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How numerous tree species can coexist in diverse forest communities is a key question in community ecology. Whereas neutral theory assumes that species are adapted to common field conditions and coexist by chance, niche theory predicts that species are functionally different and coexist because they are specialized for different niches. We integrated biophysical principles into a mathematical plant model to determine whether and how functional plant traits and trade-offs may cause functional divergence and niche separation of tree species. We used this model to compare the carbon budget of saplings across 13 co-occurring dry-forest tree species along gradients of light and water availability. We found that species ranged in strategy, from acquisitive species with high carbon budgets at highest resource levels to more conservative species with high tolerances for both shade and drought. The crown leaf area index and nitrogen mass per leaf area drove the functional divergence along the simulated light gradient, which was consistent with observed species distributions along light gradients in the forest. Stomatal coordination to avoid low water potentials or hydraulic failure caused functional divergence along the simulated water gradient, but was not correlated to observed species distributions along the water gradient in the forest. The trait-based biophysical model thus explains how functional traits cause functional divergence across species and whether such divergence contributes to niche separation along resource gradients.

Plant hydraulic architecture has been studied extensively, yet we know little about how hydraulic properties relate to species' life history strategies, such as drought and shade tolerance. The prevailing theories seem contradictory. We measured the sapwood (K(s) ) and leaf (K(l) ) hydraulic conductivities of 40 coexisting tree species in a Bolivian dry forest, and examined associations with functional stem and leaf traits and indices of species' drought (dry-season leaf water potential) and shade (juvenile crown exposure) tolerance. Hydraulic properties varied across species and between life-history groups (pioneers vs shade-tolerant, and deciduous vs evergreen species). In addition to the expected negative correlation of K(l) with drought tolerance, we found a strong, negative correlation between K(l) and species' shade tolerance. Across species, K(s) and K(l) were negatively correlated with wood density and positively with maximum vessel length. Consequently, drought and shade tolerance scaled similarly with hydraulic properties, wood density and leaf dry matter content. We found that deciduous species also had traits conferring efficient water transport relative to evergreen species. Hydraulic properties varied across species, corresponding to the classical trade-off between hydraulic efficiency and safety, which for these dry forest trees resulted in coordinated drought and shade tolerance across species rather than the frequently hypothesized trade-off., (© 2011 The Authors. New Phytologist © 2011 New Phytologist Trust.)

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P₅₀) related to dry season leaf water potentials and stem and leaf traits. P₅₀-values ranged from -0.8 to -6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade-tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber-value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade-off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P₅₀, but pioneers and deciduous species had smaller hydraulic safety margins than shade-tolerants and evergreens. A trade-off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species., (© 2010 Blackwell Publishing Ltd.)