Precipitation mediates sap flux sensitivity to evaporative demand in the neotropics.

Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

Tree mortality from an exceptional drought spanning mesic to semiarid ecoregions.

Significant areas of the southern USA periodically experience intense drought that can lead to episodic tree mortality events. Because drought tolerance varies among species and size of trees, such events can alter the structure and function of terrestrial ecosystem in ways that are difficult to detect with local data sets or solely with remote-sensing platforms. We investigated a widespread tree mortality event that resulted from the worst 1-year drought on record for the state of Texas, USA. The drought affected ecoregions spanning mesic to semiarid climate zones and provided a unique opportunity to test hypotheses related to how trees of varying genus and size were affected. The study was based on an extensive set of 599 distributed plots, each 0.16 ha, surveyed in the summer following the drought. In each plot, dead trees larger than 12.7 cm in diameter were counted, sized, and identified to the genus level. Estimates of total mortality were obtained for each of 10 regions using a combination of design-based estimators and calibrated remote sensing using MODIS 1-yr change in normalized difference vegetation index products developed by the U.S. Forest Service. As compared with most of the publicized extreme die-off events, this study documents relatively low rates of mortality occurring over a very large area. However, statewide, regional tree mortality was massive, with an estimated 6.2% of the live trees perishing, nearly nine times greater than normal annual mortality. Dead tree diameters averaged larger than the live trees for most ecoregions, and this trend was most pronounced in the wetter climate zones, suggesting a potential re-ordering of species dominance and downward trend in tree size that was specific to climatic regions. The net effect on carbon storage was estimated to be a redistribution of 24-30 Tg C from the live tree to dead tree carbon pool. The dead tree survey documented drought mortality in more than 29 genera across all regions, and surprisingly, drought resistant and sensitive species fared similarly in some regions. Both angiosperms and gymnosperms were affected. These results highlight that drought-driven mortality alters forest structure differently across climatic regions and genera.

Nocturnal transpiration in riparian Tamarix thickets authenticated by sap flux, eddy covariance and leaf gas exchange measurements.

Tamarix chinensis Lour., which is common throughout the southwestern USA, is a phreatophytic riparian tree capable of high water use. We investigated temporal congruence between daily total evapotranspiration (E) estimated from stem sap flux (J(s)) measurements (E(sf)) and eddy covariance (E(cv)), both seasonally and immediately following rain events, and used measurements of leaf-level gas exchange, stem water content and diurnal changes in leaf water potential to track drivers of transpiration. In one study, conducted near the end of the growing season in a pure T. chinensis stand adjacent to the Rio Grande River in central New Mexico, nighttime E(sf) as a proportion of daily E(sf) increased with water availability to a peak of 36.6%. High nighttime E(sf) was associated with underestimates of nighttime E(cv). A second study, conducted in west Texas, beside the Pecos River, investigated the relationships between nighttime J(s) and stem tissue rehydration, on the one hand, and nighttime E, on the other hand. Leaf gas exchange measurements and stomatal impressions suggested that nighttime J(s) was primarily attributed to concurrent transpiration, although there were small overnight changes in stem water content. Both vapor pressure deficit and soil water availability were positively related to nighttime J(s), especially following rainfall events. Thus, both studies indicate that T. chinensis can transpire large amounts at night, a fact that must be considered when attempting to quantify E either by eddy covariance or sap flux methods.

A mesic maximum in biological water use demarcates biome sensitivity to aridity shifts.

Biome function is largely governed by how efficiently available resources can be used and yet for water, the ratio of direct biological resource use (transpiration, E T) to total supply (annual precipitation, P) at ecosystem scales remains poorly characterized. Here, we synthesize field, remote sensing and ecohydrological modelling estimates to show that the biological water use fraction (E T/P) reaches a maximum under mesic conditions; that is, when evaporative demand (potential evapotranspiration, E P) slightly exceeds supplied precipitation. We estimate that this mesic maximum in E T/P occurs at an aridity index (defined as E P/P) between 1.3 and 1.9. The observed global average aridity of 1.8 falls within this range, suggesting that the biosphere is, on average, configured to transpire the largest possible fraction of global precipitation for the current climate. A unimodal E T/P distribution indicates that both dry regions subjected to increasing aridity and humid regions subjected to decreasing aridity will suffer declines in the fraction of precipitation that plants transpire for growth and metabolism. Given the uncertainties in the prediction of future biogeography, this framework provides a clear and concise determination of ecosystems' sensitivity to climatic shifts, as well as expected patterns in the amount of precipitation that ecosystems can effectively use.

Leaf surface traits and water storage retention affect photosynthetic responses to leaf surface wetness among wet tropical forest and semiarid savanna plants.

While it is reasonable to predict that photosynthetic rates are inhibited while leaves are wet, leaf gas exchange measurements during wet conditions are challenging to obtain due to equipment limitations and the complexity of canopy-atmosphere interactions in forested environments. Thus, the objective of this study was to evaluate responses of seven tropical and three semiarid savanna plant species to simulated leaf wetness and test the hypotheses that (i) leaf wetness reduces photosynthetic rates (Anet), (ii) leaf traits explain different responses among species and (iii) leaves from wet environments are better adapted for wet leaf conditions than those from drier environments. The two sites were a tropical rainforest in northern Costa Rica with ~4200 mm annual rainfall and a savanna in central Texas with ~1100 mm. Gas exchange measurements were collected under dry and wet conditions on five sun-exposed leaf replicates from each species. Additional measurements included leaf wetness duration and stomatal density. We found that Anet responses varied greatly among species, but all plants maintained a baseline of activity under wet leaf conditions, suggesting that abaxial leaf Anet was a significant percentage of total leaf Anet for amphistomatous species. Among tropical species, Anet responses immediately after wetting ranged from -31% (Senna alata (L.) Roxb.) to +21% (Zamia skinneri Warsz. Ex. A. Dietr.), while all savanna species declined (up to -48%). After 10 min of drying, most species recovered Anet towards the observed status prior to wetting or surpassed it, with the exception of Quercus stellata Wangenh., a savanna species, which remained 13% below Anet dry. The combination of leaf wetness duration and leaf traits, such as stomatal density, trichomes or wax, most likely influenced Anet responses positively or negatively. There was also overlap between leaf traits and Anet responses of savanna and tropical plants. It is possible that these species converge on a relatively conservative response to wetness, each for divergent purposes (cooling, avoiding stomatal occlusion, or by several unique means of rapid drying). A better understanding of leaf wetness inhibiting photosynthesis is vital for accurate modeling of growth in forested environments; however, species adapted for wet environments may possess compensatory traits that mitigate these effects.

Epiphyte host preferences and host traits: mechanisms for species-specific interactions.

We investigated species-specific relationships among two species of vascular epiphytes and ten host tree species in a coastal plain forest in the southeastern United States. The epiphytes Tillandsia usneoides and Polypodium polypodioides were highly associated with particular host species in the field, but host traits that favored colonization were inadequate to fully explain the epiphyte-host associations for either epiphyte. Field transplant experiments that bypassed epiphyte colonization demonstrated that the growth of epiphytes was significantly higher on host tree species that naturally bore high epiphyte loads than on host species with few or no epiphytes. These species-specific relationships were highly correlated with the water-holding capacity of the host tree's bark. Positive and negative effects of throughfall, light attenuation by the canopy, and bark stability did not explain the overall patterns of host specificity, but did correlate with some epiphyte-host species relationships. The relative importance of particular host traits differed between the "atmospheric epiphyte" Tillandsia, and the fern Polypodium, which roots in the bark of its hosts. Species-specific interactions among plants, such as those described here, suggest that communities are more than individualistic assemblages of co-occurring species.

Structural and compositional controls on transpiration in 40- and 450-year-old riparian forests in western Oregon, USA.

Large areas of forests in the Pacific Northwest are being transformed to younger forests, yet little is known about the impact this may have on hydrological cycles. Previous work suggests that old trees use less water per unit leaf area or sapwood area than young mature trees of the same species in similar environments. Do old forests, therefore, use less water than young mature forests in similar environments, or are there other structural or compositional components in the forests that compensate for tree-level differences? We investigated the impacts of tree age, species composition and sapwood basal area on stand-level transpiration in adjacent watersheds at the H.J. Andrews Forest in the western Cascades of Oregon, one containing a young, mature (about 40 years since disturbance) conifer forest and the other an old growth (about 450 years since disturbance) forest. Sap flow measurements were used to evaluate the degree to which differences in age and species composition affect water use. Stand sapwood basal area was evaluated based on a vegetation survey for species, basal area and sapwood basal area in the riparian area of two watersheds. A simple scaling exercise derived from estimated differences in water use as a result of differences in age, species composition and stand sapwood area was used to estimate transpiration from late June through October within the entire riparian area of these watersheds. Transpiration was higher in the young stand because of greater sap flux density (sap flow per unit sapwood area) by age class and species, and greater total stand sapwood area. During the measurement period, mean daily sap flux density was 2.30 times higher in young compared with old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Sap flux density was 1.41 times higher in young red alder (Alnus rubra Bong.) compared with young P. menziesii trees, and was 1.45 times higher in old P. menziesii compared with old western hemlock (Tsuga heterophylla (Raf.) Sarg.) trees. Overall, sapwood basal area was 21% higher in the young stand than in the old stand. In the old forest, T. heterophylla is an important co-dominant, accounting for 58% of total sapwood basal area, whereas P. menziesii is the only dominant conifer in the young stand. Angiosperms accounted for 36% of total sapwood basal area in the young stand, but only 7% in the old stand. For all factors combined, we estimated 3.27 times more water use by vegetation in the riparian area of the young stand over the measurement period. Tree age had the greatest effect on stand differences in water use, followed by differences in sapwood basal area, and finally species composition. The large differences in transpiration provide further evidence that forest management alters site water balance via elevated transpiration in vigorous young stands.