Integrating the host plant phylogeny, plant traits, intraspecific competition and repeated measures using a phylogenetic mixed model of field behaviour by polyphagous herbivores, the leaf-cutting ants

Herbivores use a wide range of factors to choose their host, including their own physiological states, physical characteristics of plants, and the degree of competition. Field observations of herbivores in their native habitats provide a means for simultaneously estimating the relative importance of these factors, but statistical analysis of all these factors may be challenging. Here we used a 7-wk dataset of leaf-cutting ant (Atta cephalotes) foraging in a diverse neotropical arboretum containing 193 tree species (822 trees) to examine the relative role of tree phylogeny, territoriality and tree functional characteristics using a phylogenetic generalized least squares (PGLS) model. We observed that 54 tree species (117 trees) were foraged by the ants. This pattern was not random, but reflected known features of leaf-cutting ant foraging behaviour, such as the preference for larger trees and the decreased likelihood of foraging at the periphery of a colonys territory. However, random effects such as tree phylogeny, the identity of individual trees and colony-specific effects, explained most of the variation in foraging data. A significant phylogenetic effect on foraging likelihood ({lambda} = 0.28), together with repeated measures of foraging on the same tree species, allowed estimation of relative palatability for each plant species. PGLS models can be flexibly scaled to include other covariates for even more complex investigation of foraging behaviour, and the link function can be modified to include the amount of plants foraged. As a result, PGLS can be used as a flexible framework for the study of LCA foraging.