Water taken up through the bark is detected in the transpiration stream in intact upper-canopy branches.

Alternative water uptake pathways through leaves and bark complement water supply with interception, fog or dew. Bark water-uptake contributes to embolism-repair, as demonstrated in cut branches. We tested whether bark water-uptake could also contribute to supplement xylem-water for transpiration. We applied bandages injected with 2 H-enriched water on intact upper-canopy branches of Pinus sylvestris and Fagus sylvatica in a boreal and in a temperate forest, in summer and winter, and monitored transpiration and online isotopic composition (δ2 H and δ18 O) of water vapour, before sampling for analyses of δ2 H and δ18 O in tissue waters. Xylem, bark and leaf waters from segments downstream from the bandages were 2 H-enriched whereas δ18 O was similar to controls. Transpiration was positively correlated with 2 H-enrichment. Isotopic compositions of transpiration and xylem water allowed us to calculate isotopic exchange through the bark via vapour exchange, which was negligible in comparison to estimated bark water-uptake, suggesting that water-uptake occurred via liquid phase. Results were consistent across species, forests and seasons, indicating that bark water-uptake may be more ubiquitous than previously considered. We suggest that water taken up through the bark could be incorporated into the transpiration stream, which could imply that sap-flow measurements underestimate transpiration when bark is wet.

A novel optimization approach incorporating non-stomatal limitations predicts stomatal behaviour in species from six plant functional types.

The primary function of stomata is to minimize plant water loss while maintaining CO2 assimilation. Stomatal water loss incurs an indirect cost to photosynthesis in the form of non-stomatal limitations (NSL) via reduced carboxylation capacity (CAP) and/or mesophyll conductance (MES). Two optimal formulations for stomatal conductance (gs) arise from the assumption of each type of NSL. In reality, both NSL could coexist, but one may prevail for a given leaf ontogenetic stage or plant functional type, depending on leaf morphology. We tested the suitability of two gs formulations (CAP versus MES) on species from six plant functional types (C4 crop, C3 grass, fern, conifer, evergreen, and deciduous angiosperm trees). MES and CAP parameters (the latter proportional to the marginal water cost to carbon gain) decreased with water availability only in deciduous angiosperm trees, while there were no clear differences between leaf ontogenetic stages. Both CAP and MES formulations fit our data in most cases, particularly under low water availability. For ferns, stomata appeared to operate optimally only when subjected to water stress. Overall, the CAP formulation provided a better fit across all species, suggesting that sub-daily stomatal responses minimize NSL by reducing carboxylation capacity predominantly, regardless of leaf morphology and ontogenetic stage.

Plant size and leaf area influence phenological and reproductive responses to warming in semiarid Mediterranean species.

Changes in vegetative and reproductive phenology rank among the most obvious plant responses to climate change. These responses vary broadly among species, but it is largely unknown whether they are mediated by functional attributes, such as size or foliar traits. Using a manipulative experiment conducted over two growing seasons, we evaluated the responses in reproductive phenology and output of 14 Mediterranean semiarid species belonging to three functional groups (grasses, nitrogen-fixing legumes and forbs) to a ~3°C increase in temperature, and assessed how leaf and size traits influenced them. Overall, warming advanced flowering and fruiting phenology, extended the duration of flowering and reduced the production of flowers and fruits. The observed reduction in flower and fruit production with warming was likely related to the decrease in soil moisture promoted by this treatment. Phenological responses to warming did not vary among functional groups, albeit forbs had an earlier reproductive phenology than legumes and grasses. Larger species with high leaf area, as well as those with small specific leaf area, had an earlier flowering and a longer flowering duration. The effects of warming on plant size and leaf traits were related to those on reproductive phenology and reproductive output. Species that decreased their leaf area under warming advanced more the onset of flowering, while those that increased their vegetative height produced more flowers. Our results advance our understanding of the phenological responses to warming of Mediterranean semiarid species, and highlight the key role of traits such as plant size and leaf area as determinants of such responses.