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Vicissitudes of leaves in a tropical rain forest in Madagascar

Published online by Cambridge University Press:  08 October 2009

Harold Heatwole*
Department of Zoology, North Carolina State University, Raleigh, NC 27695-7617, USA
Sybille Unsicker
Department of Biochemistry, Max Planck Institute for Chemical Ecology, Hans Knoell Str. 8, 07745 Jena, Germany
Lala Roger Andriamiarisoa
Departement Biologia Végétal, Université d'Antananarivo, Antananarivo, Madagascar
Margaret D. Lowman
Division of Natural Sciences, New College of Florida, 5800 Bay Shore Road, Sarasota FL 34243-2197, USA
1Corresponding author. Email:


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 unscathed by any agent. Individual leaves were attacked by up to five agents. There was a sharp decline in proportion of leaves affected by fungi, leaf miners, epiphyllae and mechanical breakage with increasing category of severity. 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 saplings at ground level, but with increasing height above the ground fungal attack on leaves becomes more prevalent, but attack by grazing insects less so.

Research Article
Copyright © Cambridge University Press 2009

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