Predator Species Richness Research Articles

Container-breeding mosquitoes and predator community dynamics along an urban-forest gradient: The effects of habitat type and isolation

Environmental disturbances such as deforestation, urbanization or pollution have been widely acknowledged to play a key role in the emergence of many infectious diseases, including mosquito-borne viruses. However, we have little understanding of how habitat isolation affects the communities containing disease vectors. Here, we test the effects of habitat type and isolation on the colonization rates, species richness and abundances of mosquitoes and their aquatic predators in water-filled containers in northwestern Thailand. For eight weeks water-filled containers were monitored in areas containing forest, urban and agricultural habitats and mixtures of these three. Mosquito larvae of the genera Aedes and Culex appeared to be differentially affected by the presence of the dominant predator; Toxorhynchites splendens (Culicidae). Therefore, a predation experiment was conducted to determine predator response to prey density and its relative effects on different mosquito prey populations. Colonization rates, species richness and abundances of mosquito predators were strongly related to forest habitat and to the distance from other aquatic habitats. Areas with more tree cover had higher predator species richness and abundance in containers. Containers that were close to surface water were more rapidly colonized than those further away. In all habitat types, including urban areas, when predators were present, the number of mosquito larvae was much lower. Containers in urban areas closer to water-bodies, or with more canopy cover, had higher predator colonization rates and species richness. T. splendens (Culicidae) preyed on the larvae of two mosquito genera at different rates, which appeared to be related to prey behaviour. This study shows that anthropogenic landscape modification has an important effect on the natural biological control of mosquitoes. Vector control programmes and urban planning should attempt to integrate ecological theory when developing strategies to reduce mosquito populations. This would result in management strategies that are beneficial for both public health and biodiversity.

Contribution of predator identity to the suppression of herbivores by a diverse predator assemblage.

Studies manipulating predator diversity and measuring the impact on herbivore abundance have found that enhancing predator species richness often increases the strength of prey suppression. This relationship may be due to mechanisms such as complementarity or facilitation, which are considered "true" benefits of diversity because greater prey suppression is an emergent property of the multispecies predator community. Or it may be due to an identity effect, an "apparent" benefit of diversity that results from the greater likelihood of including one particularly voracious predator species as the total number of predator species increases. In separate greenhouse and field experiments, we simultaneously manipulated the species richness and species composition of predators attacking bird cherry-oat aphids (Rhopalosiphum padi) (L.) on wheat (Triticum aestivum L.). We found that on average aphid suppression by species-rich predator assemblages was greater than suppression by single-species monocultures. However, the performance of individual predator species varied and the species-rich assemblages did not outperform all single-species compositions, suggesting an identity effect. In particular, single-species compositions of the lady beetle Coleomegilla maculata (DeGeer) exhibited high performance across experiments, and on average predator assemblages that contained a lady beetle predator had lower overall aphid abundance than compositions where lady beetles were absent. Taken together, these results provide evidence for the dominant role of lady beetles, especially C. maculata, in natural pest suppression and suggest that predator species composition and identity are important factors to consider in efforts to conserve this valuable ecosystem service.

Tree diversity promotes functional dissimilarity and maintains functional richness despite species loss in predator assemblages

The effects of species loss on ecosystems depend on the community's functional diversity (FD). However, how FD responds to environmental changes is poorly understood. This applies particularly to higher trophic levels, which regulate many ecosystem processes and are strongly affected by human-induced environmental changes. We analyzed how functional richness (FRic), evenness (FEve), and divergence (FDiv) of important generalist predators-epigeic spiders-are affected by changes in woody plant species richness, plant phylogenetic diversity, and stand age in highly diverse subtropical forests in China. FEve and FDiv of spiders increased with plant richness and stand age. FRic remained on a constant level despite decreasing spider species richness with increasing plant species richness. Plant phylogenetic diversity had no consistent effect on spider FD. The results contrast with the negative effect of diversity on spider species richness and suggest that functional redundancy among spiders decreased with increasing plant richness through non-random species loss. Moreover, increasing functional dissimilarity within spider assemblages with increasing plant richness indicates that the abundance distribution of predators in functional trait space affects ecological functions independent of predator species richness or the available trait space. While plant diversity is generally hypothesized to positively affect predators, our results only support this hypothesis for FD-and here particularly for trait distributions within the overall functional trait space-and not for patterns in species richness. Understanding the way predator assemblages affect ecosystem functions in such highly diverse, natural ecosystems thus requires explicit consideration of FD and its relationship with species richness.

Noncrop flowering plants restore top-down herbivore control in agricultural fields.

Herbivore populations are regulated by bottom-up control through food availability and quality and by top-down control through natural enemies. Intensive agricultural monocultures provide abundant food to specialized herbivores and at the same time negatively impact natural enemies because monocultures are depauperate in carbohydrate food sources required by many natural enemies. As a consequence, herbivores are released from both types of control. Diversifying intensive cropping systems with flowering plants that provide nutritional resources to natural enemies may enhance top-down control and contribute to natural herbivore regulation. We analyzed how noncrop flowering plants planted as “companion plants” inside cabbage (Brassica oleracea) fields and as margins along the fields affect the plant–herbivore–parasitoid–predator food web. We combined molecular analyses quantifying parasitism of herbivore eggs and larvae with molecular predator gut content analysis and a comprehensive predator community assessment. Planting cornflowers (Centaurea cynanus), which have been shown to attract and selectively benefit Microplitis mediator, a larval parasitoid of the cabbage moth Mamestra brassicae, between the cabbage heads shifted the balance between trophic levels. Companion plants significantly increased parasitism of herbivores by larval parasitoids and predation on herbivore eggs. They furthermore significantly affected predator species richness. These effects were present despite the different treatments being close relative to the parasitoids’ mobility. These findings demonstrate that habitat manipulation can restore top-down herbivore control in intensive crops if the right resources are added. This is important because increased natural control reduces the need for pesticide input in intensive agricultural settings, with cascading positive effects on general biodiversity and the environment. Companion plants thus increase biodiversity both directly, by introducing new habitats and resources for other species, and indirectly by reducing mortality of nontarget species due to pesticides.This study provides a comprehensive assessment of how habitat manipulation affects biocontrol services of a natural enemy community including both parasitoids and generalist predators. The trophic interactions between pests, parasitoids and predators were determined to achieve a better systemic understanding of top-down herbivore control, which can be strengthened when natural enemies complement each other or dampened by intraguild interactions. Our approach to selectively enhance the third trophic level to counteract specific herbivores was successful for both predators and parasitoids. Our results show significant positive effects of companion plants on predation of pest eggs and parasitism of pest larvae. Importantly, our data also suggest that carabids, staphylinids and spiders do not substantially interfere with parasitoid biocontrol as parasitoid DNA was rarely detected in predator guts.

Mammal predator and prey species richness are strongly linked at macroscales

Predator-prey interactions play an important role for species composition and community dynamics at local scales, but their importance in shaping large-scale gradients of species richness remains unexplored. Here, we use global range maps, structural equation models (SEM), and comprehensive databases of dietary preferences and body masses of all terrestrial, non-volant mammals worldwide, to test whether (1) prey bottom-up or predator top-down relationships are important drivers of broad-scale species richness gradients once the environment and human influence have been accounted for, (2) predator-prey richness associations vary among biogeographic regions, and (3) body size influences large-scale covariation between predators and prey. SEMs including only productivity, climate, and human factors explained a high proportion of variance in prey richness (R2=0.56) but considerably less in predator richness (R2=0.13). Adding predator-to-prey or prey-to-predator paths strongly increased the explained variance in both cases (prey R2=0.79, predator R2=0.57), suggesting that predator-prey interactions play an important role in driving global diversity gradients. Prey bottom-up effects prevailed over productivity, climate, and human influence to explain predator richness, whereas productivity and climate were more important than predator top-down effects for explaining prey richness, although predator top-down effects were still significant. Global predator-prey associations were not reproduced in all regions, indicating that distinct paleoclimate and evolutionary histories (Africa and Australia) may alter species interactions across trophic levels. Stronger cross-trophic-level associations were recorded within categories of similar body size (e.g., large prey to large predators) than between them (e.g., large prey to small predators), suggesting that mass-related energetic and physiological constraints influence broad-scale richness links, especially for large-bodied mammals. Overall, our results support the idea that trophic interactions can be important drivers of large-scale species richness gradients in combination with environmental effects.

Herbivore species richness, composition and community structure mediate predator richness effects and top-down control of herbivore biomass

Changes in predator species richness can have important consequences for ecosystem functioning at multiple trophic levels, but these effects are variable and depend on the ecological context in addition to the properties of predators themselves. Here, we report an experimental study to test how species identity, community attributes, and community structure at the herbivore level moderate the effects of predator richness on ecosystem functioning. Using mesocosms containing predatory insects and aphid prey, we independently manipulated species richness at both predator and herbivore trophic levels. Community structure was also manipulated by changing the distribution of herbivore species across two plant species. Predator species richness and herbivore species richness were found to negatively interact to influence predator biomass accumulation, an effect which is hypothesised to be due to the breakdown of functional complementarity among predators in species-rich herbivore assemblages. The strength of predator suppression of herbivore biomass decreased as herbivore species richness and distribution across host plants increased, and positive predator richness effects on herbivore biomass suppression were only observed in herbivore assemblages of relatively low productivity. In summary, the study shows that the species richness, productivity and host plant distribution of prey communities can all moderate the general influence of predators and the emergence of predator species richness effects on ecosystem functioning.

Intercropping With Aromatic Plants Decreases Herbivore Abundance, Species Richness, and Shifts Arthropod Community Trophic Structure

Habitat management is important for the regulation of arthropod community structures to reduce pest populations in orchard ecosystems, but there is limited information on how intercropping with aromatic plants regulates arthropod community structure at trophic levels. We selected four aromatic plants, Mentha canadensis L., Agerarum houstonianum Mill, Tagetes patula L., and Ocimum basilicum L., as intercrops in apple orchard to analyze the effects of intercropping on herbivore and predator abundance, species richness, and to measure the changes of trophic levels of the associated arthropod community. We found that intercropping with aromatic plants, compared with intercropping with natural herb vegetation, significantly reduced herbivore abundance and species richness by 25.62 and 11.6%, respectively. Intercropping significantly increased predator abundance and species richness by 18.78 and 15.6%, respectively, with predator abundance most notably affected during the flowering period of aromatic plants. Furthermore, herbivore abundance and species richness were strongly negatively correlated with predator species richness. Intercropping affected herbivore-predator dynamics, with an observed increase in the ratios of predator abundance and richness to herbivore abundance and richness. Our results indicate that intercropping with aromatic plants in apple orchards may play a role in the observed shift from a herbivore-dominated to a predator-dominated trophic structure, which is likely to have important flow-on effects on arthropod community structure.

Effects of an invasive alien tree on the diversity and temporal dynamics of an insect assemblage on an oceanic island

Native vegetation is frequently replaced by alien plants on isolated oceanic islands. The effects of such replacements by invasive plants on the diversity and temporal dynamics of island-endemic insects remain unclear. We examined flying insect communities using Malaise traps on the small island of Nishi-jima in the oceanic Ogasawara Archipelago in the northwestern Pacific. On the island, an alien tree, Casuarinaequisetifolia, has become dominant, occupying 57.3 % of the vegetation area. The species richness, composition, and abundance of pollinators (bees), predators (wasps), and wood-boring beetles (cerambycids, mordellids, and elaterids) were compared in each summer season of 4 years among three vegetation types: C. equisetifolia forest, natural forest, and grassland. In the traps, 82.3 % of species captured were endemic to the archipelago. The grassland harbored the highest species richness of native bees and wasps, whereas the natural forest had the highest species richness of native wood-boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups. Principal response curves indicated that differences in species composition among the three vegetation types were consistent through time for all insect groups. Most insect species were more abundant in natural forest or grassland than in C. equisetifolia forest. Standard deviations in both the numbers of individuals and species estimated under a Bayesian framework suggested that annual fluctuations of abundance and species density were similar among vegetation types (except for elaterid abundance). Therefore, replacement by C. equisetifolia has likely altered insect species composition but has not necessarily dramatically affected the temporal dynamics of insect assemblages on the island.

A simple plant mutation abets a predator‐diversity cascade

Resource consumption often increases with greater consumer biodiversity. This could result either from complementarity among consumers or the inclusion of particular key species, and it is often difficult to differentiate between these two mechanisms. We exploited a simple plant mutation (reduced production of surface waxes) to alter foraging within a community of aphid predators, and thus perhaps shift the nature of resulting predator diversity effects. We found that greater predator species richness dramatically increased prey suppression and plant biomass only on mutant, reduced-wax pea plants (Pisum sativum). On pea plants from a sister line with wild type, waxier plant surfaces, predator species richness did not influence predators' impacts on herbivores or plants. Thus, a change in plant surface structure acted to turn on, or off, the cascading effects of predator diversity. Greater predator richness encouraged higher densities of true predators but did not lead to greater reproduction by a parasitoid, Aphidius ervi; fecundity of each natural enemy species was similar for the two plant types. Behavioral observations indicated that although A. ervi was less likely to forage within species-rich predator communities, low-wax plants mitigated this interference by encouraging generally greater A. ervi foraging and thus high rates of aphid dislodgement (aphids dropped from plants to escape A. ervi, but not the other predators). Thus, only species-rich, low-wax plants simultaneously encouraged strong species-specific effects of A. ervi, and strong complementarity among the other predator species. In summary, our study provides evidence that diversity effects in predator assemblages are sensitive to habitat characteristics. Further, we show that a simple plant morphological trait, controlled by a single gene mutation, can dramatically alter the cascading effects of predator species richness on herbivores and plants.

Ecological release in White Sands lizards.

Ecological opportunity is any change that allows populations to escape selection from competition and predation. After encountering ecological opportunity, populations may experience ecological release: enlarged population size, broadened resource use, and/or increased morphological variation. We identified ecological opportunity and tested for ecological release in three lizard colonists of White Sands, New Mexico (Sceloporus undulatus, Holbrookia maculata, and Aspidoscelis inornata). First, we provide evidence for ecological opportunity by demonstrating reduced species richness and abundance of potential competitors and predators at White Sands relative to nearby dark soils habitats. Second, we characterize ecological release at White Sands by demonstrating density compensation in the three White Sands lizard species and expanded resource use in White Sands S. undulatus. Contrary to predictions from ecological release models, we observed directional trait change but not increased trait variation in S. undulatus. Our results suggest that ecological opportunity and ecological release can be identified in natural populations, especially those that have recently colonized isolated ecosystems.

Entomopathogen biodiversity increases host mortality

In biological control communities, greater predator species richness often strengthens pest suppression. The impacts and importance of species richness in insect-killing pathogen (entomopathogen) communities, however, has received less attention. Here, we manipulated species richness within a community of three soil-dwelling entomopathogenic nematodes (Heterorhabditis megidis, Steinernema carpocapsae, Steinernema feltiae) and one fungus (Beauveria bassiana), and measured resulting effects on mortality of Colorado potato beetle (Leptinotarsa decemlineata) and wax moth (Galleria mellonella) hosts. When potato beetles were the focal host, increasing pathogen species richness led to a linear increase in host mortality. This diversity effect appeared to result primarily from the pairing of nematodes and fungus within diverse communities; these pairings always produced host mortality that exceeded predictions based on the impact of nematodes or fungus alone. We then conducted an experiment using wax moth as the focal host, because this allowed greater replication, and included two different soil types to see if changing the foraging environment altered niche differences (and thus complementarity) among pathogen species. We again found an increase in host mortality at higher pathogen richness levels due to the apparent nematode–fungus synergism, and this effect was detected across both soil types. Our findings suggest that the positive effects of species richness commonly observed among predator species may extend to pathogen communities as well, such that conserving pathogen biodiversity may carry additional benefits for biological control.

Top-Down Control of Herbivory by Birds and Bats in the Canopy of Temperate Broad-Leaved Oaks (Quercus robur)

The intensive foraging of insectivorous birds and bats is well known to reduce the density of arboreal herbivorous arthropods but quantification of collateral leaf damage remains limited for temperate forest canopies.We conducted exclusion experiments with nets in the crowns of young and mature oaks, Quercus robur, in south and central Germany to investigate the extent to which aerial vertebrates reduce herbivory through predation. We repeatedly estimated leaf damage throughout the vegetation period.Exclusion of birds and bats led to a distinct increase in arthropod herbivory, emphasizing the prominent role of vertebrate predators in controlling arthropods. Leaf damage (e.g., number of holes) differed strongly between sites and was 59% higher in south Germany, where species richness of vertebrate predators and relative oak density were lower compared with our other study site in central Germany. The effects of bird and bat exclusion on herbivory were 19% greater on young than on mature trees in south Germany.Our results support previous studies that have demonstrated clear effects of insectivorous vertebrates on leaf damage through the control of herbivorous arthropods. Moreover, our comparative approach on quantification of leaf damage highlights the importance of local attributes such as tree age, forest composition and species richness of vertebrate predators for control of arthropod herbivory.

Disentangling the effects of predator hunting mode and habitat domain on the top-down control of insect herbivores

1. Polyphagous predatory invertebrates play a key role in the top-down control of insect herbivores. However, predicting predation risk for herbivores is not a simple function of predator species richness. Predation risk may be reduced or enhanced depending on the functional characteristics predator species. We predict that where predator species spatially overlap this will reduce predation risk for herbivores by allowing negative inter-specific interaction between predators to occur. Where increased predation risk occurs, we also predict that this will have a cascading effect through the food chain reducing plant growth. 2. We used a substitutive replicated block design to identify the effect of similarity and dissimilarity in predator hunting mode (e.g. 'sit and wait', 'sit and pursue', and 'active') and habitat domain (e.g. canopy or ground) on the top-down control of planthoppers in grasslands. Predators included within the mesocosms were randomly selected from a pool of 17 local species. 3. Predation risk was reduced where predators shared the same habitat domain, independent of whether they shared hunting modes. Where predators shared the same habitat domains, there was some evidence that this had a cascading negative effect on the re-growth of grass biomass. Where predator habitat domains did not overlap, there were substitutable effects on predation risk to planthoppers. Predation risk for planthoppers was affected by taxonomic identity of predator species, i.e. whether they were beetles, spiders or true bugs. 4. Our results indicated that in multi-predator systems, the risk of predation is typically reduced. Consideration of functional characteristics of individual species, in particular aspects of habitat domain and hunting mode, are crucial in predicting the effects of multi-predator systems on the top-down control of herbivores.

Plant diversity and the stability of foodwebs

Insect outbreaks in forest and agriculture monocultures led Charles Elton to propose, a half-century ago, that higher plant diversity stabilized animal foodweb dynamics in natural ecosystems. We tested this hypothesis by studying arthropod community dynamics in a long-term experimental manipulation of grassland plant species diversity. Over the course of a decade, we found that higher plant diversity increased the stability (i.e. lowered year-to-year variability) of a diverse (>700 species) arthropod community across trophic levels. As the number of plant species increased, the stability of both herbivore and predator species richness and of total herbivore abundance increased. The underlying mechanisms driving these diversity-stability relationships were plant diversity, via effects on primary productivity and plant community stability, and portfolio effects. Taken together, our results show that higher plant diversity provides more temporally consistent food and habitat resources to arthropod foodwebs. Consequently, actively managing for high plant diversity may have stronger than expected benefits for increasing animal diversity and controlling pest outbreaks.

Increased primary production shifts the structure and composition of a terrestrial arthropod community

Numerous studies have examined relationships between primary production and biodiversity at higher trophic levels. However, altered production in plant communities is often tightly linked with concomitant shifts in diversity and composition, and most studies have not disentangled the direct effects of production on consumers. Furthermore, when studies do examine the effects of plant production on animals in terrestrial systems, they are primarily confined to a subset of taxonomic or functional groups instead of investigating the responses of the entire community. Using natural monocultures of the salt marsh cordgrass Spartina alterniflora, we were able to examine the impacts of increased plant production, independent of changes in plant composition and/or diversity, on the trophic structure, composition, and diversity of the entire arthropod community. If arthropod species richness increased with greater plant production, we predicted that it would be driven by: (1) an increase in the number of rare species, and/or (2) an increase in arthropod abundance. Our results largely supported our predictions: species richness of herbivores, detritivores, predators, and parasitoids increased monotonically with increasing levels of plant production, and the diversity of rare species also increased with plant production. However, rare species that accounted for this difference were predators, parasitoids, and detritivores, not herbivores. Herbivore species richness could be simply explained by the relationship between abundance and diversity. Using nonmetric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM), we also found significant changes in arthropod species composition with increasing levels of production. Our findings have important implications in the intertidal salt marsh, where human activities have increased nitrogen runoff into the marsh, and demonstrate that such nitrogen inputs cascade to affect community structure, diversity, and abundance in higher trophic levels.

Reduced predator species richness drives the body gigantism of a frog species on the Zhoushan Archipelago in China

1. Shifts in the body size of insular vertebrates have been an interesting theme in ecological and evolutionary studies. Four primary factors, including predation pressures, resource availability, inter-species competition and immigrant selection, have been proposed to explain the trend in insular body size. Life-history theory predicts that body size, average age, the proportion of old-aged members and the density of insular populations are negatively correlated with predator species richness, and that body size and population density are positively related to resource availability. The niche expansion hypothesis argues that a positive relationship is expected to exist between insular body size and prey size, which varies in response to extinction due to small or large competitors. The immigrant hypothesis predicts that insular body size is positively correlated with distance to the mainland. 2. We tested these hypotheses by using populations of rice frogs Rana limnocharis on 20 islands in the Zhoushan Archipelago and two sites of nearby mainland China. 3. The body size (snout-vent length) of rice frogs on half of the islands was larger before and after the variable of age was controlled for; rice frog density and prey availability was higher and prey size was larger on most of the islands as compared to the two mainland sites. On the islands, the body size and other features [e.g. average age, the proportion of old-aged frogs (ages 3 and 4) and density] of the rice frogs were negatively associated with predator species richness; female body size and other features were positively associated with prey availability. The inference of multivariate linear models based on corrected Akaike Information Criterion (AIC(c) ) showed that the relative importance of predator species richness on body size and each of the other features was larger than that of prey availability, prey size and distance to the mainland. In addition, the parameters for predator species richness were all negative. 4. The results provided strong support for the life-history theory of predation pressures, but weak evidence for the life-history theory of prey availability, the niche expansion or the immigrant hypothesis. The reduced predator species richness was a dominant factor contributing to the body gigantism of rice frogs on the islands.

Arthropod colonisation of a debris-covered glacier

The largest debris-covered glacier in the Alps (Miage Glacier, western Italian Alps) has been studied to explore the effects of debris-cover extent and depth on the spatial distribution of ground-dwelling arthropods. A multitaxa approach has been used to compare taxa richness and distribution to the functional role (dietary habits) of each taxon along the glacier tongue. Spiders and ground beetles have been studied in detail. Taxa richness declines with distance from the wooded sites (in front of the glacier tongue) to those above the glacier tongue. At each of the supraglacial sites, spiders, ground beetles, aphids, springtails and flies were found. A change in the dominance of the different functional roles was observed along the tongue. Wooded sites are characterised by predatory (e.g. spiders, beetles), detrivore (e.g. springtails and certain flies), phytophagous (e.g. aphids, certain beetles) and parasitoid (e.g. certain wasps) assemblages, whereas at the debris-covered sites, aphids, flies and springtails are likely to be prey for spiders and beetles. The species richness of the predominant predators (spiders and beetles) shows a positive relationship with vegetation cover and debris thickness. Two mutually exclusive spider and ground beetle assemblages were found; one within the debris cover and one within the wooded sites. In our opinion, debris-covered glaciers are acting as a refuge area for the cryophil stenotherm species living at higher altitudes which descend the glacial tongue to lower elevations. A similar hypothesis supports the biogeographical interpretation of the distribution of many boreo-alpine relict species in the Alps. We discuss our results in the light of possible future scenarios which suggest an increase in debris cover with global warming.

Correlated geographic variation in predation risk and antipredator behaviour within a wide‐ranging snake species (Notechis scutatus, Elapidae)

AbstractGeographic variation in antipredator behaviour within wide‐ranging species may be driven by both genetic and environmental influences. We quantified antipredator responses in neonatal (laboratory born, n = 555) and adult (field caught, n = 346) tiger snakes (Notechis scutatus) from 11 mainland and island sites in southern Australia. We used these data to test predictions from Bonnet et al.'s hypothesis that the vigour of antipredator responses in this species reflects behavioural plasticity (in turn, driven by an individual snake's exposure to predators during its lifetime) rather than by genetic variation in this trait. We used the number of predator taxa in each area as an index of predator risk. As predicted, adult snakes from predator‐rich areas had more vigorous defensive responses when handled, whereas neonatal behaviour (although also variable among populations) was unrelated to predator species richness. Adult males bit more readily than adult females (as expected from the greater predation exposure of males during mate searching) but no such sex difference was evident in neonates. Although alternative models remain possible, our data are most consistent with the hypothesis that geographic divergence in antipredator tactics within this species primarily reflects developmentally plastic responses to local predation risk.

Cascading diversity effects transmitted exclusively by behavioral interactions

Consumer diversity generally increases resource consumption. Consumers can also impact other species by altering their behavior, but it is unclear how such nonconsumptive effects scale with diversity. We independently manipulated predator species richness and the consumptive and nonconsumptive effects of predator communities to measure the role of each factor in protecting Brassica oleracea plants from caterpillar herbivory. Plant biomass was greatest when diverse predator assemblages induced antipredator behaviors in herbivores, an effect not further strengthened when predators could also kill caterpillars. Predators within diverse communities were more likely to forage on plants and to disrupt herbivore feeding, reflecting greater aversion to foraging among conspecific than heterospecific competitors. Predator diversity, therefore, initiated behavioral changes at the predator and then herbivore trophic levels, both to the benefit of plants. Our results indicate that strong, emergent species-richness effects can be transmitted entirely through behavioral interactions, independent of resource consumption.

Conserving the benefits of predator biodiversity

Organisms at higher trophic levels often face a disproportionate risk of local or regional extinction, while at the same time many ecosystems are being invaded by non-native predators. Global environmental change fosters both processes, further altering predator biodiversity. Thus, there has been growing interest in how predator species richness impacts ecosystem functioning. Manipulative experiments have revealed that complementarity and sampling effects, two mechanisms commonly found to underlie diversity effects at other trophic levels, also commonly impact the relationship between predator diversity and prey suppression. Intraguild predation and non-consumptive (behaviorally-mediated) effects on prey, two mechanisms without direct analogs among plants, also strongly impact predator-diversity effects. Predator diversity studies are particularly relevant to conservation because they focus on the trophic group that is most prone to extinction, and because they nearly always measure diversity effects that span several trophic levels. Predator invasions may partly offset species-richness losses to extinction, but because invasive predators typically reach much higher densities and exert stronger impacts on prey than do native species, and because they also displace ecologically-similar native predators, invasion is likely to disrupt natural predator function. A framework for predicting which predator-diversity mechanisms are likely to operate in a given community, and experiments that span more realistic spatiotemporal scales and include large vertebrate predators, are needed to improve the relevance of predator-diversity experiments to conservation decision-making in the future.