Characterising sclerophylly: some mechanical properties of leaves from heath and forest.

Although sclerophylly is widespread through the world and is often the dominant leaf-form in mediterranean climates, the mechanical properties of sclerophyllous leaves are poorly understood. The term "sclerophyllous" means hard-leaved, but biologists also use terms such as tough, stiff and leathery to describe sclerophyllous leaves. The latter term has no precise definition that allows quantification. However, each of the former terms is well-defined in materials engineering, although they may be difficult or sometimes inappropriate to measure in leaves because of their size, shape or composite and anisotropic nature. Two of the most appropriate and practically applicable mechanical properties of sclerophyllous leaves are "strength" and "toughness", which in this study were applied using punching, tearing and shearing tests to 19 species of tree and shrub at Wilson's Promontory, Australia. The results of these tests were compared with leaf specific mass (LSM) and a sclerophylly index derived from botanists' ranks. Principal components analysis was used to reduce the set of mechanical properties to major axes of variation. Component 1 correlated strongly with the botanists' ranks. Overall, leaves ranked as sclerophyllous by botanists were both tough and strong in terms of punching and tearing tests. In addition, tough and strong leaves typically had high toughness and strength per unit leaf thickness. There was also a significant correlation between component 1 and LSM. Although more detailed surveys are required, we argue that sclerophylly should be defined in terms of properties that have precise meanings and are measurable, such as toughness and strength, and that relate directly to mechanical properties as implicit in the term.