Regeneration patterns, environmental filtering and tree species coexistence in a temperate forest.

Forest ecologists researching the functional basis of tree regeneration patterns and species coexistence often attempt to correlate traits with light-gradient partitioning. However, an exclusive focus on light can overlook other important drivers of forest dynamics. We measured light, temperatures, humidity and sapling densities in each of four phases of a forest dynamic mosaic in New Zealand: shaded understoreys, tree-fall gaps, treefern groves and clearings. We then measured leaf, wood and seed traits, as potential predictors of species' regeneration patterns. Saplings of 18 out of 21 species were significantly associated with one or other of the four phases, and associations were best predicted by a two-trait model (leaf size, wood density) explaining 51% of observed variation. Species associated with treefall gaps had traits favouring light pre-emption (large leaves, low-density wood), whereas those establishing in clearings mostly had small leaves and dense wood, traits probably conferring resistance to the frosts and summer water deficits that saplings were exposed to there. The dynamics of some forests cannot be explained adequately by light-gradient partitioning through a growth vs shade tolerance tradeoff, underpinned by the leaf economics spectrum. Consideration of multiple environmental filters and multiple traits will enhance understanding of regeneration patterns and species coexistence.