functionally similar predators. Differences in prey density can alter the strength of competition, suggesting that prey abundance may modulate the effect of combining functionally diverse species. The experiment documented here examined the potential for functional differences among predator species to promote suppression of an insect pest, the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), at different prey densities. Predation was compared at two prey densities between microcosms that contained one predator species or two functionally distinct species: the lady beetle, Coleomegilla maculata De Geer (Coleoptera: Coccinellidae) that kills early L. decemlineata instars, and the soldier bug, Podisus maculiventris Say (Hemiptera: Pentatomidae) that kills late instars. The data show that combining these predators increased predation only when prey densities were low. This suggests that multiple predator species may only provide greater biological control than single species in systems where prey is limiting.

Additive effects of predator diversity on pest control caused by few interactions among predator species

1. Studies of the impact of predator diversity on biological pest control have shown idiosyncratic results. This is often assumed to be as a result of differences among systems in the importance of predator–predator interactions such as facilitation and intraguild predation. The frequency of such interactions may be altered by prey availability and structural complexity. A direct assessment of interactions among predators is needed for a better understanding of the mechanisms affecting prey abundance by complex predator communities.

Influence of nonhost plant diversity and natural enemies on the potato leafhopper, Empoasca fabae, and pea aphid, Acyrthosiphon pisum, in alfalfa

We examined the effects of nonhost plant diversity and predators on the potato leafhopper, Empoasca fabae (Harris), the pea aphid, Acyrthosiphon pisum (Harris), and their primary host plant, alfalfa, Medicago sativa L. Potato leafhopper intensity (i.e., leafhoppers/alfalfa stem density) and plant damage (i.e., hopperburn) were significantly greater in alfalfa mono- than in polycultures of alfalfa mixed with nonhost plant species. There was no significant effect of nonhost plant diversity on pea aphid intensity or on predator abundance. Predator:prey ratios were higher in poly- than in monoculture. One predator, Nabis sp., was selected for further study. A microcosm experiment indicated that Nabis is an effective predator of both herbivores, and suggested that nonhost plant diversity may enhance the predation of leafhoppers by Nabis. These results suggest that nonhost plant diversity and Nabis limit leafhopper populations and protect alfalfa from herbivory. The potential for nonhost plants to increase herbivore movement and vulnerability to predation is discussed.