Marcescence and prostrate growth in tree ferns are adaptations to cold tolerance

Abstract

Cold tolerance strategies in plants vary from structural to biochemical permitting many plants to survive and grow on sites that experience freezing conditions intermittently. Although tree ferns occur predominantly across the tropics, they also occur in temperate zones and occasionally in areas that experience sub-zero temperatures, and how these large ferns survive freezing conditions is unknown. Many temperate tree fern taxa are marcescent – retaining whorls of dead fronds encircling the upper trunk – or develop short or prostrate trunks, possibly to insulate against frost damage to their trunks and growing crowns. We asked the following questions: 1) do global growth patterns and traits of tree ferns respond to freezing conditions associated with latitude and elevation, 2) do growth patterns of tree ferns in New Zealand vary along a temperature-related gradient, and 3) do marcescent tree fern skirts insulate the growing crown from sub-zero temperatures? To establish what morphological adaptations permitted the Cyatheales to occur in biomes that experience intermittent sub-zero temperatures and frost, we 1) reviewed the global distributions of these structural and morphological traits within the tree ferns (Cyatheales); 2) assessed the patterns of tree fern marcescence, and other traits potentially associated with cold tolerance (no trunk, prostrate, short-trunked) of nine taxa of the Cyatheales along environmental gradients across New Zealand; and 3) conducted a field experiment to assess the thermal insulation properties of tree fern marcescent skirts. We identified significant trends among growth forms, marcescence, and environmental gradients consistent with our hypothesis that these are adaptations to tolerate cold. Our field experiments provide quantitative evidence that marcescent skirts have a strong insulating effect on tree fern trunks. The Cyatheales have evolved several strategies to protect the pith cores of their trunks from extreme cold temperatures in temperate forests allowing them to capture niche space in environments beyond the tropics.