Variation in xylem structure from tropics to tundra: evidence from vestured pits

Bordered pits play an important role in permitting water flow among adjacent tracheary elements in flowering plants. Variation in the bordered pit structure is suggested to be adaptive in optimally balancing the conflict between hydraulic efficiency (conductivity) and safety from air entry at the pit membrane (air seeding). The possible function of vestured pits, which are bordered pits with protuberances from the secondary cell wall of the pit chamber, could be increased hydraulic resistance or minimized vulnerability to air seeding. These functional hypotheses have to be harmonized with the notion that the vestured or nonvestured nature of pits contains strong phylogenetic signals (i.e., often characterize large species-rich clades with broad ecological ranges). A literature survey of 11,843 species covering 6,428 genera from diverse climates indicates that the incidence of vestured pits considerably decreases from tropics to tundra. The highest frequencies of vestured pits occur in deserts and tropical seasonal woodlands. Moreover, a distinctly developed network of branched vestures is mainly restricted to warm habitats in both mesic and dry (sub)tropical lowlands, whereas vestures in woody plants from cold and boreal arctic environments are usually minute and simple. A similar survey of the frequency of exclusively scalariform perforation plates illustrates that the major ecological trend of this feature is opposite that of vestured pits. These findings provide previously undescribed insights suggesting that vessels with vestured pits and simple perforation plates function as an efficient hydraulic system in plants growing in warm environments with periodical or continuous drought stress.