The leaf anatomy of species representing all seven genera of the Penaeaceae was studied by light and scanning electron microscopy. Due to variability and inconsistency, leaf anatomical characters are not regarded as particularly useful for systematics within or among genera in this family. Across the family, a number of taxa exhibit a trend towards amphistomatous, isobilateral leaves, generally associated with increased leaf thickness and amount of palisade mesophyll. This trend is not apparent in closely related families, e.g. Alzateaceae, Cryp-teroniaceae, Oliniaceae and Rhynchocalycaceae. Most species are found in comparatively mesic habitats and it is difficult to postulate primary xeromorphic trends in leaf anatomy. The sclerophylly encountered across the family as a whole is likely to be related to paucity of soil nutrients rather than a response to water stress.
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John Dickie (1999) studied this question.
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