Vicissitudes of leaves in a tropical rain forest in Madagascar

A sample of 1834 leaves from 83 plants in 26 families was collected in tropical rain forest in Madagascar from three vertical strata: top of emergent trees (up to 37 m), top of trees in upper canopy (about 22 m), and shrubs and saplings at ground level. These leaves were examined for damage by seven different agents: fungi, epiphyllae, mechanical injury, galls, leaf miners, grazing insects and skeletonizing insects. Fungi affected more than 60% of the leaves and grazing insects 45-65%, with other agents each affecting 3-30%; fewer than 4% of the leaves escaped unscathedbyanyagent.Individualleaveswereattackedbyuptofiveagents.Therewasasharpdeclineinproportionof leavesaffectedbyfungi,leafminers,epiphyllaeandmechanicalbreakagewithincreasingcategoryofseverity.Grazing insects, mechanical injury, and perhaps galls, had greater impact at ground level than in the canopy, with fungi and skeletonizing insects showing the opposite pattern. Leaf miners had lower incidence in the canopy than elsewhere. The observed vertical stratification means that a tree not only needs to balance its defences to meet multiple threats in any stratum, but must adapt to a different suite of challenges during its lifetime. Attack by grazing insects and fungi are major challenges for saplingsat groundlevel, butwithincreasing heightabove thegroundfungal attackon leaves becomes more prevalent, but attack by grazing insects less so.