The validity of optimal leaf traits modelled on environmental conditions.

The ratio of leaf intercellular to ambient CO2 (χ) is modulated by stomatal conductance (gs ). These quantities link carbon (C) assimilation with transpiration, and along with photosynthetic capacities (Vcmax and Jmax ) are required to model terrestrial C uptake. We use optimization criteria based on the growth environment to generate predicted values of photosynthetic and water-use efficiency traits and test these against a unique dataset. Leaf gas-exchange parameters and carbon isotope discrimination were analysed in relation to local climate across a continental network of study sites. Sun-exposed leaves of 50 species at seven sites were measured in contrasting seasons. Values of χ predicted from growth temperature and vapour pressure deficit were closely correlated to ratios derived from C isotope (δ13 C) measurements. Correlations were stronger in the growing season. Predicted values of photosynthetic traits, including carboxylation capacity (Vcmax ), derived from δ13 C, growth temperature and solar radiation, showed meaningful agreement with inferred values derived from gas-exchange measurements. Between-site differences in water-use efficiency were, however, only weakly linked to the plant's growth environment and did not show seasonal variation. These results support the general hypothesis that many key parameters required by Earth system models are adaptive and predictable from plants' growth environments.

Variation in bulk-leaf 13 C discrimination, leaf traits and water-use efficiency-trait relationships along a continental-scale climate gradient in Australia.

Large spatial and temporal gradients in rainfall and temperature occur across Australia. This heterogeneity drives ecological differentiation in vegetation structure and ecophysiology. We examined multiple leaf-scale traits, including foliar 13 C isotope discrimination (Δ13 C), rates of photosynthesis and foliar N concentration and their relationships with multiple climate variables. Fifty-five species across 27 families were examined across eight sites spanning contrasting biomes. Key questions addressed include: (i) Does Δ13 C and intrinsic water-use efficiency (WUEi ) vary with climate at a continental scale? (ii) What are the seasonal and spatial patterns in Δ13 C/WUEi across biomes and species? (iii) To what extent does Δ13 C reflect variation in leaf structural, functional and nutrient traits across climate gradients? and (iv) Does the relative importance of assimilation and stomatal conductance in driving variation in Δ13 C differ across seasons? We found that MAP, temperature seasonality, isothermality and annual temperature range exerted independent effects on foliar Δ13 C/WUEi . Temperature-related variables exerted larger effects than rainfall-related variables. The relative importance of photosynthesis and stomatal conductance (gs ) in determining Δ13 C differed across seasons: Δ13 C was more strongly regulated by gs during the dry-season and by photosynthetic capacity during the wet-season. Δ13 C was most strongly correlated, inversely, with leaf mass area ratio among all leaf attributes considered. Leaf Nmass was significantly and positively correlated with MAP during dry- and wet-seasons and with moisture index (MI) during the wet-season but was not correlated with Δ13 C. Leaf Pmass showed significant positive relationship with MAP and Δ13 C only during the dry-season. For all leaf nutrient-related traits, the relationships obtained for Δ13 C with MAP or MI indicated that Δ13 C at the species level reliably reflects the water status at the site level. Temperature and water availability, not foliar nutrient content, are the principal factors influencing Δ13 C across Australia.

Embolism recovery strategies and nocturnal water loss across species influenced by biogeographic origin.

Drought-induced tree mortality is expected to increase in future climates with the potential for significant consequences to global carbon, water, and energy cycles. Xylem embolism can accumulate to lethal levels during drought, but species that can refill embolized xylem and recover hydraulic function may be able to avoid mortality. Yet the potential controls of embolism recovery, including cross-biome patterns and plant traits such as nonstructural carbohydrates (NSCs), hydraulic traits, and nocturnal stomatal conductance, are unknown. We exposed eight plant species, originating from mesic (tropical and temperate) and semi-arid environments, to drought under ambient and elevated CO2 levels, and assessed recovery from embolism following rewatering. We found a positive association between xylem recovery and NSCs, and, surprisingly, a positive relationship between xylem recovery and nocturnal stomatal conductance. Arid-zone species exhibited greater embolism recovery than mesic zone species. Our results indicate that nighttime stomatal conductance often assumed to be a wasteful use of water, may in fact be a key part of plant drought responses, and contribute to drought survival. Findings suggested distinct biome-specific responses that partially depended on species climate-of-origin precipitation or aridity index, which allowed some species to recover from xylem embolism. These findings provide improved understanding required to predict the response of diverse plant communities to drought. Our results provide a framework for predicting future vegetation shifts in response to climate change.

Transpiration of Eucalyptus woodlands across a natural gradient of depth-to-groundwater.

Water resources and their management present social, economic and environmental challenges, with demand for human consumptive, industrial and environmental uses increasing globally. However, environmental water requirements, that is, the allocation of water to the maintenance of ecosystem health, are often neglected or poorly quantified. Further, transpiration by trees is commonly a major determinant of the hydrological balance of woodlands but recognition of the role of groundwater in hydrological balances of woodlands remains inadequate, particularly in mesic climates. In this study, we measured rates of tree water-use and sapwood 13C isotopic ratio in a mesic, temperate Eucalypt woodland along a naturally occurring gradient of depth-to-groundwater (DGW), to examine daily, seasonal and annual patterns of transpiration. We found that: (i) the maximum rate of stand transpiration was observed at the second shallowest site (4.3 m) rather than the shallowest (2.4 m); (ii) as DGW increased from 4.3 to 37.5 m, stand transpiration declined; (iii) the smallest rate of stand transpiration was observed at the deepest (37.5 m) site; (iv) intrinsic water-use efficiency was smallest at the two intermediate DGW sites as reflected in the Δ13C of the most recently formed sapwood and largest at the deepest and shallowest DGW sites, reflecting the imposition of flooding at the shallowest site and the inaccessibility of groundwater at the deepest site; and (v) there was no evidence of convergence in rates of water-use for co-occurring species at any site. We conclude that even in mesic environments groundwater can be utilized by trees. We further conclude that these forests are facultatively groundwater-dependent when groundwater depth is <9 m and suggest that during drier-than-average years the contribution of groundwater to stand transpiration is likely to increase significantly at the three shallowest DGW sites.

Soil moisture controls on phenology and productivity in a semi-arid critical zone.

The Earth's Critical Zone, where physical, chemical and biological systems interact, extends from the top of the canopy to the underlying bedrock. In this study, we investigated soil moisture controls on phenology and productivity of an Acacia woodland in semi-arid central Australia. Situated on an extensive sand plain with negligible runoff and drainage, the carry-over of soil moisture content (θ) in the rhizosphere enabled the delay of phenology and productivity across seasons, until conditions were favourable for transpiration of that water to prevent overheating in the canopy. Storage of soil moisture near the surface (in the top few metres) was promoted by a siliceous hardpan. Pulsed recharge of θ above the hardpan was rapid and depended upon precipitation amount: 150mm storm(-1) resulted in saturation of θ above the hardpan (i.e., formation of a temporary, discontinuous perched aquifer above the hardpan in unconsolidated soil) and immediate carbon uptake by the vegetation. During dry and inter-storm periods, we inferred the presence of hydraulic lift from soil storage above the hardpan to the surface due to (i) regular daily drawdown of θ in the reservoir that accumulates above the hardpan in the absence of drainage and evapotranspiration; (ii) the dimorphic root distribution wherein most roots were found in dry soil near the surface, but with significant root just above the hardpan; and (iii) synchronisation of phenology amongst trees and grasses in the dry season. We propose that hydraulic redistribution provides a small amount of moisture that maintains functioning of the shallow roots during long periods when the surface soil layer was dry, thereby enabling Mulga to maintain physiological activity without diminishing phenological and physiological responses to precipitation when conditions were favourable to promote canopy cooling.

Functional Traits and Water Transport Strategies in Lowland Tropical Rainforest Trees.

Understanding how tropical rainforest trees may respond to the precipitation extremes predicted in future climate change scenarios is paramount for their conservation and management. Tree species clearly differ in drought susceptibility, suggesting that variable water transport strategies exist. Using a multi-disciplinary approach, we examined the hydraulic variability in trees in a lowland tropical rainforest in north-eastern Australia. We studied eight tree species representing broad plant functional groups (one palm and seven eudicot mature-phase, and early-successional trees). We characterised the species' hydraulic system through maximum rates of volumetric sap flow and velocities using the heat ratio method, and measured rates of tree growth and several stem, vessel, and leaf traits. Sap flow measures exhibited limited variability across species, although early-successional species and palms had high mean sap velocities relative to most mature-phase species. Stem, vessel, and leaf traits were poor predictors of sap flow measures. However, these traits exhibited different associations in multivariate analysis, revealing gradients in some traits across species and alternative hydraulic strategies in others. Trait differences across and within tree functional groups reflect variation in water transport and drought resistance strategies. These varying strategies will help in our understanding of changing species distributions under predicted drought scenarios.