Interspecific and intraspecific variation in leaf toughness of Arctic plants in relation to habitat and nutrient supply

Leaf toughness is an important functional trait that confers resistance to herbivory and mechanical damage. We sought to determine how species composition, climate, seasonality, and nutrient availability influence leaf toughness in two types of tundra in northern Alaska. We measured leaf toughness as force to punch for 11 species of Arctic plants in tussock tundra and dry heath tundra at 17 sites distributed along a latitudinal gradient. Rubus chamaemorus L. and the graminoids occupied opposite ends of the leaf toughness spectrum, with Rubus chamaemorus requiring the least force to punch, whereas one of the graminoids, Eriophorum vaginatum L., required the most. Leaf toughness increased with mean summer temperature for Eriophorum vaginatum and Betula nana L., whereas it declined with warmer temperatures for the other species. Toughness of mature leaves of Eriophorum vaginatum did not vary through the growing season but declined significantly after senescence. Application of N and P fertilizer in an experimental site decreased leaf toughness in three species but had no effect on four others. Leaf toughness of four out of five species in dry heath was greater than for the same species in tussock tundra, but there was no difference in community-weighted mean toughness between tussock tundra and dry heath.