Predicting the impact of carnivores on plants has challenged community and food web ecologists for decades. At the same time, the role of predators in the evolution of herbivore dietary specialization has been an unresolved issue in evolutionary ecology. Here, we integrate these perspectives by testing the role of herbivore diet breadth as a predictor of top-down effects of avian predators on herbivores and plants in a forest food web. Using experimental bird exclosures to study a complex community of trees, caterpillars, and birds, we found a robust positive association between caterpillar diet breadth (phylodiversity of host plants used) and the strength of bird predation across 41 caterpillar and eight tree species. Dietary specialization was associated with increased enemy-free space for both camouflaged (n = 33) and warningly signaled (n = 8) caterpillar species. Furthermore, dietary specialization was associated with increased crypsis (camouflaged species only) and more stereotyped resting poses (camouflaged and warningly signaled species), but was unrelated to caterpillar body size. These dynamics in turn cascaded down to plants: a metaanalysis (n = 15 tree species) showed the beneficial effect of birds on trees (i.e., reduced leaf damage) decreased with the proportion of dietary specialist taxa composing a tree species' herbivore fauna. We conclude that herbivore diet breadth is a key functional trait underlying the trophic effects of carnivores on both herbivores and plants.ecological specialization | host specificity | plant-herbivore interactions | tritrophic interactions | trophic cascade P redicting the strength of trophic interactions is a major goal in ecology. Because most natural ecosystems contain numerous coexisting species at each trophic level, achieving this goal necessarily involves the integration of theory in evolutionary, community, and food web ecology. In this context, evolutionary ecology explains how traits of organisms adapt them to a fundamental trade-off between resource acquisition and mortality risk from natural enemies (1, 2); community ecology theory links the many patterns and consequences of species interactions to the diversity of traits of those species (2); and food web ecology subsumes this diversity into patterns of trophic structure and dynamics, such as a trophic cascade (3). The recognition that functional traits of species can drive the indirect positive effect of carnivores on plant biomass [trophic cascades broadly defined (4, 5)] provides important insight into the causes of variation in these dynamics (1,6). An emerging understanding of the functional traits mediating trophic cascade strength includes traits of herbivores that facilitate predator avoidance (7-10), or provide constitutive (11, 12) or induced resistance to predation (13). These examples identify antipredator traits of herbivores as an important mediator of top-down effects on plants within individual tritrophic food chains. However, the role of antipredator (or other) traits of herbivores is curr...

The effectiveness of anti‐predator traits, such as warning signals and camouflage, has rarely been quantified from a phylogenetic community ecology perspective. Here we use a phylogenetic comparative analysis to test the association between several putative anti‐predator traits and bird predation risk in an assemblage of caterpillar species. We synthesize eight years of field and laboratory study of a temperate forest community, including a four‐year bird exclusion experiment that provided comparative measures of bird predation risk for 38 caterpillar species from a phylogenetic community. We then conducted a phylogenetic generalized least‐squares and information‐theoretic model selection analysis of warning signals (aposematism or mimicry), camouflage (crypsis or masquerade), and behavioral responses to physical attack as predictors of bird predation, while also accounting for putatively important effects of the abundance, mean body size, and phenology of caterpillar species. The most behaviorally specialized caterpillar species possessing warning signals experienced the lowest bird predation risk, supporting aposematism theory and highlighting the role of prey behavior in the visual signaling of predators. Among the camouflaged caterpillar species, those with the greatest latency to detection by human proxy predators experienced the lowest bird predation risk, supporting camouflage theory. Caterpillar behavioral responses to physical attack, however, predicted increased bird predation risk among camouflaged caterpillars. Although caterpillar abundance, body size, and phenology were expected to be important based on inference from optimal foraging theory and previous field observations, these factors had limited predictive power. This study provides methodologically unique evidence for the importance of morphological and behavioral components of primary, visual defenses of caterpillars against their avian predators in a natural community.

Abstract.Direct and indirect effects of predators are highly variable in complex communities, and understanding the sources of this variation is a research priority in community ecology. Recent evidence indicates that herbivore community structure is a primary determinant of predation strength and its cascading impacts on plants. In this study, we use variation in herbivore community structure among plant species to experimentally test two hypotheses in a temperate forest food web. First, variation in the strength of predator effects, such as ant predation of caterpillars, is predicted to be density dependent, exhibiting stronger effects when prey abundance is high (density-dependent predation hypothesis). Second, mutualistic interactions between ants and sap-feeding herbivores are expected to increase the abundance of predatory ants, strengthening predation effects on herbivores with cascading effects on host plants (keystone mutualism hypothesis). Using a large-scale predator exclusion experiment across eight dominant tree species, we tracked changes in insect density on 862 plants across two years, recording 2,322 ants, 1,062 sap-feeders, 5,322 caterpillars, and quantifying herbivory on 199, 338 leaves. In this experiment, density-dependent predation did not explain variation in the direct or indirect effects of ants on caterpillars and herbivory. In partial support of the keystone mutualism hypothesis, sap-feeders strengthened top-down effects of ants on caterpillars under some conditions. However, stronger ant predation of caterpillars did not lead to measurable trophic cascades on trees occupied by sap-feeders. Instead, the presence of sap-feeders was associated with increased per capita feeding damage by caterpillars, and this bottom-up effect attenuated the indirect effects of ants on host plants. These findings demonstrate that examining the multi-trophic impacts of mutualisms and predation in the context of the broader community can reveal patterns otherwise masked by compensatory interactions.

Rock daisies (Perityleae; Compositae) are a diverse clade of seven genera and ca. 84 minimum‐rank taxa that mostly occur as narrow endemics on sheer rock cliffs throughout the southwest United States and northern Mexico. Taxonomy of Perityleae has traditionally been based on morphology and cytogenetics. To test taxonomic hypotheses and utility of characters emphasized in past treatments, we present the first densely sampled molecular phylogenies of Perityleae and reconstruct trait and chromosome evolution. We inferred phylogenetic trees from whole chloroplast genomes, nuclear ribosomal cistrons, and hundreds of low‐copy nuclear genes using genome skimming and target capture. Discordance between sources of molecular data suggests an underappreciated history of hybridization in Perityleae. Phylogenies support the monophyly of subtribe Peritylinae, a distinctive group possessing a four‐lobed disc corolla; however, all of the phylogenetic trees generated in this study reject the monophyly of the most species‐rich genus, Perityle, as well as its sections: Perityle sect. Perityle, Perityle sect. Laphamia, and Perityle sect. Pappothrix. Using reversible jump MCMC, our results suggest that morphological characters traditionally used to classify members of Perityleae have evolved multiple times within the group. A base chromosome number x = 9 gave rise to higher base numbers in subtribe Peritylinae (x = 12, 13, 16, 17, 18, and 19) through polyploidization, followed by ascending or descending dysploidy. Most taxa constitute a monophyletic lineage with a base chromosome number of x = 17, with multiple neo‐polyploidization events. These results demonstrate the advantages and obstacles of next‐generation sequencing approaches in synantherology while laying the foundation for taxonomic revision and comparative study of the evolutionary ecology of Perityleae.