Leaf morphological traits as adaptations to multiple climate gradients.

Leaf morphological traits vary systematically along climatic gradients. However, recent studies in plant functional ecology have mainly analysed quantitative traits, while numerical models of species distributions and vegetation function have focused on traits associated with resource acquisition; both ignore the wider functional significance of leaf morphology.A dataset comprising 22 leaf morphological traits for 662 woody species from 92 sites, representing all biomes present in China, was subjected to multivariate analysis in order to identify leading dimensions of trait covariation (correspondence analysis), quantify climatic and phylogenetic contributions (canonical correspondence analysis with variation partitioning) and characterise co-occurring trait syndromes (-means clustering) and their climatic preferences.Three axes accounted for >20% of trait variation in both evergreen and deciduous species. Moisture index, precipitation seasonality and growing-season temperature explained 8%-10% of trait variation; family 15%-32%. Microphyll or larger, mid- to dark green leaves with drip tips in wetter climates contrasted with nanophyll or smaller glaucous leaves without drip tips in drier climates. Thick, entire leaves in less seasonal climates contrasted with thin, marginal dissected, aromatic and involute/revolute leaves in more seasonal climates. Thick, involute, hairy leaves in colder climates contrasted with thin leaves with marked surface structures (surface patterning) in warmer climates. Distinctive trait clusters were linked to the driest and most seasonal climates, for example the clustering of picophyll, fleshy and succulent leaves in the driest climates and leptophyll, linear, dissected, revolute or involute and aromatic leaves in regions with highly seasonal rainfall. Several trait clusters co-occurred in wetter climates, including clusters characterised by microphyll, moderately thick, patent and entire leaves or notophyll, waxy, dark green leaves. . The plastic response of size, shape, colour and other leaf morphological traits to climate is muted, thus their apparent shift along climate gradients reflects plant adaptations to environment at a community level as determined by species replacement. Information on leaf morphological traits, widely available in floras, could be used to strengthen predictive models of species distribution and vegetation function.