The evolutionary relations of sunken, covered, and encrypted stomata to dry habitats in Proteaceae.

Sunken, covered, and encrypted stomata have been anecdotally linked with dry climates and reduced transpiration and therefore have been used to infer dry palaeoclimates from fossils. This study assesses the evolutionary and ecological associations of such stomatal protection in a model system-the diverse southern hemisphere family Proteaceae. Analyses were based on the morphology of over 1400 Australian, South African, New Caledonian, New Zealand, and South American species, anatomy of over 300 of these species, and bioclimatic data from all 1109 Australian species. Ancestral state reconstruction revealed that five or six evolutionary transitions explain over 98% of the dry climate species in the family, with a few other, minor invasions of dry climates. Deep encryption, i.e., stomata in deep pits, in grooves, enclosed by tightly revolute margins or strongly overarched by cuticle, evolved at least 11 times in very dry environments. Other forms of stomatal protection (sunken but not closely encrypted stomata, papillae, and layers of hairs covering the stomata) also evolved repeatedly, but had no systematic association with dry climates. These data are evidence for a strong distinction in function, with deep encryption being an adaptation to aridity, whereas broad pits and covered stomata have more complex relations to climate.

Solar radiation as a factor in the evolution of scleromorphic leaf anatomy in Proteaceae.

Species of the major Southern Hemisphere family, Proteaceae, have many scleromorphic anatomical structures in their leaves. Many of these structures (very thick cuticles and five anatomically distinct structures beneath the epidermis) are associated with the leaf surface exposed to direct light. These structures increase the path through which solar radiation must pass before reaching the mesophyll. In this study, such structures are proposed to protect the mesophyll from excess solar radiation, including photosynthetically active, ultraviolet, and possibly infrared radiation. Scleromorphic structures of the upper leaf surface and nonscleromorphic photoprotective structures (dense trichomes and papillae of the upper surface) occur almost exclusively in open vegetation. Open vegetation species of Proteaceae occur in oligotrophic and/or cold and/or dry places, where protection from light in excess of photosynthetic capacity and damage from ultraviolet light should be most important. Data from 123 species and a supertree constructed from available molecular phylogenies are used to show that the proposed photoprotective structures evolved many times within Proteaceae. In tests of correlated evolution, the proposed photoprotective structures are significantly associated with open vegetation, but not with dry habitats.