Seeing the trees for the leaves - oaks as mosaics for a host-specific moth

From the perspective of a specialist herbivore, how homogenous are individual tree crowns as patches of habitat? We partitioned variation in physical and chemical host leaf traits and in the abundance and performance of a specialist oak leaf miner, Tischeria ekebladella, into variation at different hierarchical levels. For the phenolic contents of the leaves, we examined variation among oak stands, among trees within stands and among branches within trees. For leaf size and water content, we assessed variation among trees within a single stand, among shoots within trees, and among leaves within shoots. For moth abundance and performance, we examined variation across all levels: among oak stands, among trees within stands, among branches within trees, among shoots within branches and among leaves and insect individuals. For measures of phenolic contents, we found little variation among stands but substantial variation among individual trees. Yet, a tree particularly rich in a given compound was often comparatively poor in another. At a finer spatial scale, the phenolic composition of individual parts of a single tree was quite consistent, whereas leaf weight and water content varied widely within individual tree crowns. Moth abundances varied more among shoots within branches than at any other spatial level, whereas moth survival showed equal levels of variation within individual shoots as among separate oak stands. Likewise, for four other measures of larval performance (assessed at the level of trees and lower), we found more variation within than between trees. In conclusion, the large variation observed in the performance of a specialist moth and in the physical traits of the leaves among different parts of single tree crowns refutes the image of an oak tree as an 'island' of internally homogeneous quality. Hence, we may expect little evolutionary adaptation of T. ekebladella at the scale of individual trees. The moths may instead evolve to behaviourally select their resource at a very fine scale. Large variation within trees also calls for extensive replication within trees in ecological sampling designs and/or the sampling of maximally similar leaves.